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THE COCCIDAE
TABLES FOR THE IDENTIFICATION OF THE SUBFAMILIES AND SOME OF
THE MORE IMPORTANT GENERA AND SPECIES TOGETHER WITH
DISCUSSIONS OF THEIR ANATOMY AND LIFE HISTORY
BY
ALEX. D. MACGILLIVRAY
I will speak what I helieve to-day,
even if it contradicts everything that
I said yesterday. Gladstone.
SCARAB COMPANY
URBANA, ILLINOIS
1921
Copyright, 1921
BY
SCARAB COMPANY
All Rights Reserved
Including Those of Translation
ISSUED
January 10, 1921
printed by
flanigan-pearson company
champaign, ILLINOIS
S3u
en
TO
MY FORMER STUDENTS OF THE COCCIDAE WHO THROUGH
THEIR INSISTENCE ARE RESPONSIBLE FOR THE
COMPLETION AND PUBLICATION OF
THIS VOLUME
PREFACE
The following materials were originally collected for tlie use
of students in the indentification of coccids. They were based
upon printed descriptions and original studies of specimens. There
were, when the tables were first prepared, no extensive or limited
analyses that could be used for this purpose. The first plan was
to arrange analyses of all the genera and species of all the sub-
families, but it was soon found that this was impractical. The
first draft was prepared about fifteen years ago. This was later
greatly extended, modified, revised, and mimeographed for issu-
ance to students.
The mimeographed edition contained the following preface:
*'The tables given herewith are compilations to facilitate the deter-
mination of species. In making the compilations, numerous studies
were made on the phylogeny of the different subfamilies, genera,
and species and the results of the studies thus made are embodied
herein. Although compilations from descriptions in the main, yet
they contain names not used elsewhere (all those marked MacG.)
and hence unpublished. The preparation of mimeographed papers
does not constitute publication according to the rules of systematic
zoologists. Hence these new names and this manuscript has no
standing in entomological literature. In justice to the writer, there-
fore, the names and tables given herewith should be considered the
personal property of the writer and are not to be referred to by
any person using the tables in any paper that may be published
until after the names and facts here given have been printed in
such a way by the writer as to be recognized as published by
zoologists." This statement is included in order to fix the status
of the mimeographed copies in case any of them should find their
way into libraries.
Besides the preparation of the tables there was accumulated
from various sources in addition to my own studies upon speci-
mens, many facts bearing upon the preparation, anatomy, meta-
morphosis, and habits of coccids. These facts were presented to the
students in the form of a course of lectures. Most of the students
who have taken this coccid course have been provided with a set of
the mimeographed tables. I have had many requests from former
students and others to purchase the tables or asking for permis-
V
VI PREFACE
sion to be allowed to copy them and, when these requests have been
denied have urged that the material might be printed and made
available for the use of all. For this reason the accumulated
materials have been again revised and extended and are offered
on the following pages.
The number of separate publications or papers dealing with
the family Coccidae is legion. Mrs. M. E. Fernald prepared a
catalogue which gives references to the literature dealing with all
the species. This catalogue is of inestimable value to every
student of the Coccidae. The numerous titles to papers have not
been accumulated thus far by any one and I have given a consider-
able list of the books and papers dealing with the family. No
attempt has been made to make it complete.
The fact that no figures are included in this volume may
appear to many as an error. The tables were prepared primarily
for the use of students. Those who have had any experience in
teaching know that most students will not undertake anything
they are not forced to do. The omission of figures makes it neces-
sary for them to study their specimens rather than figures.
A careful examination of the figures and descriptions of the
species of Diaspidinae has made it possible to refer a considerable
number of the species to the generic groups recognized in this
volume. A number of species, however, either from the meagerness.
of their descriptions or the imperfections of their figures, have
been included under the generic group where they were originally
described. Further investigation will show that some of these
species are misplaced.
The materials accumulated are from such varied sources, that it
is impossible to name all the authors. I take this opportunity to
acknowledge my indebtedness to all students of the Coccidae for
the material that I have culled from their writings. The following
are deserving of special mention : Berlese, Cockerell, Comstock,
Green, Hempel, Kuwana, Leonardi, Marlatt, Maskell, Newstead,
and Signoret. I am also greatly indebted to all my former students
of the Coccidae for many suggestions and criticisms of the form and
arrangement of the various tables and of the characterizations used
therein.
A. D. M.
University of Illinois,
UrBx\na, Illinois,
July 15, 1920.
CONTENTS
Page
CHAPTER I.
Introduction _— _ 1
CHAPTER II
PEEPABATION 11
CHAPTER III
ExTEENAii Anatomy 24
CHAPTER IV.
Classification 40
CHAPTER V.
Subfamily Monophlebinae 62
CHAPTER VI.
Subfamily Kuwanhnae 76
CHAPTER VII.
Subfamily Xylococcinae 80
CHAPTER VIII.
Subfamily Mabgarodinae 88
CHAPTER IX.
Subfamily Callapappinae 94
CHAPTER X.
Subfamily Coccinae 99
CHAPTER XI.
Subfamily Ortheziinae 105
CHAPTER XII.
Subfamily Phenacoleachiinae 115
CHAPTER XIII.
Subfamily Ebiococcinae 118
CHAPTER XIV.
Subfamily Tachardiinae 148
CHAPTER XV.
Subfamily Lecaniinae 155
CHAPTER XVI.
Subfamily Asterolecaniinae 183
Page
CHAPTER XVII.
Subfamily Kebmesiinae 191
CHAPTER XVIII.
Subfamily Apiomorphinae 199
CHAPTER XIX.
Subfamily Cylindrococcinae 205
CHAPTER XX.
Subfamily Conchaspinae 212
CHAPTER XXI.
Subfamily Diaspidinae 217
CHAPTER XXII.
Tribe Pablatoriini 245
CHAPTER XXIII.
Tribe Leucaspini 258
CHAPTER XXIV.
Tribe Lepidosaphini 269
CHAPTER XXV.
Tribe Diaspidini 297
CHAPTER XXVI.
Tribe Fiobiniini 367
CHAPTER XXVII.
Tribe Aspidiotini 380
Bibliography 486
Index 475
THE COCCIDAE
CHAPTER I
INTRODUCTION
The subdivision of the body into a series of somewhat similar
rings or areas and the modification of the external surface of the
rings into a hardened skeleton or cuticle are important character-
istics. They denote that any animal possessing them is an
Arthropod. The rings, known as segments, are separated by dis-
tinct constrictions or infoldings. A portion of each infolded
portion, known as a conjunctiva or coria, is generally different in
color and more flexible. The presence of a pair of segmented
appendages on each or a few of the segments is a further indi-
cation of an arthropodan relationship. The fact that the segments
are assembled into three groups or regions, a cephalic region or
head which is apparently unsegmented but in reality consists of
six segments, an intermediate region or thorax consisting of three
segments, and a caudal region or abdomen consisting typically of
eleven segments, is an indication of an affinity with the hexapods
or insects. This relationship is usually emphasized by the presence
of a pair of legs on each segment of the thorax, the absence of legs
on the abdomen, and the modification of the appendages of the
segments of the head into mouth-parts. The evidences of relation-
ship of coccids to arthropods and insects are in part wanting in
many coccids and are all wanting in some.
The insects of the order Hemiptera, to which the family Coc-
cidae belongs, possess typically the following characteristics. The
constriction between the head and prothorax, the procoria, is dis-
tinct. The coria between the prothorax and mesothorax, the meso-
coria, is also distinct, while the one between the mesothorax and
metathorax, the metacoria, as well as the coria between the met-
athorax and the first abdominal segment, the unacoria, are usually
more or less obscure. The head bears a pair of compound eyes,
a pair of antennae of varying size and shape, and three pairs of
mouth-parts. These latter are of the sucking type. Two of the
^ THE COCCIDAE
pairs of mouth-parts, the mandibles and the maxillae, are long,
slender, bristle-like appendages that are similar in appearance.
The third pair, the labium, is in the form of a long segmented tube
which is fitted for sucking and piercing and is known as the ros-
trum. This tube is normally folded against the ventral aspect of
the head and thorax. It contains a longitudinal furrow which is
open on the dorsal aspect. The sides of this furrow are adjacent,
forming a lumen in which the bristle-like mandibles and maxillae
are held and supported. Each segment of the thorax bears a pair
of legs, which consist of the usual number of sclerites or segments.
The mesothorax and metathorax usually bear also a pair of wings.
The thorax generally bears two and the abdomen a varying num-
ber, never more than eight and usually about six, pairs of
spiracles. Certain coccids lack all these features indicating rela-
tionship with the Hemiptera except the presence of two pairs of
thoracic spiracles and of a thin external body skeleton or cuticle.
The great diversity and peculiarity in the form of many
coccids and the similarity to them in general external appearance
of certain species of some of the related families of Hemiptera has
led even some specialists into the error from time to time of
describing insects other than coccids as such. All orders of insects
can be divided into two large groups according to the number of
tarsal segments, in one group would fall those with five segments,
assumed to be the primitive condition, and in the other those with
less than five segments. The Hemiptera and the closely related
orders of Exometabola with sucking mouth-parts belong to the
group with less than five tarsal segments. This latter group can
be divided into two series upon the size of the pronotum and the
condition of the wings. In one of the series the mesothoracic wings
are modified into veinless wing-covers or elytra and the pronotum
is a large subquadrangular area loosely articulated to the
mesonotum, while in the other group the wings, although sometimes
thickened, are never modified into elytra and the pronotum is
reduced in size, collar-like and immovably united with the
mesothorax. The Hemiptera and their allies, which belong to the
latter series, are easily separated from all other insects with tarsi
of less than five segments, because their mouth-parts, in immature
individuals as well as adults, are fitted for sucking. It is very
unusual to have immature insects with sucking mouth-parts.
The form and structure of the sucking mouth-parts of the
orders with less than five segments in their tarsi are sufficiently
distinctive and typical to differentiate them from each other. The
INTRODUCTION 3
characters recorded in the following table will serve not only to
separate these orders from each other, but will also serve to
separate them from all the other orders of insects: —
a. Legs normal in form, fitted for walking or swimming, never scan-
sorial or with tibia and tarsus and claw of each leg combined to
form an opposed thumb and finger for clinging to hairs; labium
modified into a distinct alv/ays exposed sucking tube, if wanting,
thorax with a single pair of wings or the tarsi with a single claw
or legs wanting.
b. Tarsi with the distal segment bladder-like or hoof -like in form;
wings long, narrow, subequal in size, margins parallel and fringed
with setae as long as the width of wing; labium a conical tube
enclosing one mandible and two maxillae; maxillae and labium
provided with palpi; clypeus asymmetrical in the adult; legs and
mouth-parts always present Physopoda.
bb. Tarsi with the distal segment never bladder-like or hoof-like in
form; wings triangular or subtriangular in outline, one of the
wings of each side usually larger than the other, never with
margins parallel and fringed with long setae; labium a segmented
tube enclosing bristle-like mandibles and maxillae; maxillae and
labium never provided with palpi; clypeus always symmetrical;
legs and mouth-parts sometimes wanting Hemipteba.
aa. Legs abnormal in form, never fitted for walking or swimming,
always scansorial or with tibia and tarsus and claw of each leg
combined to form an opposed thumb and finger for clinging to
hairs ; labium modified into a sucking tube that is usually retracted
and concealed; wings always wanting and legs always present _
Anopluba.
The Anoplura or lice are generally included as a suborder
of Hemiptera under the name of Parasitica. The evidence that
they are degraded Hemiptera is not very conclusive. The form of
their mouth-parts, a closed sucking tube without mandibles or
maxillae, is very different from that of the species of this order.
The order Hemiptera can be characterized more definitely and the
Anoplura more easily differentiated from the Hemiptera if it is
considered as a distinct order.
The order Hemiptera is usually divided into three suborders,
Heteroptera, Homoptera, and Parasitica. The last of these has
been recognized as a distinct order as already indicated. The
texture of the wings, the shape and position of the head, the point
of attachment and structure of the labium or rostrum, the form of
the tarsi, and the form, size, and presence of the antennae are
the characteristics generally used to distinguish the suborders
Heteroptera and Homoptera. The difference in the form of the
jk THE COCCIDAE
antennae and in the general appearance of the families of this
suborder, as generally recognized, has led to its subdivision into
two groups or suborders. The structure of the wings, the form
and origin of the mouth-parts, and the presence of many other
minor differences emphasize the desirability of this subdivision: —
SUBORDERS OF HEMIPTERA
a. Head with rostrum attached to its cephalic end, distant from the
prothoracic legs, ventro-meson of the head distinctly longer than
the distance on the venter between compound eyes; mesothoracic
wings, when present, with proximal portion thickened and distal
portion membranous. Hetekoptera.
aa. Head with rostrum attached to its ventral aspect near the articula-
tion of prothoracic legs, ventro-meson distinctly shorter than distance
on the venter between compound eyes; mesothoracic wings, when
present, always of same texture throughout, never thickened on
proximal and membranous on distal portion,
b. Antennae minute and inconspicuous, setiform or awl-shaped; tarsi
with three segments; prothorax large and conspicuous
HOMOPTERA.
bb. Antennae long and filiform, rudimentary, or wanting, never seti-
form or awl-shaped; tarsi with one or two segments; prothorax
small and frequently inconspicuous. Gularostria.
The suborder Heteroptera as here defined includes the super-
families and families usually assigned to it. The suborder
Homoptera includes the superfamilies Cicadoidea, Jassoidea, Ful-
goroidea, and Membracoidea, while to the suborder Gularostria is
assigned the Psyllidae, Aphididae, Aleyrodidae, and Coccidae.
These groups can be separated by means of the following table : —
FAMILIES OF GULAROSTRIA
a. Tarsi of adult rarely with less than two segments and usually with
two claws, legs never wanting; wings, when present, four in number;
eyes in adult always compound, if present in nymph, compound;
rostrum always present in nymphs and adults, usually greatly elon-
gated and projecting beyond the mesocoxae; adult insects never
scale-like, grub-like, or gall-like in form and never immovably fixed
to the food plant; nymphs sometimes scale-like, legless, and immov-
able, if so, abdomen always provided with a vasiform orifice,
b. Wings, when present, membranous; the metathoracic wings al-
ways much smaller than the mesothoracic; body of adult and its
wings sometimes bearing filaments, threads, or tufts of wax,
never densely covered with a fine, white, and powdery wax;
nymphs and adults frequently associated, subsimilar in form, both
provided with antennae and legs, both capable of locomotion,
nymphs never provided with a vasiform orifice.
INTRODUCTION O
c. Antennae with eight to ten segments; rostrum with three
segments, reaching to mesocoxae; metacoxae near together,
about length of their coxae apart; adults always provided with
wings, veins arising as branches from both sides of a short
median proximal stem; nymphs with body usually strongly
depressed, wing pads horizontal and usually ending beyond
the body. Psyllidae.
cc. Antennae with three to six segments; rostrum with four or
five segments, reaching to and frequently beyond the mesocoxae ;
metacoxae distant, four or five times as far apart as the length
of their coxae; adults winged or wingless, veins arising as
branches from the caudal margin only of a large cephalic vein;
nymphs with body cylindrical, rarely depressed, wing pads not
horizontal and usually held against the body Aphididae.
bb. Wings membranous and rarely absent in adult; metathoracic
wings subequal in size to mesothoracic; body of adult and wings
always densely covered with a fine white, powdery wax; nymphs
and adults sometimes closely associated, always different in form,
adults insect-like, nymphs always scale-like or grub-like in form
and never covered with a white waxy powder, frequently with
marginal plates of wax, always without antennae and legs, and
abdomen always provided with a vasiform orifice Aleyrodidae.
aa. Tarsi of the adult rarely with more than one segment and alv/ays
with a single claw, legs frequently wanting; wings, when present,
two in number; eyes in adult and nymphal female never compound
and frequently wanting, wanting in nymphal male and rarely com-
pound in adult male; rostrum very shorit, rarely reaching mesocoxae
or as far as metathoracic spiracles if mesocoxae are wanting in fe-
male, wanting in older nymphal stages and always in adult male and
rarely wanting in adult female; adult male usually winged and
insect-like, adult female scale-like, grub-like, or gall-like in form,
frequently inbedded in mass of wax of its own excretion, frequently
fixed to the food plant and immovable; legs and antennae frequently
greatly reduced in size, rudimentary, or wanting in nymphal and
adult females; abdomen never provided with vasiform orifice
COCCIDAE.
The members of the family Coccidae live upon the stems,
leaves and roots of plants. While a great majority of the species
are fixed to the host-plant for at least a part of their life, some
are always active. The fixity or quiesence of the different species
is confined for the most part to the female sex and is always
associated with the production of eggs and young. There is the
greatest diversity in the external form of coccids. This is due
more to the form of the waxy excretion with which the body is
covered than to the general form of the body itself. The amount
of wax and the form it assumes are also associated with repro-
duction. There is frequently considerable variation in the external
6 THE COCCIDAE
appearance of different individuals of the same species due to
their being crowded into crevices and depressions upon the host-
plant and also to the close crowding together of the different
individuals. While there is considerable variation in the form and
structure of the body, these differences are always so minute that
they can not be seen without the aid of a miscroscope and many
of them are difficult to interpret even by the use of a microscope
of considerable magnification. In the classification of coccids the
early systematists used the superficial characters, those based on
the shape and consistency of the wax. Comstock demonstrated
not only the validity of the structural characters of the pygidium,
but used them in differentiating species of Diaspidinae so that it
is now possible with a considerable degree of accuracy to identify
species of this subfamily. The genera and species of the other
subfamilies are still based to a large extent upon superficial
characters.
The greatest development in number of genera and species is
found in tropical and subtropical regions, but many genera and
species extend into and are peculiar to temperate regions. The
number of peculiar or bizarre species that have been described
from Australia and the islands connecting this continent with the
Asiatic mainland is large. While certain species of coccids are
found only upon certain plants and frequently only upon certain
parts of the plant, other species whether limited to a particular
part of the plant or not may occur on all the species of plants of
a genus, of a iamily, or all the families of an order. Some species
are found apparently indiscriminately upon almost any plant,
regardless of its taxanomic association and upon any part of the
plant but the roots. Only a few coccids feed upon the roots of
plants. They are peculiar to this situation and belong as a rule
to particular coccid genera.
The individual insects are usually small and inconspicuous
and easily overlooked. While this is true of most coccids, some
of the species of the generalized subfamilies attain a size of one
inch or more. Many of the large or medium sized species are
conspicuous in the adult stage because of the mass of white wax
which surrounds their body or its attachment to it. This is partic-
ularly true of colonies of Pulvinaria which are usually overlooked
until they begin to form their ovisacs. The amount of wax
excreted by the female before and during the deposition of her
eggs varies with the subfamily and to a certain extent with the
species. Some females excrete a long ribbon of doughy wax five
INTRODUCTION 7
or six inches long as in Takahashia. In certain of the species
forming an abundance of wax, it is a thick homogenous mass and
in the case of Tachardia, the shellac forming insects, the wax_is
an important article of commerce. A long time, several weeks
to three months, is sometimes required in the production of the
ovisac, as in Icerya. The small species may become conspicuous
from the large number of individuals present in a given area,
this is also increased sometimes by the color, as in the case of the
male scales of Chionaspis and Aulacaspis. The number may be
so large that the surface is completely covered or encrusted and
hence the name of scale insects. Most of the species that occur in
considerable numbers, as the species of Aspidiotus, Lepidosaphes,
or Chionaspis, form only a minute leaf-like disk of wax and molted
skins. The females of Icerya and of other large species when they
are ready to form their large conspicuous ovisacs, seem to con-
gregate in the same place so that many adjacent ovisacs are
frequently found close together.
The coccids and aleyrodids of all the insects that undergo
a direct or incomplete metamorphosis show the farthest departure
from the generalized orthopteran type of metamorphosis. In the
case of the Coccidae there is more difference between the two sexes
throughout the greater part of their life than exists between many
species of insects of other families or orders.
When the recently emerged coccid is compared with the
diagnosis of a typical hemipterous insect and of a gularostrian,
it is seen that there is still retained most of the features peculiar
to these types. The most striking change is the modification of
the compound eyes into a pair of simple eyes or ocellanae, the
great reduction in the size of the constrictions between the head
and prothorax and the prothorax and mesothorax, the procoria
and the mesocoria, so that it is often difficult to determine the
limits of the head and of the thoracic segments. The antennae and
legs, while they are typical in form so far as segmentation and
number of parts are concerned, are different in appearance. They
are large and ungainly in appearance and are apparently out of
all proportion in size with the remainder of the body. This is not
true of other nymphal Hemiptera or of nymphal insects in general.
The two sexes, while so different throughout the greater part
of their life, are indistinguishable during the first nymphal stage.
The nymphs, when they emerge from the egg, remain perfectly
quiescent for a time, a few hours to two days, in the place where
the eggs were laid. These young nymphs are suddenly seized with
8 THE COCCIDAE
a desire to get out and wander. It acts upon them suddenly as
if they were seized with hysteria, they become not only active but
they literally swarm out over the branches of the host-plant until
frequently it is completely covered. The function of this migration
period is apparently for the dissemination of the species and to
find an unencumbered portion of the host-plant where the nymph
can attach itself and begin to draw food.
The generalized coccids are active throughout their entire
nymphal life and in some even throughout their entire life as the
females of the genus Orthezia. The great majority are quiescent
during the adult stage. It would not be strange, therefore, to find
that the length of the active period had been shortened and the
quiescent period lengthened from just the adult condition to
practically the entire life of the insects. The quiescent period in
many specialized coccids begins just as soon as the migrating
nymphs of the first nymphal period &x themselves to the host-
plant. In many species the appendages, including the legs and
antennae, and even the mandibles and maxillae in the male, but
not the labium or rostrum, are lost at the first molt, so that the
animals are capable of only slight movements.
The males are easily identified after their entrance into the
quiescent period which is after the first molt in the Eriococcinae,
because the mouth-parts are wanting. They can usually be identi-
fied a short time before the first molt in mounted specimens. The
developing mandibles and maxillae of the female which are func-
tional during the second stage can be identified as a coiled watch-
spring-like structure within the cuticle on each side of the rostrum.
These coiled structures, since the mandibles and maxillae are
wanting in the second and following stages of the male, are never
present in the first nymphal stage.
The nymphal males usually molt about four times and there
are four nymphal stages. The young nymphs form a cocoon or
scale from threads of wax at sometime before or near the beginning
of the quiescent period. This quiescence may begin early in the
second nymphal period or it may be delayed to as late as the
fourth nymphal period. The molted skins are usually pushed out
under the nymphal case or scale as they are cast. The nymphal
appendages and legs are lost at the molt when the male enters the
quiescent period. The appendages of the adult, including a single
pair of wings in most males, are developed from imaginal discs.
It has been maintained by some that the male coccid passes
through an indirect or complete metamorphosis. Their determi-
nation is based upon the presence of a quiescent period during the
INTRODUCTION
9
latter part of the nymphal life and by some to the fact that the
wings are developed from imaginal discs. The great variation in
the length of the quiescent period and the development of the
wings of all insects with an incomplete metamorphosis from
imaginal discs would show that the hypothesis that the male coccM
has a complete metamorphosis is without foundation. This is
further accentuated by the fact that the wings of the male coccid
are always developed externally, never internally, as in all insects
with an incomplete metamorphosis.
The adult male is identical in form with the hemipterous type
except that the bristle-like mandibles and maxillae are wanting
and the rostrum is represented by a slight conical projection. The
mouth-parts are often incorrectly stated to be wanting. The state-
ment that the ventral pair of ocellanae are developed from and
replace the mouth-parts is groundless. The transverse constrictions
between the segments of the body, the coriae, are of the usual
insect ean type and always more distinct than in the first nymphal
stage.
The females are generally stated to have an incomplete meta-
morphosis. This has probably arisen through the absence of wings
which are the most important external structures that can be used
in determining what type of metamorphosis exists. It is safe to
conclude, however, since the male has an incomplete metamorphosis
and that the members of the other families of this order have a
typical incomplete metamorphosis that the females of this family
have a similar type. The females of the different genera show a
greater diversity in the number of nymphal stages than the males.
There may be two, three, or four molts and a corresponding num-
ber of nymphal stages. It is an interesting fact that the generalized
female coccids have the same number of molts as most of the males
and that it approximates the condition found in most insects.
There is a direct correlation between the specialization of the
structures of the female and the reduction in the number of molts.
The most specialized species exist in only three conditions or
stages, two of which are nymphal. The first nymphal stage is an
active one for migration and location, the second nymphal stage
is for feeding and growth, and the third stage or adult stage is for
the production of eggs or young. The developing female differs
from the male in that she departs farther and farther after each
molt from the form and structure of hemipterous and other insects
while the males approximate them closer and closer in form. The
females become grub-like or bag-like while the males become
insect-like.
10 THE COC:iDAE
The transverse constrictions of the body of the female in each
succeeding nymphal stage after the first become more and more
obscure, until in the adult females of many species the coriae
are wanting or so indefinite that it is impossible to determine,
except in a general way, the caudal extent of the head. The eyes
are represented by a single ocellana on each side of the head or
are wanting. The body increases in size and becomes plumper so
that the antennae and legs do not appear so long and large and
out of proportion to the remainder of the body. The adult females
of many coccids are without legs and frequently, where they are
retained as in most Coccinae, they do not project beyond the
lateral margin of the body.
The female like the male may pass through a quiescent period
and, as in the case of the male, it varies greatly in length. The
legs and antennae which are wanting in most of the nymphal
stages of Margarodes are regained in the adult. The function of
the quiescent period is entirely different, not only for the making
over of the organs and tissues of the body, but for the production
and laying of eggs or for the giving birth to young. In a few
cases, as the Ortheziinae, the female never becomes quiescent. She
excretes a sack of wax which remains attached to her body. The
eggs are deposited in this sack, which is carried about until the
eggs hatch and the young nymphs emerge from the free and open
end. In the great majority of the species, generalized as well as
specialized, the female forms a mass or sack of wax or a scale which
is attached to the host-plant. This wax, which varies greatly in
consistency, is either interposed between the body of the female
and the host-plant or forms a covering over her body or completely
encases her body in a flocculent or dense mass. It is the color of
the wax, which is generally white, that makes the individuals or
colonies of coccids conspicuous and attracts attention to their
presence.
The members of the family Coccidae are of great economic
importance because of the injuries that they do to growing plants.
They subsist upon the sap which they obtain by means of the
tube formed by the close apposition of the bristle-like mandibles
and maxillae. Even if the great diversity and complexity of their
metamorphosis, their complicated anatomy, and the different ways
in which they produce substances useful to man were not con-
sidered of sufficient importance to warrent spending so much time
on the study of these insects, sufficient reason is found in the great
number of injurious species belonging to this family and the
enormous amount of damage which they do.
CHAPTER II
PREPARATION
The characters used for the separation of the various sub-
families, genera, and species of Coccidae are confined almost
entirely to the females. The reason for this is that the males are
rarely encountered unless special efforts are made to locate them
or to breed them. They are consequently not so well known and
have not been subjected to such extensive studies as the females
which, if found at all, usually occur in considerable numbers. It
is not strange, therefore, that the classification has been based on
the structures and excretions of the female. All individuals,
either from their size or from the presence of the excretions which
cover them, need to be treated with chemicals before their finer
structures can be examined.
The body in the great majority of the species is small. In
many it is microscopic in size. Whether the body is large or small,
it has been greatly modified, specialized by reduction, so that there
is an astonishing simplification of the external form. In certain
adult females the body is hardly more than a simple cuticular bag.
The external appendages and the structures associated with them
have been modified, greatly reduced in size, or are wanting. The
great number of species and the dearth of usable characters, because
of the simplification of their external form and structure, makes
it necessary to employ every available structure. The females
have their bodies covered with wax. The amount of wax varies
with the species, the volume excreted by some is much greater
than the volume of their bodies. The wax must be removed and
the body and its appendages clarified so that the form and struc-
ture of the various parts can be determined.
The difficulty is apparently further increased by the fact that
in most cases the specimens usually available for study are the
dried wrinkled bodies of females that have laid their eggs and
that have been stored in museums from a few to many years. The
bodies of such individuals must be removed so that all parts can
be examined. It is desirable that the body be returned as nearly
as possible to its original form, to that of the living individual.
11
12 THE COCCIDAE
This can be accomplished fortunately just as readily from speci-
mens that have been dried for a long period of years as from fresh
or living specimens.
The dried specimen is transformed by treating it with chemicals
which will not only dissolve the wax, but soften the cuticle, expand
the wrinkles until the body is of its original size and shape, and
clarify the dark and clouded portions. The making of a good
preparation of a coccid is not an easy matter. It does not consist
in dumping some specimens roughly and carelessly into a dish
with certain chemicals, and, then by juggling them through a series
of other dishes and chemicals, expect them to come out a finished
product. Those wRt) undertake the task with this attitude of mind
have a sad awakening before them and the fewer number of
bunglers of this type who undertake the preparation and study of
the classification of the Coccidae, the better off the subject will be.
The making of a good preparation is an operation that requires
time, unbounded patience, and skill in manipulation. It frequently
occurs even when all of these are applied to the very best of one^s
ability that a large proportion or all of the resulting mounted
specimens may be practically worthless for study. There is no
easy road to follow at such times. The only thing to do is to try
again and to keep on trying until preparations that will answer
every necessity of the case are secured.
Equipment. — The making of good microscopic preparations
presupposes that the preparator is provided with all the necessary
tools, chemicals, and glassware. A considerable equipment is
desirable but not essential in the preparation of eoccids. Such an
outfit is not in use continuously and it is frequently advantageous
to remove the bottles and glassware at such times from the table.
This is more easily accomplished if it is arranged in a drawer that
can be shoved into a locker or cabinet. The outfit named below
can be arranged compactly in a drawer eighteen inches long, twelve
inches wide, and three inches deep. If an inch board with holes
bored in it in which the various bottles and jars will fit, is placed
in the bottom of the drawer, there will be less shifting about and
the various vessels will always be in their proper places.
Such a drawer should be provided with the following
articles : —
1. Rubber stoppered bottle for ten per cent caustic potash
and holding 500 cc.
2. Glass stoppered bottle for 95 per cent alcohol and holding
500 cc.
PREPARATION 13
3. Glass stoppered bottle for distilled water and holding
500 cc.
4. Glass stoppered bottle holding 100 cc. for staining
solution.
5. Bottle with dropper for carbol-xylol and holding_250 cc.
6. Stender jar, 60 x 90 mm. for clean slides.
7. Stender jar, 60 x 35 mm. for clean cover-glasses.
8. Stender jar, 60 x 90 mm. for burnt matches.
9. Ointment jar for waste. A large covered jar, preferably-
one of the colored glass jars sold as ointment jars, about three and
one-half inches broad and about six inches high, is needed into
which can be drained waste liquids as used caustic potash, water,
and alcohol. A quart fruit jar can be used if nothing else is
available.
10. A jar of xylol balsam with a solid glass rod.
11. Box of safety matches.
12. Bunsen burner stand with shelf. The best type is that
where one of the legs is provided with a shelf which can be raised
or lowered and fastened in place with a set screw.
13. Alcohol lamp. Better results can be obtained by the use
of an alcohol lamp than by the use of a gas flame. The alcohol
lamp should be of good size so as not to need refilling too frequently.
Care should be used not to fill the lamp too full or when it gets
hot the alcohol may explode.
14. Casserole with cover and wooden handle. The best vessel
in which to boil coccids is a casserole about three and one-half
inches in diameter and holding about four ounces. Any sort of
chemical glassware, as beakers or test tubes, can be used, but
when considerable heat is applied to caustic potash, causing it to
boil vigorously, it is liable to pop out of any vessel with a restricted
mouth or neck and to carry the specimens with it. There is no
danger from popping-out if a wide-mouthed vessel such as a
casserole is used. It will be found that the wooden handle will
save burnt fingers and temper.
15. Syracuse Watch-glasses. The watch-glasses with flat
bottoms and grooved so that they can be stacked up are the most
convenient receptacles in which to perform many of the operations
necessary in the treatment of the specimens, as dehydrating,
staining, or clearing. A considerable number, twenty-five or more,
will be found useful. Those with the beveled surface ground, upon
which the name of the substance or the name or number of the
14 THE COCCIDAE t
insect can be written with an ordinary pencil, are the most
convenient.
The student should supply himself with a good stock of slides
and cover-glasses. The slides should be of the ordinary size,
75 X 25 mm., with ground edges. There are two kinds, those known
as thin or extra thin which have the advantage that twice as
many can be stored in an ordinary slide box, those known as thick
slides which require more space but they are not so easily broken.
The cover-glasses should be No. 1 as to thickness and not
over 15 mm. in diameter and preferably only 12 mm. The use of
small covers, where the great majority of the objects to be mounted
are minute, saves time in locating the specimens under the cover.
Slides and cover-glasses as they come from the dealer are
greasy and should always be cleaned before using. The slides
should be placed in water containing about five per cent of
ammonia or in 95 per cent alcohol acidulated with one per cent
of hydrochloric acid. The cover-glasses should be placed in acidu-
lated alcohol. They should be added to the solution one at a time so
that the solution will cover all parts and should remain in it for
five or six hours. The slides and covers should be wiped with a
towel or piece of cheese cloth and stored in covered jars until
needed.
Tools. — The person making preparations of coccids should be
provided with the following tools : —
1. Forceps. — The forceps should have fine smooth points
without corrugations of any sort. The spring should be weak.
This serves a double purpose, they will not tire the hand so
quickly and there is less danger of injuring the specimen, since
only slight force is needed to close them. Two pairs of such
forceps will often .be found convenient.
2. Dissecting Needles. — The needles with a fixed ebony or
aluminum handle with straight points are best. Two are essential,
more will be found convenient.
3. Embryo Knives. — Dissecting needles with a spear point,
one with a double cutting edge and one with a single cutting edge.
4. Section Lifter. — One with a small, thin, flexible blade.
5. Scissors. — A pair 115 mm. long with fine curved points.
6. Pipettes. — The rubber bulb should hold 2 cc. and the
glass tube should be long, straight, and with a large opening. One
at least for each kind of solution used.
7. Brushes. — Two artist's sable brushes of different sizes will
PREPARATION 15
be found very convenient at times for handling, transferring, and
cleaning specimens.
Clarifying. — The removal or dissolving of the wax excreted
by all or certain cells of the hypodermis and the making of the
body translucent is known as clarifying. This can be accomplished
by the use of several substances, the best of which is a solution of
caustic potash.
The body of the insect or the wax surrounding it is usually
closely attached to the host-plant. It must be detached in such a
way as not to injure the finer microscopic parts which are fre-
quently located along the margin of the body. This margin is
ordinarily distinct since the body is generally more or less
depressed in the adult female. In many species the characters
used for differentiation are the presence and absence and form
of minute setae or setae-like projections which are easily destroyed.
The specimens should be separated from the host-plant with a fine
dissecting needle under a binocular or simple microscope or a hand
lens. Where the body is encased in a homogenous mass of wax,
this should be placed directly into the clarifying solution. In the
Diaspidinae the body of the insect is covered by a composite
scale consisting of wax excreted by the insect and one or two cast
nymphal exuviae which are imbedded in the wax. The body of the
insect is located beneath the scale and can be exposed in most
cases by simply turning the scale over. In certain species the body
can be separated from the scale only with great difficulty, this is
due to the presence of a pronounced ventral scale. Where it is
difficult to separate the insect from the scale, the scale and the
enclosed insect should be placed in the clarifier. When the insect
is exposed, the dried body should not be lifted or handled by
holding it between the file-like or even smooth points of a pair
of forceps. Moisten the point of the dissecting needle or of the
forceps with water, alcohol, or caustic potash, or on the tongue and
lift the specimen with the moistened tool. It is frequently
desirable to mount some of the nymphal exuviae of the female
attached to the scale, since they frequently show characters not
found on the body of the insect itself. In certain genera of the
Diaspidinae the last or second nymphal exuvia is not molted and
it must be broken open to obtain the body of the insect.
The best solvent of the wax is a solution of caustic potash.
It is also the best substances for expanding the body and making
it translucent. A ten per cent solution, made by adding ten grams
of stick potash to ninety cubic centimeters of distilled water, is
16 THE COCCIDAE
the best strength to use. The effect of a very strong solution is
different from that of a weak solution which destroys the cellular
structures. Distilled water should always be used in making the
solution, if this is not available, fresh rain water should be used.
Water containing lime or tap water that has been filtered with
chemicals should never be used in making a solution of caustic
potash or used in washing specimens. When such water is used,
the various chemicals form linear or spine-like crystals in the
specimens. It is often desirable to use a weaker solution than ten
per cent, such a solution can readily be made by diluting the stock
solution one-half, one-third, or one-fourth as desired, making about
five or three or two per cent solutions.
In large species and those with globular bodies, it will often
be found desirable to separate the dorsal and ventral surfaces and
mount the two portions side by side under the same cover-glass.
Large species after they have been boiled for a time often have
large opaque masses within them. The body in such cases should
be punctured and the hard opaque mass removed. The source of
these masses is not known. If the specimen is left in the caustic
potash until the mass is dissolved, the specimen will be ruined.
The time necessary for the clarification of any particular species
varies with the individual. Some specimens of a species will clear
quickly while others will take considerable time. The specimens
should at all times be given careful attention. The length of time
required will need to be modified with each species. This can
be determined only by observing the rapidity of the clarification.
Those specimens in which the body is soft or the cuticle is
thin must be handled carefully so as not to puncture or tear the
cuticle unnecessarily, but particularly so as not to get the body
out of shape. This applies especially where the body is bag-like
in form. Species that normally have the body depressed are liable
to become distorted or decidedly compressed. Many specimens
become so distorted that considerable time is required to unravel
the relation of the various parts. This distortion is due to one
or both of two causes, carelessness in handling the specimens with
the forceps or carelessness in boiling them too vigorously, frequently
the latter. Specimens that are badly distorted, unless they are
very rare, are not worth the time required to mount them. When
the caustic potash for clarifying or the water for washing is boiled
vigorously, the specimens are forced to the surface of the liquid
and into the greatly agitated surface film. This has the effect of
flattening out the specimens, folding them compactly, and twisting
PREPARATION 17
and mixing the appendages. If there are several specimens in the
liquid, the mixing and fastening together may be so complete that
it is practically impossible to separate them. It is a waste of
time to try and untangle such specimens, they are worthless for
mounting, unless they are very rare species.
Since the above was written, one of my students, J. Howard
Gage, has shown that specimens, no matter how badly they may
be tangled and distorted, can be separated and inflated in the fol-
lowing manner. The specimens to be inflated should be washed
as described later and placed from the water or from alcohol in a
watch-glass containing chemically pure lactic acid. They should
remain in this solution from three-fourths to one hour and then
be placed in another watch-glass containing sulphuric ether or
chloroform, preferably the former, for fifteen to thirty minutes.
The inflation occurs while in this latter solution and when com-
pleted the specimens should be separated and as plump as in life.
An immersion in 95 per cent alcohol is needed to remove the ether
and for dehydration.
There is always difficulty, except in the case of large speci-
mens, in transferring them from one solution to another, particu-
larly if the transferring is to be done without injury to the speci-
mens. It should be done with a section lifter, pipette, sable brush, or
forceps. The forceps, while apparently more difficult to use,
will be found the best at all times for this purpose. The speci-
mens, whether large or small, should never be grasped between the
points of the forceps. This will break off fine projections or distort
the body. The specimen should be floated from the bottom and
taken up in the fluid enclosed between the points of the forceps.
This method will seem almost impossible at flrst, because of the
difficulty of getting the specimens, particularly small ones, through
the surface film. The knack of doing this is readily acquired, if
the preparator is persistent. It will soon be learned, if tried, that
it is not as easy as it may seem to transfer specimens with a section
lifter for the same reason. Float the specimens around in the fluid
after each change is made so that they will return, if slightly
distorted, to their original form.
The clarifying with hot caustic potash is the method most
generally followed and may be known as the fast method. The
use of a cold solution may be known as the slow method. The heat
hastens the action of the caustic potash and it is possible to make
preparations in a few hours by the fast method which would
require several days by the slow method.
18 THE COCCIDAE
The fast method consists in placing the specimens removed
from the host-plant in a casserole into which should be poured
enough caustic potash to cover the bottom to a depth of one-fourth
to one-half inch. The cover should be added and the solution
should be heated until it just simmers. If the solution becomes in
great part evaporated before the specimens are clarified, either
more caustic potash should be added or some distilled water.
The slow method differs from the fast method in that a cold
solution is used. The specimens clarified in this way will need
from eight hours to several days depending upon the size of the
specimens, the amount of wax, and their opacity. Either Syracuse
watch-glasses or four drachm homeopathic vials should be used.
This method does away with a great part of the handling of the
specimens and the chance of breakage from the boiling solution.
The fluids can be decanted off, if desired, and the specimens left
in the vials or watch-glasses. This method is not practical for
those species that require a long treatment, even by the fast
method, as many Coccinae. It is especially useful in preparing
specimens of Diaspidinae, which are usually clarified by a
treatment over night. The action can be hastened by pouring a
hot solution into the vials containing the specimens. Hot solutions
should not be used with the watch-glasses as the danger of breakage
from the heat is considerable.
Other substances can be used for dissolving the wax, as carbon
bisulphide, sulphuric ether, xylol, or chloroform. The last of these
is the only one of much value. The powdery or cottony wax, when
present in small quantities, can be dissolved with chloroform. It
does not expand shrivelled specimens and its most important use
is in preparing specimens of Diaspidinae where good prepar-
ations of the fringe are especially to be desired. Specimens can
be mounted in xylol balsam directly from the chloroform.
Washing. — The specimens after the clarifying is completed
need to be thoroughly washed in distilled water in order to remove
all traces of the caustic potash. If the fast method was used, the
specimens can be boiled in distilled water. The washing can be
done whether the fast or slow method was used, by soaking the
specimens for an hour or more in cold distilled water. They should
be transferred from the casserole, if the fast method was used, to
watch-glasses or vials, if already in such receptables, the caustic
potash can be poured off and replaced by water. The specimens
should be carried through several changes of distilled water, three
to five, whether they are boiled in it or not.
PREPARATION 19
Hardening. — When the washing is completed, the specimens
should be transferred to 95 per cent alcohol. If the specimens
are hardened by remaining in this solution for about twenty
minutes, they, particularly the soft baggy ones, will be greatly
benefited. This step can be omitted if desired.
Staining. — If the specimens are to be stained, the general rule
is that they should be transferred from the hardening solution to
the stain if an alcoholic stain is used and to water and then to
the stain if an aqueous stain is used. This will be found unneces-
sary if the stain recommended on a following page is used as
specimens can be placed in the stain from either water or alcohol.
While it takes more time to stain specimens, the greater ease with
which they can be studied and the much greater visability of many
characters that are just or not at all perceptible in unstained
specimens, more than repays for the extra time required.
Many solutions have been suggested, as safrannin, gentian
violet, picric acid, picro-creosote, Delafield's haematoxylin,
fuchsin, saurefuchsin, and carbolic fuchsin. Some writers have
confused fuchsin and saurefuchsin and their various commercial
names. Fuchsin is a basic stain containing rosanilin, and is
variously known as fuchsin, analin red, rubin, rosein, magenta,
etc. Saurefuchsin is an acid stain containing rosanilin and known
as acid fuchsin, fuchsin S, acid rubin, saurerubin, acid magenta,
and saurefuchsin. Gage, who experimented extensively with these
various stains and most of the others named, concluded that the
best results were obtained by the use of an aqueous solution of
saurefuchsin. As he pointed out, one of the great difficulties with
stained specimens of coccids is that they may be sharply stained
and well differentiated when mounted, but in the course of a few
weeks or months they have lost all trace of the stain. The fading
of saurefuchsin was shown to be due to the presence of caustic
potash in the specimens. The washing of the specimens needs,
therefore, to be done with great care and thoroughness and, if there
is any question as to the complete removal of the caustic potash,
the specimen should be washed in acidulated water or alcohol,
1 cc. hydrochloric acid to 99 cc. distilled water.
The staining should be done while the specimens are in vials
or watch-glasses. It will require from ten to forty minutes,
depending upon the size of the specimens and the condition of their
cuticle. If the specimens are overstained, the excess stain can be
removed with a dilute solution of caustic potash, one-half per cent
20 THE COCCIDAE
to one hundred ec. of water. This should be followed by acidulated
water or alcohol.
The following formula for Gage's Saurefuchsin contains
considerable hydrochloric acid in order to reduce any cautsic potash
there may be left in the specimens : —
Saurefuchsin 0.5 gr.
Hydrochloric acid, 10 per cent 25.0 cc.
Distilled water 300.0 cc.
The excess of stain should be removed by washing with
distilled water. The use of acidulated water will sharpen the color
of the stain. The longer the specimen has to remain in the caustic
potash beyond a certain time, the more difficult it will be to stain.
A stronger solution, one gram of saurefuchsin to the same
amount of water and hydrochloric acid, will be found advan-
tageous. This is particularly true in staining individuals with a
very thin cuticle and in staining specimens of Eriococcinae from
which the anal ring is to be dissected and mounted separately
under a cover.
Dehydration. — The removal of the water from the specimens,
if they are to be mounted in a resinous media, as Canada balsam,
is known as dehydration. The water must be removed because the
balsam and water are not miscible. If a nonresinous mounting
media is used, as glycerin jelly, dehydration is unnecessary. If
the specimens are to be cleared in a clearer containing carbolic
acid, the dehydration can be done in commercial or 95 per cent
alcohol, otherwise absolute or 98 per cent alcohol should be used.
The dehydration is best done in a vial or watch-glass. The speci-
mens should remain in the alcohol for at least fifteen minutes.
If a complete dehydration is not accomplished, the specimens
become cloudy or foggy, have a whitish aqueous bubbly appearance,
a short time after they are placed in the balsam. Time will be
saved by allowing the specimens to remain in the alcohol for a
sufficient length of time to insure a complete dehydration. If the
specimens become cloudy, remove the cover-glass and return the
specimens to the clearer and then give them a thorough dehy-
dration. They should then be returned to the clearer and mounted
in balsam again. The same alcohol should not be used too long,
renew the supply from time to time.
Clearing. — The placing of the specimens in a solution that is
miscible not only with alcohol but with the resinous mounting
media is known as clearing. It can be done in a watch-glass or a
PREPARATION 21
vial or on a slide. The specimens should remain in the clearer for
fifteen minutes or more. Where the specimens are to be mounted
in a nonresinous media as glycerin jelly, clearing is unnecessary.
The following substances can be used for clearing, chloroform,
xylol, cedar oil, oil of bergamont, benzol, turpentine, synthetic oil
of wintergreen, carbol-turpentine, carbol-xylol, and oil of cloves.
The last two give excellent results.
The oil of cloves can be used after dehydration with com-
mercial alcohol. It differs greatly in shades of color, a light
colored oil should be used. Specimens cleared with clove oil become
very dark colored with age.
One of the most practical clearers for use with coccids is
carbol-xylol. It can be used after commercial alcohol, clears
rapidly, evaporates slowly, and specimens do not discolor with age.
The solution is prepared by combining one part by measure of the
melted crystals of carbolic acid with three parts of xylol.
If it is inconvenient to complete the preparation of the
specimens immediately, they should be removed from the caustic
potash, washed, and stored in alcohol. A stay of a lew hours in
alcohol will not injure stained specimens, but a stay of two or
three days in alcohol or carbol-xylol does effect the intensity
of the stain. An examination of stained cleared specimens of
large individuals in a watch-glass is often very advantageous.
Such specimens should be placed in clove oil for study. If the
specimens are to be mounted after the completion of the exam-
ination, they should be placed in 95 per cent alcohol to remove
the clove oil and for dehydration and then cleared in carbol-xylol.
Mounting. — The specimens after clearing are ready for mount-
ing in a resinous mounting media, as Canada balsam. They
in most cases are minute and the use of cover-glasses one-half inch
or twelve millimeters in diameter will be found more economical
of time when the specimens are studied. Several specimens, except
in the case of large species, can be placed under a single cover-glass.
The specimens should be arranged in a row, thus | | | | and
sufficient very thin balsam to cover them added. The balsam
should be allowed to harden for a short time and fasten the
specimens in place. When sufficiently hardened, add enough
balsam to fill the space under the cover and carefully put the
cover-glass in place. Do not use more balsam than is absolutely
necessary, the preparation must be thin if an oil immersion
objective is to be used. In order to secure thin preparations, the
balsam should be diluted until it is about as thin as water. If
22 THE COCCIDAE
spaces unfilled with balsam appear a few hours after completing
the mounting, put a minute drop of the balsam at the edge of the
cover adjacent to the unfilled spaces and allow it to run under the
cover. If the balsam gets thick, it can be thinned by the addition
of xylol, chloroform, or benzol, depending upon the solution
originally used to dissolve it. The solution in xylol is the one
most generally used. Specimens should never be mounted in the
natural or unthinned Canada balsam. Such balsam will never
harden and the preparations, where it is used, are always soft and
sticky. When the mounting is completed or before the slides are
handled much, they should be heated gently by placing them on a
steam radiator or over an alcohol lamp to drive off the xylol and
to fasten the cover-glass in place. Care must be used in heating
with a lamp not to allow the balsam to boil and fill the preparation
with bubbles.
Specimens mounted in glycerin jelly have the advantage of
being more opaque than those mounted in balsam. Such prepar-
ations are likely, especially if stored in a warm place, to have the
mounting media disintegrate and the preparations become worth-
less for study. Such specimens can be remounted by placing the
slides in hot water and dissolving the glycerin jelly. They can
then be remounted in the glycerin jelly or dehydrated and cleared
and mounted in balsam. The prepared glycerin jelly can be pur-
chased from dealers in microscopic supplies.
Labelling. — Each slide should be carefully labelled. This
should be done as they are completed. The labels should be written
with India ink or waterproof ink. Ordinary writing fluids become
illegible in a few years. It is always of great advantage to marl:
the slide in such a way that the mounted specimens can be con-
nected up with the original lot of material from which they vvere
obtained.
Rapid Method. — ^While many entomologists depend upon the
external appearance of the scale in the Diaspidinae, the only abso-
lutely safe method of idenification is a study of the characters of
the body of the insect itself. Considerable time is required to make
a preparation for such a study. A preparation showing the pygidial
fringe will often serve every purpose. Glycerin jelly alone or
combined with glacial acetic acid, nine parts of the glycerin jelly
to one of the acetic acid, is used for making such preparations.
The acetic acid acts as a clearer and its addition is an advantage.
The specimens should be removed and placed on a slide with just
sufficient of the glycerin jelly to mount the specimens. The slide
PREPARATION 23
should be heated slowly over an alcohol lamp until the glycerin
jelly is melted and surrounds the specimens. The heating should
be continued until the mounting media has thoroughly penetrated
the specimens, gently boiled if necessary. Care must be used
not to get the slide too hot and burned or heated until the glycerin
jelly is all evaporated. The cover-glass should be added when
the heating is completed. The pygidial fringe is easily followed in
such specimens and often other structures of the pygidium can be
identified.
The following formula shows the procedure for the potash
method : —
Eemove Insect from Host-plant
Treat with Caustic Potash.
Wash in Distilled Water.
Harden in 95 per cent Alcohol.
Stain in Alcoholic or Aqueous Stain.
Dehydrate in 95 per cent Alcohol.
Clear in Carbol-xylol.
Mount in Xylol Balsam.
Label Slide.
CHAPTER III
EXTERNAL ANATOMY
The sclerites and sutures of the Hemiptera even in the species
of the most generalized families are generally modified in form
and distinctness from the same parts in all insects of the orthop-
teran type. In the specialized Gularostria the sutures are effaced
for the most part. The intersegmental coriae are frequently
wanting in the Coccidae and, when present, are usually indicated
only as slight transverse furrows. A study of the external anatomy
of the coccids consists in an examination and identification of the
body regions, the appendages, and the numerous new or secondary
structures that have arisen as a result of their peculiar mode of life.
The body of a coccid, while different in external form from
that of most Hemiptera, has retained certain characters showing
conclusively their relationship to the other families of the order.
The variation in external structure is not limited to differences
from the other families of the order, but there is a great dissim-
ilarity in form even among the various subfamilies of Coccidae,
particularly among the adult females. This difference has un-
doubtedly resulted from the dissimilarity in their habits and from
the diverse ways in which the females shield their progeny. Eggs
are laid in many species and they are protected by a waxy covering
which varies greatly in form, consistency, and position in the
different genera. With the variation in the waxy excretion there
is a corresponding variation in the structures of the body of the
insect that produces the excretion.
The two sexes, as pointed out, are very different in size, form,
and structure. The adult males are distinctly insect-like in form.
Their parts are normal in form and arrangement and do not differ
markedly, except in the loss of mandibles and maxillae, from those
of other winged insects. The adult females on the contrary are
usually more or less grub-like in form. They possess many struc-
tures not found in the males. The classification is based almost
entirely upon the females and the following descriptions unless
otherwise stated refer only to them.
Body-wall. — The exterior covering or body-wall of coccids is
composed of the three layers common to other insects, cuticle,
hypodermis, and basement membrane.
24
EXTERNAL ANATOMY 25
The hypodermis is a single layer of pavement epithelium or
cuboidal cells. It is the living part of the body-wall and excretes
the cuticle and the wax in which the body is imbedded or by which
it is covered. The wax varies in form according to its method of
excretion. In such genera as Ceroplastes and Tachardia, where
the wax is a homogenous mass, it is considered as excreted by all
of the hypodermal cells and as poured out through the cuticle.
In most coccids the wax is excreted by special cells of the hypo-
dermis which have been greatly modified and enlarged. Careful
detailed studies of the form of the wax excreting hypodermal cells,
the wax cells, has been made in only a few species. Such cells
are usually more numerous during the adult than during any of
the nymphal stages and are fewest in number during the first
nymphal stage. In many genera certain wax cells are wanting
or are not functional until during the adult stage as the
genacerores of the Diaspidinae. The openings of these cells are
usually located on the ventral aspect of the abdomen or are
associated with the external opening of the oviduct, the vulva.
They excrete the wax in which the eggs are enclosed.
The inner ends of the hypodermal cells are bound together
by a thin semicuticular layer, the basement membrane. This
membrane, so far as is known, does not perform any important
function in the Coccidae.
The outer layer of the body- wall is the outer skeleton, cuticula,
or cuticle. It is generally comparatively thin and flexible in this
family. The cuticle and the internal skeleton are the only portions
not only of the body-wall but of the body remaining after the coccid
has been subjected to an extended treatment to caustic potash.
It is, therefore, incorrect to consider any of the parts remaining
after such a treatment as glandular structures or apply names to
them that would suggest such a condition. When the body is
studied in section, the cuticle is found to be indefinitely lamellate.
The outer thinner portion, hardened by the deposition of chitin-
is usually darker in color and inelastic. It is the outer portion
that is discarded when the insect molts. The much thicker inner
portion is elastic and permits of the increase in size or growth of
the body at stated intervals when the outer layer of cuticle is cast
off or molted. The cuticle covers not only the outer surface of
the body but lines the cephalic and caudal portions of the lumen of
the alimentary canal, the air tubes or tracheae, the ducts of the
salivary glands, and the caudal portion of the ducts of the repro-
ductive organs. It also forms the basis of the internal skeleton
26 THE COCCIDAE
or endoskeletoii of the head and thorax when these are present.
Setae, Spinulae. — The cuticle bears appendages of different
kinds, as setae, spinulae, and cerores, the openings through which
the wax is poured from the wax cells.
Each of the hair-like appendages borne by the cuticle is a
seta. They are generally designated as hairs or spines although
they are entirely different in origin from both of these structures.
The setae are hollow, open at the distal end, and are formed by
special hypodermal cells. Each seta consists of a slender thread-
like portion, the shaft, which is articulated in a small cup in the
cuticle, the alveolus or calyx. It appears as a depressed circular
area when the shaft is in place and when broken off the shaft
shows as a minute irregular stub in the bottom of the depression
The presence of the calyx is the only certain means of proving
whether any given projection is a seta or not. The opening in the
shaft of the seta, the lumen, connects with an opening extending
through the cuticle, the pore canal or trichopore, which terminates
at the large cell secreting the seta, a trichogen cell. This cell
forms the seta and in most cases is apparently formed by the
fusion of several adjacent cells which have lost their body-wall,
since the trichogen cell always contains several independent masses
of nuclear matter. The setae are probably primarily sensory in
function and are connected with the nervous system. The trichogen
cells often elaborate an excretion which is poured out through the
pore canal and the lumen of the seta. The wax of coccids is never
excreted from cells of this type so far as known. The cuticle
bearing each seta is sometimes produced pimple-like, as in Icerya.
The swollen portion of the cuticle together with the seta which it
bears is known as a chalaza. The extent of the cuticle can always
be determined from the position of the calyx. The wax excreted
by the wax cells is never poured out through setae, the seta-like
projections of Orthezia lack calices. The spines of insects are
extensions of the body-wall into which the hypodermis extends
and frequently bear setae. True hairs are found only in the
Vertebrata.
The surface of the cuticle may be prolonged into numerous
minute projections which are frequently seta-like. These projec-
tions are known as solid setae, solid hairs, or spinulae. They lack
not only the calyx of the seta but also its lumen, are not formed
by special hypodermal cells, and are usually inflexible.
Cerores, Ceratubae. — The setae and spinulae are projections
of the cuticle. It usually contains in addition to these in the
EXTERNAL ANATOMY 27
family Coccidae minute microscopic openings through which the
wax is poured. These openings vary greatly in form and
arrangement, are sometimes borne on minute projections, and are
known as cerores. They are also known as pores, glandular pores,
wax pores, or spinnerets. The cerores are sometimes Forne on
minute projections of the cuticle and for this reason have been
considered as appendages of the cuticle.
The simplest and probably the primitive type of ceroris is one
that consists of a single opening and in which the periphery is
flush with the ectal surface of the cuticle. Such cerores are present
in the Coccinae and Eriococcinae. The great majority of the
cerores in the Ortheziinae are at the ends of seta-like prolongations
of the cuticle. This cuticular extension, which lacks a calyx, has
its opening in the free end and is a pilaceroris. One of the
commonest types of ceroris is where there are six openings arranged
in a circle around the periphery of a central circular area. These
are known as paracerores. The cerores located in the rectum, the
anacerores, are also known as rectal wax pores, rectal spinnerets,
or honey dew glands. The powdery wax in most species of
Pseudococcus is poured out through cerores that are triangular in
outline, the tricerores, and contain numerous irregularly
arranged pores. In the Asterolecaniinae the cerores are arranged
in pairs with their adjacent margins usually flattened so that they
form a figure 8. These are the octacerores. The opening or
openings of each ceroris through which the wax is poured is an
oraceroris.
In the Diaspidinae the openings through which the
wax is poured are generally designated as wax pores or spinnerets.
These openings are the external outlets of invaginated cuticular
tubes and are not homologous with the cerores. The tubes are
known as ceratubae and their external openings as oraceratubae.
The ceratubae are not confined to the Diaspidinae, but have been
observed in species of Lecaniinae, Kermesiinae, Eriococcinae, and
Asterolecaniinae. They will probably be found in some of the
other subfamilies. So far as observed, the ceratubae of each sub-
family are distinctive in form. They are difficult to observe in
unstained specimens.
Segmentation. — The cuticle is divided by transverse constric-
tions, coriae, into three well marked regions, the head, the thorax,
and the abdomen. The segmentation of the head is lost during
embryonic development, but the three segments of the thorax and
the eight or more segments of the abdomen can usually be
28 THE COCCIDAE
identified, particularly in the early nymphal stages. The con-
stricted part of the cuticle, the coria, is usually flexible and marks
the division between adjacent segments. The coriae are named
from the segments forming their caudal boundary, as the one
cephalad of the prothorax is the procoria, the one cephalad of the
mesothorax is the mesocoria, the one cephalad of the first abdominal
segment is the unacoria, etc.
The segmentation of the body of the adult coccid is greatly
obscured through the obliteration of the coriae by the expansion of
the body to hold the ovaries or because of its covering of wax or
probably in most cases to both of these causes. In the first nymphal
stage of certain species the segments and coriae can generally be
identified, but they gradually disappear with each successive molt.
The microthorax can not be identified and the microcoria, micro-
thorax, and procoria are represented by a single continuous
transverse area, which is separated from the head by an infolded
area which has been designated the procoria. Where the thoracic
coriae are wanting, the thoracic region can be identified from the
attachment of the legs when these are present. But where the
coriae are obscure or wanting and the legs are wanting, the extent
of the thoracic segments can be approximated only by the attach-
ment of the mouth-parts and the location of the thoracic spiracles,
the mesospiracles and the metaspiracles.
Head. — The head is an area of considerable extent on both
dorsal and ventral aspects. It typically bears a pair of antennae,
a pair of simple eyes, a rostrum or labium, and a pair of bristle-
like mandibles, and a pair of bristle-like maxillae. These are
usually all present in the first nymphal stage even though all or a
part of them are wanting in all the later nymphal stages and the
adult. The caudal extent of the head is marked by the procoria,
but this is wanting in many adults, as Diaspidinae and Lecaniinae,
where the head and three thoracic segments form a continuous
undivided area. This fused area is designated by some writers as
the cephalothorax. Something as to the size and shape of the
head can frequently be determined from the appendages which
it bears and their attachment. The head is generally figured and
described as being triangular in outline and limited for the most
part to the ventral aspect. Its true size and shape are more
easily determined in nymphs of the first stage. The subtriangular
ventral area frequently described as the head is only a part of it
and represents the fused front and clypeus. It is designated as
the frons in the Homoptera and this name should be used here.
EXTERNAL ANATOMY 29
The labrum so far as observed is always obsolete. The cephalic
portion of the ventral aspect of the head bears the eyes and the
antennae and the caudal portion the mouth-parts. The latter are
attached between the first pair of legs and usually in line withr the
first pair of spiracles.
Eyes. — The eyes, when present in the female, are always
simple. They are comparatively large in the active species and
sometimes, as in Orthezia, are placed on short projections. The
portion representing the eye can usually be recognized even in
treated specimens from the presence of pigment. The eyes of the
female are sometimes designated as the primary eyes, because they
persist through out all the nymphal stages and are present in the
adult. The eyes of the male are compound in the Monophlebinae,
the Xylococcinae, the Margarodinae, and the Ortheziinae. The
external part of each separate compound eye, a facet, is strongly
convex so that the compound eye appears as a whole coarsely
granular. In the subfamilies with compound eyes certain species
are also provided with two simple eyes or true ocelli, which are
sometimes located between the compound eyes but more often
adjacent to their caudal margin. In the other subfamilies the
compound eyes are represented by groups of simple eyes which
are arranged in a semicircle on each side of the head. The number
of simple eyes varies greatly in the different genera. They are
most numerous in the Phenacoleachiinae, eight on each side,
where they form almost a complete circle, but the usual number is
from three to five on each side. There is usually a pair of large
facets on the dorsal and another on the ventral aspect. It has
been claimed by some that the ventral pair of facets is developed
from the mouth-parts which are always stated to be wanting. In the
wingless males the eyes are limited to two facets, one on each side
of the head as in the female, but there is no reason other than
similarity in position for considering them homologous with the
primary eyes of the female.
Each of the simple eyes of the nymphal and adult female and
of the first nymphal male and of the adult male with simple eyes
is known as an ocellana. This term is applied to all the simple
eyes, other than ocelli, of all Exometabola. In order to distinguish
the simple eyes other than ocelli of other insects, those found in
nymphal and adult Apterygota are known as ocellalae and those of
the larvae of Entometabola as ocellarae. The position of one pair
of ocellanae on the dorsal aspect of the head of certain males,
which are not in line with the other ocellanae, would suggest that
30 THE COCCIDAE
they are probably true ocelli. In the Diaspidinae there are three
pairs of ocellanae, a pair of large dorsal and a pair of large ventral
ocellanae and a pair of minute subdorsal ocellanae. These latter
are designated as eyes by Newstead and the large ocellanae as
ocelli while in Lichtensia, where there are five pairs of ocellanae,
two pairs of ventral ocellanae, two pairs of dorsal ocellanae, and
a pair of lateral ocellanae, the latter are considered as eyes and all
the other ocellanae as ocelli. True ocelli are never present in
immature insects of this order and when present in adults are never
more than three in number. A very different nomenclature is used
by Signoret for the ocellanae. There is no reason for considerino;
any of the pairs of ocellanae as eyes or the homologues of primary
eyes. The pair of small ocellanae in the Diaspidinae and one
pair in other male coccids may prove to be homologous with the
true ocelli of other insects.
Antennae. — The antennae are long and prominent in the
active species and usually consist of about eleven segments. This
number is greatly reduced in the inactive species where the
antennae may be wanting. In the Diaspidinae each antenna is
generally represented by a minute tubercle bearing from one to
four long setae. Where the antennae are large, the distal segment
is usually much larger than the others and is sometimes thickened
or clavate and irregularly indented or notched. The notches cause
this segment to appear as if subdivided or composed of several
subsegments. These have been described by some writers as true
segments. The indentations mark the location of the attachment
of setae. The comparative length of the various segments of the
antennae, particularly in the Eriococcinae and Lecaniinae, have
been used extensively for the separation of genera and species.
The variation in the length of the different segments is usually
expressed in the form of a formula, thus: (3, 4), 7, 2, (1, 6).
The numbers are the numbers of the segments, the longest segment
is always indicated at the left end of the formula and the suc-
cessively shorter ones are arranged in order from left to right
while those of the same length are enclosed in a parenthesis. A
formula giving the actual length in microns is used by some
authors. Kellogg and Bell have shown from an examination of
several individuals from the same colony that there may be enough
variation between the segments of the two antennae of the same
individual to place them in different genera. This was confirmed
by Thro from a study of certain species of Lecaniinae. The
number and arrangement of the setae borne by the antennal
EXTERNAL ANATOMY 31
segments has been much used by systematists. Careful studies of
representative species of several genera of Lecaniinae by Thro
showed that the setae were practically identical in number and
arrangement in the species studied and the differences described
were due to overlooking the calices of setae that had been broken
off. In certain genera there is a group or transverse row of setae
located on the ventral aspect of the head between the articulations
of the antennae which have been named by Green the interantennal
setae.
Rostrum. — The rostrum in the heteropterous Hemiptera has
been shown to be homologous with the labium of other insects.
That the rostrum in the Coccidae is homologous with the rostrum
of the Heteroptera and the labium of other insects, there is abso-
lutely no doubt. It is an elongate projection attached between
the prothoracic legs. The lateral margins of the rostrum are
normally folded so that it is U-shaped in cross-section except that
the margins meet on the dorso-meson. By this folding a closed
furrow is formed, the rostral furrow, which extends lengthwise
through the rostrum and serves for holding, guiding, and support-
ing the bristle-like mandibles and maxillae. The rostrum consists
of three segments in Coelostoma and Phenacoleachia ; of two
segments in Icerya, Pseudococcus, and many other coccids ; and of
a single segment in the Lecaniinae and Diaspidinae. In the adults
of the Margarodinae, Xylococcinae, Callipappinae, and Kuwaniinae
the rostrum is always wanting. It is usually stated that the rostrum
is lost in the male at the first or second molt and is wanting in all
subsequent stages, including the adult. But in the adults of cer-
tain species as those of the genus Pseudococcus, it is represented
by a minute conical projection. The distal end of the rostrum is
frequently provided with large setae, the labial setae. In the
Coccidae the rostrum is always much shorter than in the closely
related families, the plant lice and the psyllids, rarely if ever
extending beyond the thorax and usually only to the caudal margin
of the mesothorax.
Mandibles, Maxillae. — The mandibles and maxillae are long
and bristle-like as in all other Hemiptera. It is very difficult, if
not impossible in most cases, to distinguish the mandibles from the
maxillae, except where their articulation to the head can be de-
termined. They can be distinguished in some heteropterous
Hemiptera by the teeth or projections on their distal ends. The
margins of the mandibles and maxillae are furrowed in such a
way that they are normally so closely fastened together that it is
32 THE COCCIDAE
difficult to separate them. When they are separated, which some-
times happens from the handling of the specimen, there is
frequently apparently less than four parts present, the actual
number. The usual number is apparently three. The mandibles
in such cases are separated from each other and from the maxillae.
They form two of the three parts and the closely applied maxillae
the third. They are normally all fastened together and form a
tube through which the liquid food is drawn into the pharynx.
This structure, formed of the closely applied mandibles and
maxillae, is the rostralis. It is frequently longer than the entire
body and may be five times as long. The rostrum serves simply
as a support for the rostralis and does not function as a tube
through which the food is drawn. The free end of the rostralis
in individuals which are not attached to their host-plant usually
does not project beyond the free end of the rostrum. The mandi-
bles and maxillae are lost at the first or second molt and are
wanting in the adult male.
Crumena. — The thin cuticle of the head adjacent to the
proximal portion of the rostrum is deeply infolded or invaginated,
forming a long slender pocket which lies free in the body cavity
and generally extends through the mesothorax and metathorax.
The rostralis after its exit from the mouth extends into this pocket.
This pocket was first described by Mark and by him named the
crumena. It was later figured and described by Berlese who
designated it the guaina delle setole rostrali. I had before the
name of Mark was discovered designated it as the theca. All the
slack of the rostralis is held in the crumena and the proximal part
of the rostral furrow is thickened and forms a sort of bumper, the
plena, against which the end of the rostralis rests. The crumena
has been observed in many coccids and is probably present in all.
It is difficult to identify in unstained specimens. The rostral fold
described by many systematists is the portion of the rostralis
retained within the crumena.
Endoskeleton. — There is a well developed series of chitinized
arms or bars present within the cavity of the head of all coccids.
These bars are a part of the endoskeleton and are usually desig-
nated as the head skeleton or chitinous cradle. The parts of the
endoskeleton serve for the attachment of the muscles that operate
the rostralis, rostrum, and the parts associated with them. In the
differentiation of species or genera, the parts of the endoskeleton
have not been used. There is only a slight variation in the form
and arrangement of these parts in .the different subfamilies.
EXTERNAL ANATOMY 33
Thorax. — The thorax is divisible into three clearly marked
regions in most insects, the prothorax, the mesothorax, and the
metathorax. Such a condition does not exist among coccids. The
prothorax is usually continuous with the head, except iur some
nymphs of the first stage, and with difficulty separated from it.
The mesocoria and metacoria, the membranes connecting the pro-
thorax and mesothorax and the mesothorax and metathorax, are
usually limited to the dorsal and lateral aspects and then only
marked as slight transverse constrictions. These coriae are effaced
without passing through a suture stage. The combined head and
thorax, as in the adult female of the Diaspidinae, are sometimes
designated as a cephalothorax.
There is a strongly chitinized area extends dorsad and laterad
from the lateral margin of the proximal segment of each leg, a
coxa, in certain of the leg bearing species. This linear area has
been designated the episternum because it bears a projection for
the articulation of a coxa. This articular surface of each epis-
ternum is a coxacoila and the articular surface of the coxa which
is applied against the coxacoila is a coxartis. In those Hemiptera
where the pleural sclerites can be identified, the episternum is
usually large and forms the greater part of each pleural area,
while the epimeron is small and inconspicuous and forms a narrow
area along the caudal margin of the episternum or is covered hy it.
The coxacoila is typically formed from the episternum and epimeron
not only in the Hemiptera but in most insects, so that this area in
the Coccidae undoutedly contains a small portion of each of these
sclerites. For the sake of brevity, however, each of the entire
areas bearing a coxacoila has been called an episternum. This
sclerite is distinct in Icerya.
Those species of coccids provided with an episternum are also
usually provided with an apodema. This is the thin, vertical,
cuticular plate extending into the body cavity from the inner
surface of each episternum. These pleural plates form a part of
the endoskeleton of the thorax, which is present in most insects,
but wanting in all legless coccids. This structure has no relation
to the notal area of the male, the mesoscutellum, to which the name
of apodema has been incorrectly applied. There is on each side
of the meson of the ventral aspect of each thoracic segment of
most insects a small invaginated lobe, a furca. The furcae also
constitute a part of the endoskeleton of the thorax. In the coccids
the furcae are present only in the generalized species and then are
represented only by small finger-like projections which are difficult
34 THE COCCIDAE
to identify even in stained mounted specimens. The mesofurcae
and metafurcae are present in Icerya. The furcellae another
series of invaginated lobes forming a part of the endoskeleton of
the ventral aspect have not been identified. Each of the lobes of
the endoskeleton, apodemae or furcae, which are primitive struc-
tures, is often designated as an apodeme ; while other lobes which
are identical in form and structure but are secondary in origin
are known as parademes.
Legs. — The legs are of the ordinary type and sequence of parts
of other insects. They appear much larger and longer in the
nymph of most species than in the adult because of the size of the
body. The profemur is greatly enlarged and dilated in the Mar-
garodinae and the protibia is fitted with a hook-like tarsus for
digging.
The proximal segment of each leg, a coxa, is subglobular and
its proximal end is usually provided with a thickened rim. The
middle of the lateral margin of this rim or ring is provided with
an articular surface, an artis or coxartis, which articulates with a
coxacoila. The second segment or trochanter is oval or triangulai
in outline. The largest sclerite or segment of the leg is the third
segment or femur. It is usually more or less compressed. The
suture between the femur and the trochanter is obsolete in certain
species of Orthezia. The fourth segment, the tibia, is usually longer
than the femur but smaller in diameter. The fifth segment is the
tarsus. Its distal end frequently bears long slender clavate setae,
which are known as tenant hairs, digitules, or empodial hairs.
They are probably the outlets of glands secreting a sticky substance
which aids them in clinging to smooth surfaces. The digitules
are more frequently present in the males than in the females.
The suture between the tibia and tarsus is always wanting in
species of Conchaspis and in certain species of Orthezia. The
tarsus consists of a single segment except the protarsus of the
males of Coccus and all the tarsi of the females of Exaeretopus,
which have two. The hook borne by each tarsus, a claw, is usually
not prominent and in some species bears digitules. The digitules
of the legs are distinguished as the tibial digitules and the ungual
digitules.
The comparative length of the various segments of the leg
is frequently indicated by systematists and used in the separa-
tion of species. This is usually stated in microns in a formula, as
follows : c. 120-140 ; tr. 68 ; f em. 180-240 ; tib. 120-152 ; tar. 80-88 :
claw 12-12.
EXTERNAL ANATOMY 35
Abdomen. — The third, largest, and most important region of
the body is the abdomen. It is of importance because it contains
the organs of reproduction and their external opening and the
greater part of the structures that excrete the wax for covering tlie
body and the eggs. The abdomen consists typically of ten seg-
ments. In the Diaspidinae the segments of the caudal half are
fused and the coriae are wanting. Systematists have made no use
of the number of abdominal segments and usually those who have
described and figured the abdomen have not always been consistent
in giving the same number for the same species. The dorsal aspect
of each segment is known as a tergum and the ventral aspect as a
sternum.
The abdomen in the Diaspidinae is divided into two regions.
The cephalic portion consists of four segments with distinct flexible
coriae, the preabdomen, and a strongly chitinized caudal portion
without coreae, the pygidium. The adult females of the Conchas-
pinae are described as having a pygidium, but the coriae can be
identified between practically all of the segments and they are
provided with cerores instead of ceratubae as in the Diaspidinae.
In the abdomen of the adult Lecaniinae all the segments are fused
and the coriae obsolete, but the coriae can frequently be identified
in the nymphal stages, particularly the first.
The segmentation of the body can be determined or approxi-
mated in many species of coccids even if the legs are wanting
and the intersegmental coriae are obsolete. This can be accomp-
lished by means of the row of setae, cerores, or setae and cerores
placed along each lateral margin or around the periphery of the
body. The setae of these rows are generally larger or different
in form from the other setae of the body so that they can be
readily recognized and their intersegmental homology determined.
In the first stage nymphs of Icerya these structures are large setae
placed on prominent chalazae while in Pseudococcus they are
groups of short conical setae, usually one to six, surrounded by the
tricerores which form the prominent pencils of wax projecting
from the periphery of the body in all the nymphal stages and the
adult. While there is some variation among the different species
and genera as to the number and nature of these groups, they are
always constant for a given species. The usual arrangement is a
single group on each side of each abdominal segment, two to three
groups on each side of each thoracic segment, and four to six
groups on each half of the head. The components of the cephalic
36 THE COCCIDAE
group of the head and the caudal group of the abdomen are
generally subadjacent.
Anus. — The external opening of the caudal end of the alimen-
tary canal, the anus, is normally located in the tenth segment.
In the Diaspidinae the caudal portion of the abdomen is strongly
depressed and the anus is usually a circular opening located on the
dorsal aspect. It varies considerable in position, in some species
it is located near the cephalic portion of the strongly chitinized
pygidium, in others near the caudal end, and in still others it is
intermediate in position between these two. In the Lecaniinae
and Asterolecaniinae the anus is located at the cephalic end of the
invaginated barrel-shaped anal tube. It is usually surrounded by
an anal ring at this point and is not located on the exterior of the
body as might be expected.
Vulva. — The external opening of the female reproductive
organs is the vulva. It is located on the ventral aspect generally
near the caudal margin of the sixth segment. There are frequently
accessory structures associated with it which make its location more
easily identified. Figures of the caudal portion of the dorsal or
ventral aspects of the abdomen of coccids generally show the
position of both anus and vulva on the same drawing, this does not
mean that they are both situated on the same aspect. In the
Diaspidinae the vulva is usually located near the middle of the
pygidium.
Spiracles. — The external openings of the respiratory system.
the spiracles, also known as stigmata, are usually distinct and are
always four in number. The cephalic pair, the mesopiracles.
belong to the mesothorax and the caudal pair, the metaspiracles,
to the metathorax. These spiracles are normally located in the
mesocoria and metacoria. In the leg bearing species, the
mesospiracles are located about midway between the articulations
of the prothoracic and mesothoracic legs and the metaspiracles
about midway between those of the mesothoracic and metathoracic
legs. In the Diaspidinae the adults of which are always legless,
the mesothoracic spiracles are located one on each side of the
rostrum, sometimes designated as the head spiracles, near where the
prothoracic legs are ordinarily^ larticulated. The metaspiracles
are apparently not different in location from that of the species
with legs. The spiracles are the only landmarks available in
many legless coccids for indicating the thoracic segmentation.
They are greatly reduced in size in the genus Xylococcus and were
EXTERNAL ANATOMY 37
described by Pergande as wanting, while in Taehardia the
mesospiracles leave their normal position during nymphal devel-
opment and through a readjustment of the bag-like body assume
a position near the anus.
Peritreme. — The spiracles of most insects are surrounded by
a thickened area of cuticle which forms a rim about the opening
and is known as a peritreme. This rim varies somewhat in form,
but in most coccids is simply a circular plate or may be wanting.
The portions of the peritreme bordering the two sides of the
opening or slit, the cephalic and the caudal, may be modified into
lips or labiae for closing the spiracle.
Spiracularia. — The prominent parademe extending from the
ental surface of the peritreme into the body cavity in certain
coccids may be known as the spiracularia. They are distinct in
Icerya and Monophlebus, serve for the attachment of muscles, and
are so far as known peculiar to the thoracic spiracles.
Cavaera. — The tracheal tube connecting with each spiracle, a
spiracular trachea, is frequently expanded in the coccids into a
chamber of varying size and shape. This chamber is usually
trumpet-shaped or globular and its walls are generally smooth,
not transversely striate like the ordinary tracheal tube. This
structure may be known as the cavaera. It is also known as the
collar chamber. The cavaera constitutes the greater part of the
structure usually described and figured by students of coccids as
a spiracle. This is particularly true of species of Lecaniinae.
The term spiracle is limited here as with other insects simply to the
opening. Thisre are folds sometimes present in the cavaera which
serve for closing the spiracle, these should not be confused with
the labiae which are modifications of the peritreme.
Abdominal Spiracles. — The abdominal spiracles are generally
smaller than those of the thorax and may be provided with or lack a
distinct peritreme. The normal number of abdominal spiracles
for insects in general is eight pairs, a pair on each of the first eight
abdominal segments. When less than this number is present, it
is due to the reduction or fusion of the first abdominal segment
with the adjacent segment and the loss of its spiracles or the
atrophy of the spiracles or their loss through the increase in size
and the encroachment of the metathorax or the atrophy of the
spiracles at the caudal end of the abdomen. Since the abdomen
is not used extensively in classification, it is not strange that the
number of records of the presence of abdominal spiracles in coccidf^
is not large. They have been recorded more frequently from
38 THE COCCIDAE
among the generalized subfamilies, particularly the Monophlebinae,
than from the specialized subfamilies. Eight pairs are recorded as
present in species of Stigmacoccus, seven in Perissopneumon,
Monophlebus, and Droschia, and two or three in Icerya on segments
six to eight or seven to eight. Abdominal spiracles are also re-
ported as present in Cryptokermes, Margarodes, and on the first
segment of Conchaspis. All the species of Orthezia examined have
seven pairs and there are eight pairs in all the American species
of Xylococcus. Abdominal spiracles are also present in certain
species of Steingelia, Stomatacoccus, Coelostomidea, and Kuwania.
Cavaerae are rarely if ever associated with the abdominal spiracles.
Anal Ring. — The anus in certain genera is surrounded by an
elevated ring-like structure, known as the anal ring. It was
formerly known as the genital ring or genito-anal ring from the
mistaken notion that it surrounded the vulva. The anal ring
generally bears a number of setae, typically six, which are longer
and stouter than the other setae of the body. There are cerores
which vary in form and position in the different genera and species
which are arranged irregularly or in rows between the setae.
The wax excreted by the anal ring is generally different in
character from that excreted by the cerores on other parts of the
body and frequently forms a tube through which the honey-dew or
excrement is passed. The prominent anal wax-tube formed by
species of Xylococcus is not excreted in this way but by anacerores.
The anal ring is usually wanting in the adults of the Kermesiinae
but present in a rudimentary condition in the nymphs.
Anal Ring Setae. — The setae borne by the anal ring are knov/n
as the anal ring spines, the anal ring hairs, or the anal ring setae.
They are usually six in number. There may be as few as two and
as many as eight, ten, twelve, or even thirty. Newstead reported
tv/enty to twenty-four in Aclerda, but Green does not consider
these as anal ring setae. Some authors have recorded a large
number of anal ring setae as present in certain genera of
Lecaniinae, but these are according to Green longitudinal corruga-
tions of the eversible sac of the rectum, the anal tube.
Anal Lobes. — Most coccids with an anal ring are also provided
with anal lobes. These are the shoulder-like projections or lobes
located at the caudal end of the abdomen on each side of the anal
ring. They may be present in the nymphal stages and wanting in
the adult, in fact they are usually larger and more prominent in
the nymphal stages. They vary greatly in form and size. In some
species they are so large that the caudal end of the abdomen
EXTERNAL ANATOMY 39
appears cleft or emarginate. Each anal lobe bears one or more
large setae, the anal setae. These setae probably serve as a core
to support the wax filaments excreted by the cerores of the «nal
lobes.
The presence of an anal ring with anal ring setae and of
anal lobes with anal setae seems to be closely correlated. The
modification of these structures shows them to be much older in
their development than the pygidium of the Diaspidinae which is
typically perfect only in the adult female. The anal ring and
anal lobes, if present in any stage, are more likely to be found in
the first nymphal stage and, if atrophy or modification of parts
has begun to take place, it starts with the adult and finally extends
into the older nymphal stages. Such changes may be confined to
the adult alone. The typical number of anal ring setae is six in
the first nymphal stage, although there may be more or less in the
other nymphal stages and the adult. A large number of the
species lacking an anal ring and anal ring setae in the adult have
six anal ring setae in the first nymphal stage. The modification
of these structures may extend into the first nymphal stage which
is shown in the reduction in the size of the anal ring setae and
later in the number of these setae to four, two, or none, and going
hand in hand with this change is a segmentation of the anal ring
and finally the complete loss of the anal ring itself. There is
generally a change in the form and a reduction in the size of the
anal lobes associated with the modification of the anal ring, but
the anal lobes like the anal ring reach their most perfect develop-
ment in the first nymphal stage. Some of the subfamilies, as the
Diaspidinae, which have evidently been derived from the line with
anal lobes, an anal ring, and anal ring setae, in the first nymphal
stage of many species have the anal lobes either obscure or so they
can be traced, but are always provided with two large prominent
anal setae. The position and form of these setae show unques-
tionably their homology. They are peculiar to the first nymphal
stage.
CHAPTER IV
CLASSIFICATION
The early students of the Coccidae were not particularly-
interested in the describing of new species and new genera, but in
discussing the habits of certain common species. This was par-
ticularly true of the cochineal insects which were introduced into
Spain, Italy, and northern Africa after the discovery of America.
Upon these a large number of treatises were based. Signoret
after deliberating over all the literature of the family published
before 1868, gives a list of the described species. This list contains
the names of about two hundred and twenty-eight species. Some
of these names have been referred to the synonomy and many others
were applied to species that were so poorly described that they
have not been identified since. The species were included in
thirty-five genera. Some idea of the amount of work and study
that has been given to this family in recent years can be obtained
from a comparison of the number of species and genera listed by
Signoret and those included in the catalogue of Mrs. Fernald
published in 1903, thirty-five years later. This list contains 1514
species and 164 genera.
The subdivision of the family into sections, tribes, or sub-
families varies considerable with the author. The variation
consists not only in the rank assigned the various groups and the
distribution of the groups, but in their number and the genera
apportioned to them.
The first subdivision of the family worthy of consideration
was that of Targioni Tozzetti, '68, where four groups are recog-
nized: Orthezites, Coccites, Lecanites, and Diaspites. Signoret
prepared the first comprehensive account of the family, the
publication of which was begun in 1868, consists of seventeen parts,
and was completed in 1875. In a portion of this work published
in 1868 four sections are recognized: Diaspides, Brachyscelides.
Lecanides, and Coccides. Targioni Tozzetti in 1869 added another
tribe to those which he had previously proposed, Lecanodiaspites.
placing it between his Lecanites and Diaspites. Comstoek, '81,
adopted the sections proposed by Signoret, arranged them in the
40
CLASSIFICATION 41
same order, but ranked them as subfamilies. The new tribe of
Targioni Tozzetti was not recognized and the species that had been
referred to this tribe were considered as belonging to one of the
other sections or subfamilies. The Orthezites of Targioni Tozzetti
were included among the Coccinae of Signoret and Comstock.
W. M. Maskell between the years 1879 and 1897 published
many papers dealing with the Coccidae. In **An account of New
Zealand scale insects" this author divides the family into four
groups, Diaspidinae, Lecanidinae, Hemicoccidinae, and Coeeidinae.
The first two of these groups are the same as the Diaspinae and
Lecaninae of Comstock and the second and third represent the
Coccinae of this author and Signoret. The Brachyscelinae of
Signoret, the Apiomorphinae of this work, which is peculiar to
Australia, was omitted. The group Lecanidinae was subdivided
by Maskell into three groups : Lecanodiaspidae which is the same
as the Lecanodiaspidites of Targioni Tozzetti, Lecanidae, and
Lecano-coccidae ; the Hemicoccidinae into two groups: Kermetidae
and Cryptokermetidae ; the Coeeidinae into five groups: Acantho-
coccidae, Dactylopidae, Coccidae, Monophlebidae, and Porphyro-
phoridae. These names although terminating with the usual
suffixes applied to subfamilies and families are specifically desig-
nated as groups and subdivisions. If all the groups and
subdivisions of this classification are taken into account, this would
make eleven divisions of the family Coccidae. In a list of New
Zealand species published by the same author in 1895, a similar
arrangement is followed. There are six subfamilies recognized :
Diaspidinae, Lecaninae, Hemicoccinae, Coccinae, Monophlebinae.
and Brachyscelinae. The Lecaninae is subdivided into three
sections as in the first classification; the Cryptokermetidae of the
Hemicoccinae is omitted; the Monophlebinae of the Coccinae is
raised to the rank of a subfamily and the Porphyrophoridae the
Margarodinae of later writers, which does not occur in the
Australian region, is omitted ; and the Brachyscelinae omitted from
the previous list is recognized as a subfamily.
Cockerell, '96, in a *' Check-list of the Coccidae" recognized
ten subfamilies arranged as follows in an ascending order: Mono-
phlebinae, Porphyrophorinae, Coccinae, Hemicoccinae, Ortheziinae
Asterolecaniinae, Brachyscelinae, Idiococcinae, Lecaniinae and
Diaspinae. The very different order in which the subfamilies are
arranged, also the appearance of the Orthezites of Targioni Tozzetti
again but as a subfamily and the substitution of the name
42 THE COCCIDAE
Asterolecaniinae for the Lecanodiaspidites of Targioni Tozzetti and
the Lecanococcidae of Maskell, is worthy of note.
Berlese and Leonard!, '98, in their ''Notizie intorno alle
cocciniglie americane che minaciano la frutticultura europea"
recognize eight groups, tribes as they designate them, but with
subfamily suffixes, which are arranged in the following order:
Monophlebinae, Porphyrophorinae, Coccinae, Hemicoccinae,
Ortheziinae, Asterolecaniinae, Lecaninae, and Diaspinae. In a
table for the separation of the subfamilies two additional tribes,
Brachyscelinae placed between the Lecaninae and Coccinae and
Idiococcinae placed before the Monophlebinae, are recognized.
This arrangement does not differ from that of Cockerell. The five
subgroups of their tribe Diaspinae which are recognized by
Leonardi in his ''Generi e specie di Diaspiti," the first part of
which was printed early in 1898, are adopted in the ''Notizie"
and arranged in the same order: Parlatoriae, Aspidioti, Leucas-
pides, Diaspides, and Mytilaspides.
Green, '96, in the first volume of his "Coccidae of Ceylon"
proposes eleven subfamilies arranged as follows in an ascending
order: Conchaspinae, Diaspinae, Lecaniinae, Hemicoccinae,
Dactylopiinae, Tachardiinae, Coccinae, Idiococcinae, Brachy-
sceliinae, Ortheziinae, and Monophlebinae. In remarking upon
the classification of. Cockerell, he states, **I should be inclined to
include the Porphyrophorinae with the Monophlebinae .... nor
does the separation of the Asterolecaniinae from the Coccinae (as
generally constituted) seem necessary-" In volume four of this
work, published in 1909, Green, however, adopts the subfamily
Asterolecaniinae and places it between the Dactylopiinae and
Tachardiinae. This is the first recognition of a subfamily
Tachardiinae which includes the genus Tachardia, the lac insects.
The name of Conchaspis also appears in this list of subfamilies for
the first time, erected to include the genus Conchaspis described in
1893. It is here associated by Green with his subfamily Diaspinae.
In a series of tables for the identification of genera, Cockerell
in 1899 shows eight subfamilies arranged in the following order:
Ortheziinae, Monophlebinae, Margarodinae, Conchaspinae, Coccinae,
Tachardiinae, Lecaniinae, and (Diaspinae). The name Mar-
garodinae is substituted for Porphyrophorinae and the subfamily
divided into two tribes, Margarodini and Xylococcini. The sub-
family Coccinae includes five tribes : Brachyscelini, Asterolecaniini,
Kermesiini, Eriococcini, and Dactylopiini. It will be noted that
CLASSIFICATION
43
the arrangement of the subfamilies is quite different from that
proposed in 1896. The subfamily Conchaspinae, which did not
appear in the previous list, is placed between the Margarodinae
and the Coccinae. Four of the subfamilies of the previous list -are
distributed among the five tribes of the subfamily Coccinae: (1)
Hemicoccinae is changed to Kermesini, (2) Asterolecaniinae to
Asterolecanini, (3) Brachyscelinae to Brachyscelini, and (4)
Idiococcinae is included with the Eriococcini. A new tribe.
Dactylopini, is added to this subfamily. The subfamily name
Tachardiinae, to include the lac insects and first used by Green,
is adopted here. The arrangement of both of the classifications of
Cockerell are evidently intended to show a disposition in an
ascending order instead of a descending order as was evidently
intended by Signoret, Comstock, and Maskell.
Newstead, '01, in his ''Coccidae of the British Isles'' has
''with slight alterations, adopted the very clear and comprehensive
divisions recently given by Mr. E. E. Green, adding thereto the
subdivision Margarodince, which, from the absence of mouth-parts
in the adult female, I agree with Professor Cockerell naturally
calls for a separate division. I should also add that Mr. Green
places the Conchaspince before the Diaspincu; but it seems to me,
although I may be quite wrong, that the insects comprising this
subfamily are more naturally placed in the ascending order above
the BiaspincB, and I have therefore made this additional deviation
from Mr. Green 's classification. ' '
The first complete ''Catalogue of the Coccidae of the World"
was published in 1903 by Mrs. M. E. Fernald. There are nine
subfamilies recognized in this work arranged in the following order :
Monophlebinae, Margarodinae, Ortheziinae, Phenacoleachiinae.
Conchaspinae, Dactylopiinae, Tachardiinae, Coccinae, and Dias-
pinae. The arrangement of the subfamilies and genera is in an
ascending order. The first three subfamilies of this list are similar
in extent to the same group of Cockerell, Green, and Newstead.
The fourth subfamily, Phenacoleachiinae, erected by Cockerell, is
for a peculiar species and genus described from New Zealand by
Maskell. The position of the Conchaspinae is the anomalous one
of the classification of Cockerell, near the Dactylopiinae. The
Dactylopiinae includes the Brachyscelinae, Asterolecaniinae,
Kermesinae, Eriococcinae, Idiococcinae, and Dactylopinae of other
authors. The type of each genus is indicated. The genus Coccus,
which had been used by all previous writers for the cochineal
44 THE COCCIDAE
insects with cacti as type, has a species previously referred to the
genus Lecanium, hesperidum, named as its type. From this change
there has resulted a supression of the generic name Lecanium and
of the subfamily name Lecaniinae derived from it. For these the
generic name Coccus and the subfamily name Coccinae are sub-
stituted. The cochineal insects are referred by Mrs. Fernald to
the genus Dactylopius and to the subfamily Dactylopiinae. I have
examined the evidence submitted but have been unable to discover
any basis for the change and the original application of these generic
names, Coccus and Lecanium, has been restored in the following
pages. The Tachardiinae and Diaspinae are of the same extent as
these groups of Green and Cockerell. A reduction in the number of
subfamilies has resulted in the combination of unrelated groups and
marked a backward rather than a forward step in the classification
of the family. This is more than offset by the way in which this
catalogue has made the literature of the family available to the vari-
ous students of the group. The United States Bureau of Entomology
has issued five supplements, but these include only the names of
new genera and species, the place where they were described, the
localities, and the host-plants. The same arrangement of sub-
families is followed. The classification adopted by Mrs. Fernald
was evidently under the sponsorship of Cockerell as the following
from the introduction implies. ''It gives me much pleasure to
acknowledge numerous favors and great assistance received in the
preparation of this work Especially am I indebted to Prof.
T. D. A. Cockerell, who for the past two years has been constantly
sending me separata and references to foreign literature, besides
giving me all possible assistance in regard to classification and
synonomy, the result of his extensive studies on the Coccidae from
all parts of the world. To him is due much of whatever value
there may be in this catalogue. ' '
A linear arrangement such as must be used on a printed page
is not feasible for showing the phylogeny of a group of animals
or plants. This should be kept in mind when examining and
judging all of the preceeding classifications. The family Coccidae
has been divided into seventeen subfamilies on the following pages.
These have been arranged in an ascending order. The following
arrangement of the dichotomous groups is an attempt to show
graphically by means of a table something as to the lines of
modification of these groups and is not a statement of characters
to be used for the identification of specimens. The synopsis shows
CLASSIFICATION 45
what I believe to be the relation of these subfamilies and the
scientific and vernacular names that have been applied to them.
SYNOPSIS OF THE COCCIDAE
GENERALIZED COCCIDAE.— Anal Lobes and Anal Ring Undeveloped.
Males with Compound Eyes.
Rostralis Conservers. 1. The Giant Coccids. — Monophlebinae.
Rostralis Loosers.
Abdominal Spiracle Conservers.
Prothoracic Legs Normal in Form.
Spiracles closed, Spiracular Tracheae without Cerores
2. The Sinoran Coccids. — Kuwaniinae.
Spiracles Open and Spiracular Tracheae with Cerores.
3. The Meltratuban Coccids. — Xylococcinae.
Prothoracis Legs Fossorial
4. The Ground Pearls. — Mabgabodinae.
Abdominal Spiracles Loosers
5. The Marsupial Coccids. — Callapappinae.
Males with Simple Eyes. 6. The Cochineal Coccids. — Coccinae.
SPECIALIZED COCCIDS— Anal Lobes and Anal Ring Developed.
Abdomen of Anal Ring Type.
Anal Lobe Lackers.
Males with Compound Eyes.
7. The Ensign Coccids. — Ortheziinae.
Males with Simple Eyes.
8. The Ringeyed Coccids. — Phenacoleachitnae.
Anal Lobe Possessors.
Anal Ring Conservers.
Thorax typically with Cerari.
9. The Mealy Bugs. — Eeiococcinae.
Thorax with Canellae.
Abdomen with Rectal and Stigmatal Processes.
10. The Lac Insects. — Tachardiinae.
Abdomen with an Anal Cleft.
11. The Tortoise Scales. — Lecaniinae.
Anal Ring Loosers.
Nymphs Anal Ring Conservers.
Abdomen short, subequal to Head and Thorax.
Adult Female with Cerores, Body Naked
12. The Pseudogall Coccids. — Kermesiinae,
Adult Female with Octacerores, Body Covered.
13. The Ceravitreus Coccids. — Asterolecaniinae.
Abdomen longer than Head and Thorax, Top-shaped.
14. The Pegtop Coccids. — Apiomorphinae.
Nymphs Anal Ring Loosers.
15. The Lubberly Coccids. — Cylindrococcinae.
Abdomen of Pygidial Type.
Abdomen with Cerores.
16. The Exuvlaless Coccids. — Conchaspinae.
Abdomen with Ceratubae. 17. The Armored Scales. — Diaspidinae.
46 THE COCCIDAE
The coccids in common with all other groups of animals are
divisible into two series, a generalized series and a specialized
series. These two series are differentiated so far as the Coccidae
are concerned by the lack or development of anal lobes and an
anal ring, three structures that are closely correlated. The presence
of either anal lobes or of an anal ring in one or all the stages of
any given species of all but the generalized end of the specialized
series although the other, anal lobes or anal ring, is wanting, is
considered as indicative of the presence of both anal lobes and an
anal ring in the progenitors of the species.
The generalized series. The Generalized Coccids, includes those
subfamilies in which an anal ring and anal lobes are not only
wanting but the members of this series were separated off from
the progenitors of the coccids before the origin of these structures.
The subfamilies of this series, six in number, represent very
different types. The long many segmented antennae, the many
segmented body, the retention of legs, the two or three segmented
rostrum, the retention of ten pairs of spiracles in certain genera,
the presence of furcae and apodemae and episterna, and the
retention of compound eyes and usually ocelli in the adult male,
all mark the Monophlebinae as the most generalized member not
only of this series but of the family Coccidae. The four following
subfamilies all agree in that the adult females lack a rostralis and
probably a rostrum and that the adult males have compound eyes,
usually also with ocelli. Each subfamily bears the brand of a
different line of development, the Kuwaniinae contains the most
generalized species, those which are only slightly different from the
Monophlebinae, many of them apparently differ only in the loss
of their rostralis. The Xylococcinae, Margarodinae, and Callapap-
pinae have each gone along a tangent of its own. In the first the
nymphs of the species live in depressions in the solid bark of trees
and there has been developed a structure in the rectum for
excreting a long thread-like anal tube of wax through which the
excretia is passed. The presence of a similar structure in the
nymphs of certain Monophlebinae is of interest. In the second
subfamily, the loss of legs during certain nymphal stages and the
enclosure of the body in a pearly cyst is distinctive. The legs are
regained at the last molt. The presence of greatty enlarged
prothoracic legs which are fitted for digging, since all the species
live upon the roots of plants, in both males and females is not
only characteristic but distinctive. In the third subfamily the
CLASSIFICATION 47
large size of the individuals and the invagination of the abdomen
in the adult female to form a pouch for carrying the eggs so that
there are only two or at most three abdominal segments exposed
is characteristic. The presence of the anus and vulva ~at- the
cephalic end of the pouch and the apparent absence of most of the
abdominal spiracles is to be expected.
The subfamily Coccinae includes the most specialized species
of the Generalized Coccids. They differ from the four subfamilies
just described in the retention of the rostralis. In this feature
they agree with the Monophlebinae, from which they are not readily
differentiated. The most striking features are the few segmented
antennae, the absence of abdominal spiracles, the presence of short
blunt truncated setae, and the presence of ocellanae in the adult
male instead of compound eyes. This last feature separates them
from the five other subfamilies. The absence of anal lobes and an
anal ring not only in all nymphal stages but in the adult female
together with the characteristic form of the abdomen precludes the
association of the subfamily with the Specialized Coccids. The
presence of ocellanae in the adult male and the absence of
abdominal spiracles in all stages of the female would necessitate
the association of the group, if placed with the Specialized Coccids,
with the most specialized members of this series. The structure
of the species of the Coccinae makes this impossible.
As is common with other groups, the second series, the
Specialized Coccids, includes not only the largest groups, but those
showing the greatest departure from the form and structure found
in the Monophlebinae. This series is divisible into two marked
types, the Anal Ring Type and the Pygidial Type. The anal ring
is evidently older phylogenetically than the anal lobes, because in
the two subfamilies, Ortheziinae and Phenacoleachiinae, which are
recognized as the most generalized of the series, a well devloped
anal ring is present in both but there is no indication of anal lobes
in any stage of any of the species of either group. The retention
of compound eyes with ocelli, three at least in one species, in the
adult male and of several pairs of abdominal spiracles in the
females of many species is strong evidence in corroberation of the
consideration of the species of this subfamily as the most generalized
of the Specialized Coccids. This is further substantiated by the
long many segmented antennae, the distinctly segmented abdomen,
and the retention of legs in all stages. The cerores of the Orthe-
ziinae, which are characteristic in form, differentiate them, so far
CLASSIFICATION 49
The body in the adult female of all the species of Tachardiinae
is a bag with the mouth-parts at one end and three tubes at the
other. One of the tubes, the mesal one, bears the anus and anal
ring and anal ring setae. The mesothoracic spiracles are associ-
ated with the other two. These- spiracles reach their present
position through a revamping of the body. The first stage nymphs
are similar in a general way to those of the Eriococcinae, although
the anal lobes and the anal setae are wanting, showing that the
usual form of the body of the adult female is derived from a
progenitor of a type common to that of the Eriococcinae. The lack
of cerores in the adult female, although a considerable amount of
wax is excreted, the loss of legs, the presence of canellae with
cerores, and the loss of antennae mark it as a distinct line of
development. The segmentation is distinct in the first nymphal
stage, obscure in the others, and wanting in the adult female.
The first nymphal stage in the Lecaniinae, while different when
examined in detail, has many of the characteristic features of the
first nymphal females of the Eriococcinae. The presence of
canellae marks a relationship with the Tachardiinae. The segmen-
tation is generally obscure in the first nymphal stage, but legs and
antennae, while wanting in some of the highly specialized genera,
are usually present. The presence of the characteristic anal cleft
with an operculum on each side near the caudal end of the anal
cleft in the first nymphal stage and at its cephalic end in the adult
female is characteristic. The development of an eversible tubular
extension caudad of the anus and anal ring from an invagination of
the body-wall is also distinctive. The three subfamilies of the
Anal Ring Conservers have features showing their origin from a
common progenitor, but many other structures mark the develop-
ment of each along a line peculiarly its own.
Each of the four subfamilies of the Anal Ring Loosers, like
each of the subfamilies of the Anal Ring Conservers, has developed
along characteristic phylogenetic lines. Lines that are so dis-
tinctive that they can not be said to have been derived from any
one of the subfamilies of the Anal Ring Conservers. While each
of these subfamilies retains features showing their relationship
with the generalized species of Eriococcinae, these features are
confined for the most part to the first nymphal stage and must be
interpreted as indicating a common descent with the Eriococcinae
instead of an intimate relationship with or descent from this group.
While the anal ring and anal ring setae are more ancient structures
50 THE COCCIDAE
than the anal lobes and anal setae, as already noted, the latter are
more persistent and are often retained after the complete loss of
the anal ring. The anal lobes and anal ring are retained in the
nymphs of Kermesiinae, Asterolecaniinae, and Apiomorphinae, they
are usually wanting in the adults, while in the Cylindrocoecinae
the anal ring is wanting in all nymphal stages as well as in the
adult, but anal lobes are generally retained in the first nymphal
stage. The naked globular unsegmented body without or with
greatly reduced legs and antennae is characteristic of the
Kermesiinae ; the vitreous or glassy covering of wax excreted from
octacerores is characteristic of the typical Asterolecaniinae; the
greatly elongated top-shaped body with the abdomen typically
several times the length of the head and thorax together, fitting
the insect for a life in an open gall, is characteristic of the
Apiomorphinae; and the peculiar circular mite-like bodies with
the legs wanting or one pair much longer and larger than the
others is characteristic of the Cylindrocoecinae.
In the two subfamilies with the pygidial type of abdomen,
all trace of the anal ring and anal ring setae are typically lost in
all nymphal stages and the adult. Brittin has recently recorded
the presence of an anal ring with anal ring setae in Scutare, a new
genus of Conchaspinae from New Zealand. They differ from all
other coccids in having the body covered by a loose thin disk-like
scale of wax formed by the weaving together of threads of wax.
There are two subfamilies included, the Conchaspinae in which the
scale is formed from wax excreted from cerores. The legs and
antennae are present in all stages and the first abdominal segment
bears a pair of spiracles. The abdomen is depressed and the
general form of the body is that of the next subfamily, but there
is no fusion of segments to form a pygidium like that of the
Diaspidinae. In the second subfamily, the Diaspidinae, the scale
is formed from wax excreted from ceratubae. The antennae and
legs are present in the first nymphal stage but are wanting in the
second nymphal stage and the adult female. The abdomen never
bears spiracles but is always depressed and the caudal segments
are fused into a prominent pygidium which typically bears
numerous microscopic projections of great taxonomic value. The
scale differs from that of the Conchaspinae in that there is mixed
with the wax of the scale the cast skins or exuviae of the nymph.
The following tables are for the separation of the specimens
of the subfamilies listed in the above synopsis. They are artificial in
CLASSIFICATION 51
arrangement and are not intended to indicate anything as to the
relationship of the groups concerned. The tables are for three
different ages of each species, first, for those of the first nymphal
stage which would include males and females since the sexes are
indistinguishable in this stage; second, for those of the other
nymphal stages of the female, one, two or more as the case may be ;
and third, for those of the adult female. There has been included
in this last table in certain of the subheads such nymphal struc-
tures as would make the table more usable, definite, and exclusive.
I am aware that the two tables for the separation of nymphs are
very inadequate and experience will undoubtedly show that they
are defective. They are offered even with their imperfections in
hopes that they may at times prove of value and also that the various
students of the Coccidae may be led to record observations that
will make possible the removal of the imperfections, so that bettei
tables for the separation of the nymphal stages can be constructed.
Experience has shown that in many if not all of the subfamilies,
there are characters present in the nymphal stages that are not
only more easily observed but more fixed than many of the char-
acters found in the adult females.
Tables for the separation of the subfamilies as well as all the
later tables follow an exact dichotomous arrangement. While
tables of this sort require more space because of the indentation
of the paragraphs, this is more than compensated for by the ease
with which the rank of the paragraphs can be recognized through
the difference in the indentation. The difference in rank is further
shown by the numbering of the paragraphs with the letters of the
alphabet, the sequence of the letters showing the relative rank of
the paragraph. In using the tables the statement of characters
given under paragraph a and aa should be read and carefully
compared. The specimen should then be examined and compared
with the statement of characters given under a and aa and the
question decided as to which of these statements describe the
characters found in the individual examined. If the specimen
agrees with the characterization given under a, the other paragraphs
given under a should be examined and all those under aa should
be disregarded. Read the characterizations given under b and bb
and determine with which of these the specimen agrees. If the
specimen agrees with b, disregard the paragraphs under bb and then
determine in the same way whether it agrees with c or cc, d or dd,
e or ee, etc., or until the name of a subfamily has been reached.
52 THE COCCIDAE
These names are printed at the right-hand end of the last line of
certain paragraphs. If the specimen should agree with bb instead of
b, disregard the paragraphs under b and then determine in the same
way with which of the paragraphs under bb the specimen should
agree until the name of a subfamily is reached. If the specimen
should belong under aa instead of a, then all of the paragraphs
under a should be disregarded and the specimen compared with the
characterizations given under b and bb, c and cc, d and dd, etc.
of aa until its correct position has been determined.
SUBFAMILIES OF COCCIDAE
FIRST NYMPHAL STAGE
a. Abdomen, even if consisting of only six segments, never with
caudal end chitinized, depressed, and forming a pygidium bearing
two large anal setae,
b. Abdomen either with or without spiracles, but if with spiracles,
never with only a single pair located on the first segment; tibio-
tarsal sutures rarely obsolete, if so, body always with pilacerores.
c. Abdomen never longitudinally cleft at caudal end, forming
anal cleft with operculum on each side of cleft, if apparently
cleft, body always provided with octacerores.
d. Abdomen always without anal ring and anal ring setae.
e. Abdomen with caudal and lateral margins fringed with
chalazae; antennae distinctly clavate Monophlebinae.
ee. Abdomen with caudal and lateral margins never fringed
with chalazae; antennae rarely clavate, usually not en-
larged, but setaceous, or at most only slightly clavate.
f. Prothoracic legs greatly enlarged with greatly swollen
femora and fitted for digging; first pair of abdominal
spiracles dorsal in position, other abdominal spiracles
ventral in position. Maegarodinae.
ff. Prothoracic legs never greatly enlarged with greatly
swollen femora and fitted for digging; abdominal
spiracles, when present, always with those of first seg-
ment on same aspect as those of other segments.
g. Abdomen never provided with anal lobes and anal
setae.
h. Abdomen always provided with two or more pairs
of spiracles, some of them sometimes small and
inconspicuous; cuticle never provided with num-
erous short truncate setae.
1. Abdomen, if with spiracles, always with more
than three pairs; rostrum consisting of two
segments.
j. Thorax with two pairs of spiracles which are
large and normal in size; ventro-meson of
CLASSIFICATION 53
" caudal portion of abdomen not with a row
of large cerores; caudal abdominal segment
not projecting farther caudad than lateral
portions of penultimate segment; coriae be-
tween segments convex. Kuwaniinae.
jj. Thorax with two pairs of spiracles which are
minute and often inconspicuous; ventro-
meson of the caudal portion of abdomen with
a longitudinal row of large cerores; caudal
abdominal segment projecting farther caudad
than the lateral portions of penultimate seg-
ment; coria between segments straight
Xylococcinae.
ii. Abdomen never with more than three pairs of
spiracles located on segments one to three;
rostrum consisting of single segment
Callipappinae.
hh. Abdomen without spiracles; cuticle provided with
numerous short truncate setae arranged in longi-
tudinal rows; rostrum consisting of two incon-
spicuous segments. Coccinae.
gg. Abdomen provided with anal lobes and small anal
setae, often obscure; dorsal aspect of b.ody often with
prominent longitudinal or transverse rows of setae;
rostrum consisting of two segments.
Cylindrococcinae.
dd. Abdomen always with anal ring with anal ring setae.
e. Body always bearing groups of pilacerores excreting
lamellae of hard white wax which are usually more or
less fused; tibio-tarsal sutures sometimes obsolete; abdo-
men frequently with seven pairs of spiracles; rostrum
consisting of two segments. Ortheziinae.
ee. Body never bearing groups of pilacerores; abdomen, so
far as known, never provided with spiracles,
f. Rostrum consisting of three segments; abdomen not
provided with anal lobes or anal setae; antennae con-
sisting of seven segments; tarsus with toothed claw.
Phenacoleachtinae.
ff. Rostrum usually consisting of two segments, sometimes
of only single segment, never of three segments; abdo-
men provided with anal lobes and anal setae, some-
times obscure; antennae usually consisting of six seg-
ments, sometimes with less.
g. Thorax always provided with prominent canellae
with cerores extending from mesospiracles, meta-
spiracles without canellae; anal lobes and anal setae
wanting; anal ring located at caudal end of body
on protuberance, each of six anal ring setae attached
to chitinous plate. Tachardiinae.
54 THE COCCIDAE
gg. Thorax never provided with canellae; anal lobes
present, sometimes obscure; anal ring more or less
distant from caudal end of body and each anal ring
seta never attached to chitinized plate,
h. Body never provided with octacerores, these gener-
ally arranged in longitudinal transverse or margi-
nal rows; rostrum consisting of a single segment,
hh. Body never provided with octacerores; rostrum,
so far as known, always consisting of two seg-
ments.
i. Body strongly depressed, elliptical or oval, and
its periphery fringed with flattened suboval
; ' scales; anal lobes obscure or wanting, anal
setae large and distinct. Apiomorphinae.
ii. Body rarely depressed, elliptical or oval in
outline, usually elongate, and its periphery
never fringed with flattened suboval scales;
anal lobes and anal setae usually distinct,
j. Body almost naked or sparsely setaceous,
with setae spine-like and arranged in longi-
tudinal or marginal rows, body often with
cerarae; cerores abundant, so that body is
usually covered with mealy or powdered wax;
ocellana present on each side of head; anal
ring complete, rarely divided, length of anal
ring setae usually several times width of
anal ring. Eriococcinae.
jj. Body usually provided with prominent spine-
like setae, generally arranged in marginal
rows, sometimes in longitudinal rows, never
provided with cerarae; cerores never abund-
ant, always a few in number and body at
most only rarely covered with mealy or
powdery wax; ocellanae wanting; anal ring
often subdivided or obscure, anal ring setae
generally short, at most not much if any
longer than width of anal ring._KEEMESiiNAE.
cc. Abdomen always longitudinally cleft at caudal end and with
an operculum on each side near the caudal end of cleft; body
not or only rarely provided with octacerores; rostrum con-
sisting of single segment; anal ring situated at cephalic end
of lumen of an anal tube and concealed externally unless
exserted. Lecaniinae.
bb. Abdomen always with single pair of spiracles, always located on
first segment; tibio-tarsal sutures always wanting; body never
with pilacerores; rostrum consisting of two segments; body with
cerores, few in number, never with ceratubae. Conchaspinae.
aa. Abdomen consisting apparently of six segments, caudal segment
depressed and strongly chitinized, forming a pygidium, bearing two
long anal setae, decussating and folded over abdomen; rostrum
CLASSIFICATION 55
consisting of single segment; body with ceratubae, never with
cerores; legs and antennae always present, latter consisting of
six segments. Diaspidinae.
FEMALE NYMPHS
Abdomen never with caudal end strongly chitinized and flattened,
forming pygidium, even if flattened, not with pectinae, plates, or
ceratubae; legs present or wanting, if wanting, body never covered
by leaf-like scale with exuviae.
b. Legs present or wanting, if present, never with body covered by
leaf -like scale without exuviae; tibio-tarsal sutures rarely obso-
lete, if so, body with numerous pilacerores; abdominal spiracles,
if present, not limited to flrst abdominal segment.
c. Abdomen not longitudinally cleft at caudal end with operculum
on each side of cleft, if apparently cleft, body provided with
octacerores.
d. Abdomen always without anal ring and anal ring setae,
e. Legs and antennae wanting,
f. Abdomen with first pair of spiracles dorsal in position,
others ventral; body of insect usually encased in irreg-
ular cyst of over-lapping plates, somewhat pine-cone-
like; insects living in ground; rectum never chitinized
and provided with rings of anacerores excreting long
thread-like tube of wax; abdominal spiracular tracheae
never with cerores. Maegarodinae.
ff. Abdomen with first pair of spiracles, when present, on
same aspect as those of other segments; body of insect
never encased in irregular cyst of overlapping plates;
insects arboreal in their habits; rectum strongly chit-
inized and provided with rings of anacerores excreting
long thread-like tube of wax; abdominal spiracular
tracheae with cerores. Xylococcinae.
ee. Legs and antennae present; rectum never with rings of
anacerores, spiracular tracheae never with cerores, or
body never encased in irregular cyst of over-lapping
plates,
f. Abdomen never with any indication of anal lobes and
never with anal setae; legs usually present and normal
in form.
g. Cuticle of body never bearing numerous short trun-
cate setae; abdomen frequently with spiracles,
h. Abdomen either without spiracles or if with
spiracles, usually with more than three pairs, if
with three pairs, located on segments six to eight,
i. Abdomen usually with lateral and caudal mar-
gins fringed with band of setae, more numerous
and longer than those of dorsal or ventral
aspects, among which are located large cerores;
rostrum consisting of two or three segments.
MONOPHLEBINAE.
56 THE COCCIDAE
ii. Abdomen, so far as known, not with lateral
and caudal margins fringed with setae more
numerous or longer than those of dorsal or
ventral aspects; rostrum consisting of two seg-
ments. KUWANIINAE.
hh. Abdomen never with more than three pairs of
spiracles and these always located on segments
one to three; rostrum, if present, consisting of a
single segment Callipappinae.
gg. Cuticle of body bearing numerous short truncate
setae; abdomen always without spiracles; rostrum
never wanting, consisting of two inconspicuous seg-
ments. COCCINAE.
ff. Abdomen with anal lobes and small anal setae, anal
lobes often inconspicuous, rostrum always present and
consisting of two segments. Cylindrococcinae.
dd. Abdomen always with anal ring and anal ring setae.
e. Body bearing groups of pilacerores excreting lamellae of
white wax which are more or less fused; tibio-tarsal
suture sometimes obsolete; abdomen frequently with
seven pairs of spiracles Ortheziinae.
ee. Body never bearing groups of pilacerores; tibio-tarsal
suture present; abdomen, so far as known, never provided
with spiracles.
f. Rostrum consisting of three segments; abdomen not
with anal lobes or anal setae; tarsus with toothed
claw. Phenacoleachiinae.
ff. Rostrum usually consisting of two segments, sometimes
of only single segment, never of three segments; abdo-
men usually provided with anal lobes and anal setae,
anal lobes sometimes obscure or wanting.
g. Thorax provided with canellae associated with meso-
spiracles, extending onto dorsal aspect, mesospiracles
twice as far apart measured on sternum as metaspir-
acles; anal ring setae ten in number and placed on
six circular or oval plates; anal lobes and anal setae
wanting. Tachardiinae.
gg. Thorax never provided with canellae; mesospiracles
at most only slightly farther apart on ventral aspect
than metaspiracles; anal ring setae never placed on
plates, usually six in number, sometimes more or
less.
h. Body provided with octacerores, generally arranged
in longitudinal, transverse, or marginal rows;
rostrum consisting of single segment
ASTEROLECANIINAE.
hh. Body never provided with octacerores; rostrum, so
far as known, always consisting of two segments.
CLASSIFICATION ' 57
i. Abdomen elongate, much longer than head and
thorax together, lateral margins strongly con-
vergent caudad; anal lobes strongly chitinized,
divergent, pointed, and prominent; metathor-
acic coxae twice as far apart as prothoracic
coxae. Apiomorphinae.
ii. Abdomen not decidedly elongate, subequal in
length to head and thorax together; lateral
margins usually not strongly convergent caudad,
usually with anal lobes, but these not pointed,
divergent, strongly chitinized projections;
metathoracic coxae at most only slightly farther
apart than prothoracic coxae.
j. Body bearing numerous cerores excreting
powdery or mealy wax, forming covering for
the body, cerores often associated with setae
and forming cerarae; anal ring usually bear-
ing anal ring setae, setae sometimes small,
reduced in number, less than six or wanting,
usually similar to or more than in adult
Eriococcinae.
jj. Body practically without or with only few
cerores; mealy or powdery wax, if present,
limited to the ventral aspect; only first
nymphal stage and adult female have been
described. Kermesiinae.
cc. Abdomen always longitudinally cleft at caudal end and with an
operculum on each side of cleft; cuticle not or only rarely pro-
vided with octacerores; rostrum consisting of single segment;
anal ring situated at cephalic end of lumen of anal tube and
concealed externally unless exserted. Lecaniinae.
bb. Legs always present and body always covered with leaf-like
scale without exuviae; tibio-tarsal suture always obsolete; body
with only few cerores, never with pilacerores; abdominal spir-
acles limited to single pair on first abdominal segment
CONCHASPINAE.
aa. Abdomen with caudal end always strongly chitinized and fiattened,
forming pygidium fringed with pectinae or plates and bearing
ceratubae; legs and antennae always wanting and body always
covered by leaf-like scale with exuviae Diaspidinae.
ADULT FEMALES
a. Abdomen of adult female with all segments separate and flexible
or all segments fused and inflexible, but never divided into two
regions, cephalic series of four flexible movable segments forming
preabdomen and caudal series of fused inflexible segments forming
pygidium bearing plates or pectinae; dorsal aspect of abdomen with
or without cerores but never with ceratubae.
b. Legs present or wanting, if present, never with body covered
S$ THU COCCIDAE
with leaf -like scale without exuviae; tibio-tarsal sutures rarely
obsolete, if so, body bearing pilacerores.
c. Abdomen of adult female never longitudinally cleft at caudal
end with pair of opercula at cephalic end of cleft, if apparently
cleft and provided with opercula, cuticle usually bearing
octacerores in some of nymphal stages and usually also in
adult female, frequently provided with cribiform plates if
provided with octacerores.
d. Body of adult female never with mouth-parts at one end and
.three tubular projeptions with associated spine-like projec-
tion; never enclosed in resinous cell with three adjacent
openings; mesothoracic spiracles never associated with stig-
matic processes located near anus.
e. Adult female and female nymphs rarely provided with
octacerores, if so provided, also provided with ventral
cribiform plates; never with dorsal cribiform plates;
rostrum usually consisting of two or three segments, if
of only one segment, abdomen invaginated to form a
marsupium.
f. Body of adult female never flask-shaped or peg-top-
shaped; abdomen with sides uniformly convex, not
strongly convergent caudad, as broad as or broader than
thorax at cephalic end and usually as long as or longer
than thorax.
g. Adult female and female nymphs always without
anal ring and without anal ring setae; body of adult
female never naked and globular or gall-like in form.
h. Mouth-parts always present in adult female; adult
male with compound eyes or ocellanae.
i. Adult female with all legs present and subequal
in length,
j. Antennae of adult female consisting of eight
to eleven segments, rarely only seven, artic-
ulated to ventral aspect near lateral margin
of head; cuticle never bearing short truncate
setae; rostrum consisting of two or three
segments; adult male with compound eyes.
MONOPHLEBINAE.
jj. Antennae of adult female consisting of seven
short segments, articulated to ventral aspect
as far from lateral margin as from each
other; cuticle usually bearing short truncate
setae; rostrum consisting of two inconspic-
uous segments; adult male with ocellanae.
COCCINAE.
i. Adult female either with all legs wanting or
with all but one pair wanting, if all legs are
present, one pair is two or three times as
CLASSIFICATION 59
long as either of other pairs; antennae, if
present, consisting of less than seven segments,
often vestigial or wanting; adult male with
ocellanae. Cylindrocgooinae.
a. Mouth-parts always wanting in adult female;
adult male with compound eyes.
i. Prothoracic legs of adult female and male not
greatly thickened and fitted for digging, normal
in form; insects arboreal.
j. Abdomen normal in form, all segments fully
exposed, none of them infolded or invaginated
to form marsupial pouch for carrying eggs;
anus and vulva always opening to exterior;
abdominal spiracles, when present, usually
consisting of more than three pairs.
k. Abdomen of adult female usually with less
than eight pairs of spiracles, if with eight
pairs, spiracles never open and their spir-
acular tracheae never provided with one
or more rings of cerores; rectum of nym-
phal females never strongly chitinized and
forming a rectal tube, never bearing ana-
cerores; antennae consisting of eight to
ten segments. Kuwaniinae.
kk. Abdomen of adult female with six to
eight pairs of spiracles, spiracles usually
large and broad and open with spiracular
tracheae with one or more rings of cerores;
rectum of nymphal females strongly chit-
inized, forming tube bearing one or more
rings of anacerores which excrete long
glassy hair-like tube eight to ten times as
long as entire body; antennae consisting of
five to eleven segments. Xylococcinae.
jj. Abdomen abnormal in form, only two or three
cephalic segments exposed, remainder in-
folded or invaginated to form marsupial
pouch for carrying eggs; anus and vulva
never opening to exterior but within and at
cephalic end of marsupial pouch; abdominal
spiracles limited to segments one to three;
antennae consisting of ten to eleven seg-
ments. Callipappinae.
ii. Prothoracic legs of the adult female and male
always thickened and fitted for digging, fossor-
ial; insects root inhabiting; antennae consist-
ing of eleven segments Margarodinae.
gg. Adult female usually with anal ring with anal ring
setae, if without, female nymphs always provided
60 THE COCCIDAE
with anal ring with anal ring setae; body of adult
female sometimes naked and globular or gall-like in
form.
h. Adult female and female nymphs usually with
anal ring bearing anal ring setae, if anal ring
or anal ring setae are wanting in adult female,
body always bearing numerous cerores and com-
pletely covered with white mealy or powdery wax;
body of adult female never naked and globular or
gall-like in form.
i. Abdomen of adult female bearing long mar-
supium formed of fused parallel plates or
lamellae of hard white wax excreted from
pilacerores; antennae consisting of four to nine
segments; rostrum consisting of two segments;
adult males with compound eyes.__ORTHEziiNAE.
11. Abdomen of adult female never bearing mar-
supium formed of fused parallel plates of hard
wax and never provided with pilacerores; male
always with ocellanae.
j Antennae of adult female with eleven seg-
ments; female nymph and adult female with
anal lobes wanting; always with anal ring
with anal ring setae; rostrum consisting of
three segments; aduli male with ocellanae
forming almost a complete ring around head.
Phenacoleachiinae.
jj. Antennae of adult female always with less
than eleven segments, five to nine segments,
sometimes rudimentary or wanting; female
nymphs always with anal lobes, usually dis-
tinct; adult females with anal lobes, some-
times only slightly indicated; adult female
sometimes without an anal ring and anal ring
setae, but first female nymph always with
anal ring and anal ring setae; rostrum con-
sisting of two segments; adult male with
ocellanae few in number and not arranged so
as to form an almost complete ring around
head. Eriococcinae.
hh. Adult female rarely with anal ring and female
nymphs always with anal ring and anal ring
setae; body of adult female gall-like in form and
usually found only on species of oak; cerores few
in number, venter sometimes but entire body
never completely covered with powdery or mealy
wax; body usually grayish or brownish in color,
frequently polished; antennae of adult female
consisting of six segments, rudimentary, or want-
CLASSIFICATION 61
ing; rostrum consisting of two segments
Kermesiinae.
ff. Body of adult female always flask-shaped or peg-top-
shaped; abdomen with sides either strongly conyergent
caudad, narrower and longer than thorax or thorax
globular in outline and abdomen minute plug-like pro-
tuberance; thorax with prothoracic and mesothoracic
legs shorter than metathoracic legs and nearer together;
species gall inhabiting; antennae consisting of three to
five segments or wanting; males with ocellanae
Apiomorphinae.
ee. Female nymph of at least one stage and usually also adult
female provided with octacerores; body often provided
with dorsal cribiform plates but never with ventral crib-
iform plates; antennae consisting of four to six segments
or tuberculate; rostrum consisting of single segment
ASTEROLECANIINAE.
dd. Body of adult female always with mouth-parts at one end
and three tubular projections at other, one projection bearing
anus and other two mesothoracic spiracles, usually with
associated spine-like projection; always enclosed in resinous
cell with three adjacent openings; each mass of wax con-
taining one to many individuals; mesothoracic spiracles
always associated with stigmatic processes located near anus;
legs and antennae wanting; rostrum consisting of two seg-
ments. Tachardiinae.
cc. Abdomen of adult female always longitudinally cleft at caudal
end with pair of opercula at cephalic end of cleft; sides of
cleft rarely fused with loss of line of fusion; cuticle rarely
provided with octacerores, if so, never provided with dorsal
or ventral cribiform plates; antennae consisting typically of
seven to eight segments, sometimes greatly reduced or want-
ing; rostrum consisting of single segment Lecaniinae.
bb. Legs always present and body always covered with leaf-like
scale without exuviae; tibio-tarsal sutures always obsolete; abdo-
men provided with cerores but never with ceratubae; antennae
consisting of three segments; rostrum consisting of two segments.
CONCHASPINAE.
aa. Abdomen of adult female with segments divided into two groups,
cephalic series of four flexible movable segments forming preabdomen
and caudal series of fused inflexible segments forming pygidium;
dorsal aspect of abdomen without cerores and always with ceratubae;
ventral aspect sometimes provided with genacerores; body always
covered with leaf-like scale containing one or two exuviae; legs
wanting, antennae tuberculate; rostrum consisting of single seg-
ment. DiASPIDINAE.
CHAPTER V
SUBFA>nLY MONOPHLEBINAE
The Giant Coccids
The body of the adult female is rotund or strongly convex on the
dorsal and flat on the ventral aspect. The cephalic, the caudal, and the
lateral margins are broadly convex. The segments are generally well
defined. The caudal end of the body is not inflated and the cephalic and
lateral margins are not provided with a keel. The antennae are prom-
inent, usually consist of eight to eleven segments, and are articulated
to the ventral aspect of the head near each lateral margin. The eyes
are present. There is a single ocellana located on each side of the
head near the articulation of an antennae. The mouth-parts are always
present, the rostrum consists of two or three segments, rarely of only a
single segment, and the rostralis is long and slender. The legs are
present. They are subequal in length and normal in form. The profe-
mora are never enlarged and the prothoracic legs are never fltted for
digging. The trochanto-femoral and tibio-tarsal sutures of all the legs
are always distinct. The spiracles of the mesothorax and metathorax
are normal in form and ventral in position. The mesothoracic spiracles
are never located near the anus. The abdominal spiracles are located
on the lateral or dorsal aspect, are variable in number, varying from
eight pairs to none, and are never large in size. The spiracular tracheae
of the abdominal spiracles are not provided with one or more rings of
cerores. The stigmatic clefts, spinae, and canellae are wanting. The
abdomen is about as long as the head and thorax together. It is convex
on the dorsal and ventral aspects, is never depressed. The lateral mar-
gins are uniformly convex. All of the segments are fully exposed and
subsimilar in appearance with coriae between them, none are retracted
to form a marsupium. The abdomen is never provided with an anal
cleft and opercula, an anal ring and anal ring setae, or anal lobes and
anal setae. The caudal abdominal segment is never short and narrow
and extending beyond the other segments. The body is provided with
cerores none of which are octacerores or pilacerores and is never
provided with ceratubae. The body is never enclosed in a resinous cell
with three adjacent openings or covered by a scale with or without
exuviae, but may be more or less covered or associated with white
powdery wax or loose woolly excretion of wax or may rest upon a large
mass of wax with or without a fluted surface and with an internal mass
of doughy wax. The rectum is never provided with a long glassy tube
of wax or with a chitinized tube bearing rings of anacerores. The body
is never naked and gall-like in form.
The female nymphs of all stages are provided with a rostrum of two
or three segments and a long rostralis. The three pairs of legs are
62
SUBFAMILY MONOPHLEBINAE 63
always present in all nymphal stages and are similar in form. The
profemora are never greatly enlarged and the prothoracic legs are never
fitted for digging. The body is never provided with an anal cleft and
opercula, an anal ring and anal ring setae, anal lobes and anal setae,
octacerores, pilacerores, or ceratubae. The caudal end of the rectum
is sometimes modified into a strongly chitinized rectal tube provided with
one or more rings of anacerores, but not excreting a long glassy tube of
wax. The caudal abdominal segment is not short, narrow and projecting.
The male has compound eyes and may have an ocellus along the
caudal margin of each compound eye. The abdomen is either provided
with long lateral filaments or is without such structures, never provided
with tufts of long filaments of wax. The stylus is small or wanting.
The subfamily Monophlebinae contains about fifteen genera
and about ninety species. While certain of these genera are
limited to species from the old world and others to species from the
new world, the larger more important genera contain species from
all parts of the world so that they can be said to be universal in
their distribution. The species are found mainly in tropical
regions but a few extend into or have been introduced by commerce
into subtropical and the warmer temperate regions. A few species
sometimes become established under glass in north temperate
regions.
Coccids are generally considered as being of small or minute
size. This is true of the great majority of the species. The sub-
family Monophlebinae nevertheless contains some species of
considerable size. Species that are twelve millimeters or one-half
inch in length are of common occurence, while some of the species
of certain genera as Llaveia, attain a length of twenty-five
millimeters or one inch. It is not strange, since this subfamily
contains the most generalized coccids, that it should include species
of considerable size, but species that are of considerable size, not
only for coccids, but for insects in general.
The best known species of this group in America is the citrus
pest, the Cottony Cushion Scale or Fluted Scale, Icerya purchasi.
The life history of this species, because of its economic importance,
has been studied more intensely than that of any other. The
following account, based in great part upon the descriptions of this
species, is offered in place of a detailed statement of the serial
development of the group as a whole, since our knowledge is too
limited at this time to prepare such a statement.
The recently hatched nymphs are reddish yellow in color.
The body is ovate in outline with the dorsal surface convex and
the ventral aspect more or less flattened. The antennae are long
64 THE COCCIDAE
and consist of six segments which are enlarged so that the distal
portion of the flagellum is distinctly clavate. This enlargement is
located in the segment which bears several short setae and about
four setae which are longer than the entire antennae. There is a
single ocellana located on each side of the head on a tubercle
which is situated near the articulation of an antenna. The
segmentation of the body is indistinct but more distinct than is
usual with coccids and more distinct than in the later nymphal
stages of this species. The rostralis extends as a loop into the
long deeply invaginated crumena. The head, thorax, and abdomen
bear numerous cerores which are arranged in longitudinal rows,
one along each lateral margin and four others between them.
These cerores form transverse rows, two on the head, three on the
prothorax, two each on the mesothorax and metathorax, and one
on each of the nine abdominal segments. The setae are numerous,
placed among the cerores and like them arranged in transverse
rows. The cuticle bearing each seta is swollen, pimple-like, and
forms a chalaza. The calyx marking the articulation of each seta
to its swelling is distinct. On the antennae, legs, and rostrum the
setae are of the ordinary type. The setae are longer on the lateral
than on the mesal portions of the body, while the six setae at the
caudal end of the abdomen, three on each side of the meson and
known as the anal chalazae, are as long as the entire body. These
setae are always borne by large swellings. The short, smooth,
transverse areas between the cerores and chalazae mark the posi-
tion of the coriae. The legs are normal in form and all their
sclerites are easily identified. Each coxa bears a distinct coxartis
which articulates against a coxacoila located in a curved linear
episternum. The ental surface of this sclerite is produced as a
distinct apodema. The thoracic spiracles are normal in position.
Each contains a prominent spiracularia extending into the body
cavity from the cephalic margin of the peritreme, which in the
case of the metaspiracle is connected with the mesapodema. The
abdominal spiracles are very small and difficult to identfy. They
are located in two pairs on the seventh and eighth abdominal
segments. The rectum is chitinized forming a rectal tube bearing
rings of anacerores. The portion of the rectum situated between
the anus and the rectal tube is flexible and not strongly chitinized.
The sexes are indistinguishable during this nymphal stage.
The individuals of the first nymphal stage of this subfamily
are, so far is known, characteristic in form. The long clavate
SUBFAMILY MONOPHLEBINAE 65
antennae and the long prominent chalazae will serve to distinguish
them. The number of antennal segments varies with the genus
and in many cases even with the species. - _
The female nymphs of the second stage of purchasi are difficult
to recognize because of the variation in their size. The antennae
instead of being articulated to the margin of the body as in the
first nymphal stage are articulated to the ventral aspect. They
consist of the same number of segments, six, but the fourth and
fifth segments are of the same length and much shorter than the
other segments. The cerores are more numerous and, because of
the more numerous longer chalazae, the segmentation is more
difficult to determine. The cerores and chalazae are more
prominent near the margin of the body. While the swellings of
the anal chalazae are easily identified, their setae are only slightly
longer than the other setae. The thoracic and abdominal spiracles
are similar in number and position to those of the first nymphal
stage. There is a small round projecting area on the meson of the
ventral aspect of the caudal end of the body. This structure is
a discaloca and appears to be wanting or is only imperfectly
developed in the first nymphal stage.
The female individuals of the third nymphal stage, because
they intergrade in size between the large individuals of the second
nymphal stage and the small adults, are also difficult to identify.
The antennae have more than six segments, typically nine, but
individuals with seven or eight are not uncommon. A difference
in the number of segments of the two antennae of the same indi-
vidual is not unusual. The number of chalazae and cerores is
greatly increased. They are arranged in a distinct band around
the margin of the abdomen. The six anal chalazae can only be
identified with difficulty. The thoracic and abdominal spiracles
are similar in number and position. The spiraculariae are dis-
tinct. The rectal tube is trumpet-shaped in this stage and the
rings of anacerores, if present, have not been identified. The
discaloca is large and distinct. There is a smaller but similai
structure located on each side of the mesal discaloca. The mesa!
structure may be known as a mesodiscaloca and each of the others
as a latadiscaloca. The form of the discalocae has been used by
some systematists in differentiating spediies of this and ;other
genera. They have designated them as the vaginal discs, the
vaginal areoles, the ventral scars, and the subcircular scars.
The addlt females of purchasi are easily recognized by the
66 THE COCCIDAE
presence of the broad band of cerores and chalazae extending
around the margin of the abdomen and onto the sides of the
thorax. The presence of a partially or completely formed ovisac,
which is always formed in this species, is also positive proof of the
condition of the specimen. They vary from two and one-half tc
four millimeters in length. The antennae consist typically of
eleven segments, but adult females with antennae of ten segments
or even of nine are not rare. The parts of the head skeleton, the
rostrum, the rostralis, and the crumena are much larger than in
the nymphs, but do not differ in structure. The parts of the
thorax are similar but larger. The ventral aspect of the meso-
thorax and metathorax contain distinctly invaginated furcae. The
furcellae have not been identified. The spiracles of the thorax
and abdomen are similar in number, form, and position to those of
the nymphs. The vulva is located in the caudal part of the ventral
aspect, of the seventh segment. There is in certain genera a large
ventral gland, sometimes eversible, located caudad of the vulva.
This is often more distinct than the vulva and may be present in
the nymphal females as well as in the adult while the vulva is
present only in the adult. It is not impossible but that the mouth
of this gland, because of its distinctness, has been described as the
vulva in certain cases. The discalocae, three in number, are
located just caudad of the vulva. They are distinct black spots
and can be recognized in untreated specimens. The segments of
the abdomen are campanulate and the anus is located in the ninth
tergum. The chitinized rectal tube and rings of anacerores can
not be identified. The body of the untreated adult female shows
distinct transverse constrictions, the coriae, which are obscured in
mounted specimens by the stretching of the cuticle.
The eggs are deposited in a large white ovisac which is excreted
from the cerores along the margin and near the ventral aspect of
the body. As the wax is excreted the thorax and abdomen are
elevated until the body of the insect is held almost at right angles
to the surface of the host-plant. The ovisac is frequently much
larger than the body of the insect, often two or three times its
length, and according to the observations of Coquillet and Gossard
requires from sixty to ninety days for its completion. The rostralis
remains attached to the host-plant during all this time. The wax
constituting the central portion of the ovisac has a fibrous cottony
texture in ovisacs that have been dried for some time, but this wax
has a doughy stringy consistency in ovisacs covered by living
SUBFAMILY MONOPHLEBINAE 67
females. The wax is excreted from the cerores of the ventral
aspect and surrounds the eggs. The threads of wax from the
different cerores melt together as they come into contact. - The
characteristic external part of the ovisac is formed by the mar-
ginal band of cerores. The fluted external surface is produced by
the scalloped arrangement of the bands of cerores. The homo-
genous appearance of the wax is due to the melting together of the
wax from adjacent cerores. The fine granular wax of the dorsal
aspect is excreted by the cerores of this region. The long glassy
threads of wax are poured from the very large cerores of the dorsal
aspect. The cerores excreting these glassy threads are more
abundant upon the head and thorax. The difference in the
appearance of the wax of the dorsal aspect and that of the ovisac
is probably due to the fact that in the former the cerores are
distant from each other and there is no opportunity for the threads
from adjacent cerores to melt together.
The large fluted ovisac of purchasi, while characteristic of
this species, is not found even in all the species of the genus
Icerya. The females of the great majority of the species of this
subfamily do not form ovisacs of the purchasi type, although all
of them form a greater or less abundance of white wax. This wax
is usually described as cottony and as forming indefinite masses
which are used as much by the female in covering her body as in
covering her eggs. A striking exception to this is found in the
genus Stigmacoccus, where the body of the adult female is enclosed
in a thin more or less spherical hard waxy test with an apical
aperture through which half of the body of the female may be
extruded. A few species live in ant 's nests and the amount of wax
excreted is always small.
The female, as soon as a thin sheet of wax marking the
beginning of the ovisac is formed, begins to deposit her eggs. The
number of eggs in purchasi varies according to the condition of
the plant upon which the female has been feeding. Gossard
counted 600 to 800 while Koebele counted over 1,000 in a single
ovisac.
The life history of the male has not been so accurately
described as that of the female. In the first nymphal stage, the
males are indistinguishable from the females, but in the second
nymphal stage differ in having a longer more slender body and
longer and stouter antennae and legs. The antennae consist of
six segments and are similar in form to those of the female.
68 THE COCCIDAE
The nymphal males of the third stage are said to be easily
recognizable through their longer, narrower, flatter bodies. In
each successively older nymphal stage of the female there is an
increase in the number of cerores and setae, while in the male
there is a corresponding decrease in the number so that the body
of the male at this time is almost bare. The antennae consist
typically of nine segments. The male as it approaches the close
of its third nymphal stage becomes less active and seeks a place
where it can conceal itself under some projecting piece of bark oi
under some leaves. Having found such a place, it excretes a
flossy mass of wax threads which adhere together when they come
into contact with each other. The nymph forms a covering or
cocoon from this wax by the movements of its body.
When the formation of the cocoon is completed, the nymph
becomes quiescent and soon molts and transforms to a legless indi-
vidual. The nymph is quiescent throughout this period, the fourth
nymphal stage, and differs from the individuals of the previous
period in that it has lost its rostralis and is unable to feed. It has
also lost its nymphal legs and antennae and has in place of them
the developing antennae and legs and wings of the adult. The
insect is quiescent throughout the fourth nymphal period which
ordinarily lasts about three weeks. This is the so called pupal
stage of many writers.
The adult males are reddish insects with long, slender, ten-
segmented antennae. The mesothoracic wings are comparatively
large while the metathoracic wings are represented by a pair of
halteres each of which bears two hooks which fit in a fold in a
mesothoracic wing. The abdomen is without the long slender
lateral prolongations of each side of certain of the caudal abdominal
segments so characteristic of many of the males of this subfamily.
The following table for the separation of the genera of the
Monophlebinae is very unsatisfactory. It is based for the most
part upon generalities, upon the amount and form of the wax
excreted by the adult female and the number of antennal segments.
These are characters of slight value in differentiating genera
because of their great variation even in the same species. Such
genera as Lophococcus, Aspidoproctus, Stigmacoccus, and Perissop-
neumon, which are based upon definite structural characters of the
body, are readily recognized. The size of the insects, the great
number of structural characters, the largest found in any group
of coccids, and the presence of structures not present in other
SUBFAMILY MONOPHLEBINAE 69
coccids should make it possible to define the genera so that- they
would have definite limits and without having to utilize the
covering of wax or the number of antennal segments. The form,
number, and arrangement of the cerores; the form, number, and
position of the abdominal spiracles; the apodemae; the spiracu-
lariae ; and numerous other characters should be found useful. It is
to be hoped that some investigator with typical specimens of the
genera will make a careful morphological and phylogenetic study
of the subfamily.
GENERA OF MONOPHLEBINAE
a. Abdomen of male with two or four pairs of slender lateral fila-
ments; female unknown. Monophlehus Leach.
aa. Abdomen of male with or without slender lateral filaments, if with
filaments, never with two or four pairs; female usually known,
b. Antennae of adult female normally with nine or eleven segments.
c. Antennae of adult female normally with eleven segments, if
with nine or ten segments, female provided with distinct ovisac
located between body of female and host-plant, its outer surface
frequently ribbed or fiuted.
d. Wax excreted by adult female powdery or cottony, soft and
indefinite in form, not forming definite ovisac,
e. Body of adult female with isolated dorsal areas of powdery
or cottony excretion, body never entirely concealed by
excretion,
f. Body of adult female elongate with powdery or cottony
wax; vulva normal in size and position; abdomen of
male with four pairs of slender lateral filaments
Llaveia Sign,
ff. Body of adult female strongly convex and dorsum with
transverse bands of powdery wax; vulva large and dis-
tant from caudal end of body; abdomen sometimes with
single pair of spiracles; abdomen of males without
slender lateral filaments. Palaeococcus C'kll.
ee. Body of adult female completely covered bj'' dense fine
cottony excretion of long threads of wax, entirely con-
cealing body of female; male unknown. __GwenneZZa Fern,
dd. "Wax excreted by adult female definite in form and usually
hard, forming definite ovisac located between body of female
and host-plant, its outer surface usually ribbed or fluted;
dorsal aspect of body with powdery wax and frequently mar-
gined with erect or decumbent processes of wax; abdomen
with spiracles on segments seven and eight or six to eight;
abdomen of males without slender lateral filaments or with
single pair of short blunt projections Icerya Sign.
cc. Antennae of adult female with less than eleven segments.
70 THE COCCIDAE
d. Antennae of adult female normally with ten segments.
e. Wax excreted by adult female powdery or cottony, soft
and indefinite in form,
f. Adult female without abdominal spiracles; abdomen
normal in form, not with stout erect spine on dorsum;
abdomen of male with four pairs of slender lateral fila-
ments. Tessarobelus Montr.
ff. Adult female with distinct abdominal spiracles; abdo-
men abnormal in form, with short, stout, erect spine on
dorsum shaped like the spine on a military helmet;
male unknown. Lopococcus Ckll.
ee. Wax excreted by adult female definite in form and usually
hard, in shape of conspicuous lamellae or processes which
conceal body more or less.
f. Vulva normal in position and size, not conspicuous, and
not covered by hinged flap or valve of wax; male un-
known. Walkeriana Sign.
ff. Vulva abnormal in size, covered by thin valve or flap
of wax hinged at cephalic end, supposed to form a re-
ceptacle for the eggs and nymphs; male unknown
Aspidoproctus Newst.
dd. Antennae of adult female normally v/ith nine segments,
body of adult female with powdery or cottony covering of
wax; cuticle soft; abdomen of male v/ith five pairs of slender
lateral filaments. Drosicha Walk.
bb. Antennae of adult female normally with less than nine segments.
c. Antennae of adult female normally with eight segments.
d. Abdomen of adult female whether provided v/ith spiracles
or not, never with large crescent-shaped valve-like structure
located between them,
e. Abdomen with seven or eight pairs of spiracles.
f. Abdomen of adult female with eight pairs of spiracles,
pair on first abdominal segment; body of adult female
enclosed in more or less spherical hard waxy test with
apical aperture; male unknown Stigmacoccus Hemp.
ff. Abdomen of adult female, if bearing spiracles, provided
with seven pairs of dorsal spiracles, those of first ab-
dominal segment wanting; body of adult female not
enclosed in spherical hard waxy test, but excreting a
■■' distinct ovisac formed of numerous white threads which
sometimes fuse and form a fluted surface; abdomen of
males with three or four pairs of lateral filaments
Greeniella MacG.
ee. Abdomen with six pairs of spiracles; head between and
caudad of antennae with setaceous clypeus-like lobe, cov-
ering rostrum in part; antennae consisting of eight to
nine segments; setae clavate or knobbed at distal end;
body spherical; male unknown Mimosicerya Ckll.
?
SUBFAMILY MONOPHLEBINAE 71
dd. Abdomen of adult female provided with seven pairs of
subdorsal spiracles; abdomen with large crescent-shaped
valve-like structure located between spiracles; test unknown;
male unknown. Perissoimeumon_ Newst.
cc. Antennae of adult female normally with less than eight seg-
ments,
d. Antennae of adult female normally with seven segments;
wax excreted by adult female not prominent; male unknown.
Monophlebulus Ckll.
dd. Antennae of adult female with less than seven segments.
e. Abdomen with caudal end not bearing bifurcate setae;
body with mealy covering of wax and with transverse
ridges of wax on abdomen; antennae of adult male con-
sisting of six segments, sixth segment subequal in length
to each of others; male unknown Cocker ellella MacG.
ee. Abdomen bearing bifurcate or serrate or broadly truncate
setae.
f. Abdomen with caudal end bearing three or four bifur-
cate setae on each side of meson; antennae consisting
of six segments in first and third nymphal stages, sixth
segment twice as long as any of other segments; rostrum
consisting of two segments; male unknown
Newsteadiella MacG.
ff. Abdomen of adult female margined on each side with
bluntly or sharply serrated setae or broad blunt setae,
these setae also present in all nymphal stages; antennae
of adult female consisting of four to six segments, of
second and third nymphal stages of five to six segments,
and of first nymphal stage of four to five segments;
rostrum apparently consisting of a single segment;
adult male unknown. Stictococcus Ckll.
The foregoing table to the genera of Monophlebinae has made
it necessary to change the position of certain of the species. The
following list gives their generic reference : —
Monoplebus Burm. — This genus is limited to males, two species
of which, atripennis Burm., the type, of which only males are known,
and dubius Fabr., each have two pairs of lateral filaments on the abdo-
men, and a third, saundersii Westw,, has four pairs of lateral fila-
ments. All these species are from the East Indies. The following
three species based upon females from Kiliman/ijaro, Africa, are re-
ferred to this genus by Newstead: sjostedti -Newst., pallidus Newst.. and
africana Newst. A single species, dugesi, from Mexico has been de-
scribed by Vayssiere. It is not likely that these latter species are
congeneric with the type of Monophlebus.
Lilaveia Sign. — There are eight species included in this genus, as
follows: — axin Llave from Mexico on Spondias, Jatropha, mango;
bouvari Sign, from Guatemala; cacti Ckll. (not Linnaeus) from the
72 THE COCCIDAE
West Indies on cactus; championi Ckll. from Panama, known only in
the male sex, transferred" from Tessarobelus which contains a single
species from New Caledonia; mexicanorum Ckll. from Mexico on Acacia;
primitiva Towns, from Mexico on "Nettle-tree;" uhleri Sign, from
Ecuador; luzonica Ckll. from Philippine Islands. The following table
based upon one published by Cockerell will aid in the separation of
the American species: —
SPECIES OF LLAVEIA
a. Body large, length of adult female 15 to 25 mm.
b. Body with scant pubesence; length 15 to 18 mm.; antennae short,
segments, except the distal, broader than long houvari Sign,
bb. Body with abundant pubescence; length 23 to 25 mm.; legs and
antennae reddish.
c. Species from Mexico; antennae with cuticle densely covered
with short setae, segments two to eight times as broad as long,
nine and ten broader than long. axin Llave.
cc. Species from Ecuador. uhleri Sign.
aa. Body small, length of mounted specimens of adult female about 7
mm.; legs and antennae dark sepia brown.
b. Cuticle of body densely covered with setae; antennae long, seg-
ments two to six not twice as broad as long, segments six to ten
longer than broad, segment eleyen narrow and cylindrical
primitiva Towns.
bb. Cuticle of body with minute, sparse, scarcely noticeable setae; an-
tennae short, segments two to eight twice as broad as long, seg-
ments nine and ten considerably broader than long, segment
eleven oval. mexicanorum Ckll.
^ PalaeococcTis Ckll. The following species are included in this
genus: — ^australis Mask, from Australia on Hakea; braziliensis Walk,
from South America; ewarti Newst. from West Africa on "Peppet;"
fortis Ckll. from Natal on Eucalyptus; fuller! Ckll. from Natal on grass;
fuscipennis Burm. from Europe on oak, maple, fir, pine; hellenicus Genu,
from Greece on Pinus; mexicanus Ckll. & Parr, from Mexico on Prosopis;
morrilli Ckll. from Arizona; nudatus Mask, from Australia on Cosmos,
Verbena; plucheae Ckll. from New Mexico on Pluchea; pulcher Leon,
from Java on Ilex; rosae R. & H. from Florida, Jamaica, Mexico on Am-
herstia. Euphorbia, Prosopis, rose, sugar-apple, lime, lemon, lignum-vitae;
theobromiae Newst. from Calabar, West Africa on Cacao; townsendi Ckll.
from New Mexico on Townsendia, Gutierrezia, Picradenia, Grindelia,
Bahia. Ferris has studied three of the above species, morrilli Ckll., plu-
cheae Ckll., and townsendi Ckll. Each of these species has a large
marsupium in the adult female which opens on the ventral aspect just
caudad of the metathoracic legs. The opening after it is filled with eggs
is closed by a flap of wax. These species may not be congeneric with the
type of the genus, fuscipennis Burm.
Guerinella Fern. — This name was proposed by Mrs. M. E. Fernald
\
SUBFAMILY MONOPHLEBINAE 73
for Guerinea which is preoccupied. It includes a single species, serra-
tulae Fab. from Europe and Algeria on Serratula.
Icerya Sign. — This is the largest genus of the subfamily so far as
number of species is concerned. The following are included: — aegyptiaca
Dougl. from Egypt, Ceylon, India, Australia on Ficus, Croton, Goodenia,
Aristochlia; albolutea Ckll. from West Africa on Anona; brasiliensis
Hemp, from Brazil on Liriodendron, Laurus, Codiaeum; Candida Ckll.
from Philippine Islands; colinensis Ckll. from Mexico; crocea. Green
from Ceylon on Citrus, Croton, Cocculus; genistae Hemp, from Brazil
on Genista, Lespedeza, Fragaria; jacobsoni Green from Java on Dom-
beya; koebelei Mask, from Australia on Leptospermum, Acacia; littor^lis
Ckll. from Mexico on Croton, Mimosa; longisetosa Newst. from German
East Africa on Acacia; minor Green from India on mango; montser'
ratensis R. & H. from West Indies, Mexico on Chrysophyllum, Clusia,
palms, orange, banana, avocardo pear; natalensis Dougl. from Natal on
Acacia, orange, lemon; okadae Kuw. from Japan on orange; palraeri
R. & H. from Mexico on Coufsetia, grape-vine; pilosa Green from Ceylon
on grass; purchasi Mask, from Australia, New Zeland, Fiji Islands, Ha-
waiian Islands, California, Louisiana, Florida, Mexico, West Indies,
South Africa on Pittospermum, Acacia, Cypress, orange, lemon, pine,
rose, grass, etc.; rileyi Ckll. from New Mexico, Mexico on Larrea, mes-
quite; schrottkyi "Ttemp. from Brazil; seychellarum Westw. from Sey-
chelles Islands, Mauritius, China, Formosa on citrus, Podocarpus, guava,
palms, rose, sugar cane; subandina Leon, from Argentina on Bulvesia;
tangalla Green from Ceylon on Gutierrezia; and ze^_ki Ckll. from Panama
Canal Zone. The following tables based upon tables published by Riley
and Howard may be found useful in separating the included American
species: —
SPECIES OF ICERYA
FIRST STAGE NYMPHS
a. Abdomen with lateral setae cephalad of six anal chalazae minute,
always much less than one-eighth length of anal chalazae.
b. Antennae with distal segment bearing many long setae, five of
which are especially long. seychellarum Westw.
bb. Antennae with distal segment bearing less than five especially
long setae.
c. Antennae with distal segment bearing four especially long
setae; abdomen not emarginate between caudal pair of anal
setae. . purchasi Mask.
cc. Antennae with the distal segment bearing two especially long
setae; abdomen emarginate between ithe caudal pair of anal
setae. aegyptiaca Doug.
aa. Abdomen with lateral setae cephalad of six anal chalazae not minute,
about one-fifth length of anal chalazae.
b. Antennae with distal segment not constricted near middle and
enlarged on proximal half, sides uniformly convex
montserratensis R. & H.
5
7
74 THE COCCIDAE
bb. Antennae with distal segment constricted near middle and prox-
imal half distinctly swollen, two sides of distal segment not
uniformly convex. palmerii R. & H.
ADULT FEMALES
a. Body furnished with many long glassy filaments of wax arising
from dorsal cerores.
b. Body supported by ovisac with distinct longitudinal flutings
purchasi Mask.
bb. Body supported by ovisac without longitudinal flutings
seychellarum Westw.
aa. Body not furnished with long glassy filaments of wax.
b. Body supported by ovisac with prominent longitudinal fiutings;
tufts of wax bordering dorsal aspect limited to two at each end,
each tuft usually much longer than entire ovisac
montserratensis R. & H.
bb. Body supported by ovisac with only faint longitudinal flutings;
tufts of wax bordering dorsal aspect ten or more in number,
longest not much longer than ovisac. aegyptiaca Doug.
Tessarobelus Montr. — A single species, guerinii Montr., described
from New Caledonia on Melaleucae and Filices, is included.
liophococcus Ckll. — This genus includes a single species, mirabilis
Ckll., from Natal on Mimosa. Newstead has described a variety, mirabilis
tricornis Newst., from Dutch Southwest Africa and Vayssiere a second
species, vuilleti, from West Africa on Acacia.
Walkeriana Sign. — The following species are included in this
genus: — andreae Green from Congo, Africa; compacta Green from Ceylon;
euphorbiae Green from Ceylon on Euphorbia; floriger Walk, from Ceylon
on Litzea; poleii Green from Ceylon on Dodonaea; and senex Green from
Ceylon on Dodonaea.
Aspidoproctus Newst. — A single species, pertinax Newst., was
originally described from Central Africa. Two additional species, arma-
tus Newst. on Acacia and maximus Newst, on Brachystegia, have been
added from German East Africa, while Vayssiere has added bouvieri
Vays. from French Equatorial Africa and serrei Vays. from Batavia,
Java.
Drosicha Walk. — The following species belong in this genus: —
burmeisteri Westw. from Java, China, Japan on pine; contrahens Walk,
from Ceylon, North China; corpulenta Knw. from Japan on oak; craw-
fordi Mask, from Australia on Eucalyptus; illigeri Westw. from Tas-
mania; leachii Westw. from India; lichenoides Ckll. from Philippine Is-
lands on Ficus; maskelli Ckll. from Java, China, Japan on Ficus, Gar-
denia, pine; raddoni Westw. from West Africa.
Mimosicerya Ckll. — A single species, hempeli Ckll. from Brazil
on Mimosa (?), is included. Newstead has described and figured this
coccid with considerable detail and proposed the new generic name Cly-
peococcus for it.
Stigmacoccus Hemp. — This genus includes a single species, asper
Hemp, from Brazil, where it is found in the nests of Campanotus and
upon the bark of Inga.
SUBFAMILY MONOPHLEBINAE 75
Greeiiclla MacG. — The following species, originally referred to the
genus Monophlebus, are included in this genus: — dalbergiae Green from
India; stebbingii Green from India on Ficus, Dalbergia, Areocarpus,
mango, is the type; and tamarinda Green from India on branches of
tamarind.
Perissopiieunioii Newst. — A single species, ferox Newst., described
from India from the nests of ants is the type. Newstead has described
a second species, zimmermanni Newst. from German East Africa.
Monophlebulus Ckll. — There is only a single sjpecies, fuscus
Maskell described from Australia, included in this genus.
Newstcadiella MacG. — A single species, formicarum Newst., col-
lected in the nests of ants in India and originally described as a species
of Icerya, is included.
Cockerellella MacG. — This genus includes a single species, town-
sendi Ckll. from the Philippine Islands. It was originally described as
a species of Monophlebulus.
Stictococcus Ckll. — A genus containing four species: — sjostedi
Ckll. from Cameroons, Africa from the stalks of the pods of Theobroma;
multispinosus Newst. from Kilimanjaro, East Africa; dimorphus Newst.
from German East Africa; and formicarius Newst. from Upper Congo,
Africa from the hollow stems of Barteria and Cuviera. This genus, orig-
inally referred to the Lecaniinae and later to the Dactylopiinae. appears
to belong to the Monophlebinae.
CHAPTER VI
SUBFAMILY KUWANIINAE
The Sinoran Coccids
The body of the adult female is rotund or convex on the dorsal
and flat on the ventral aspect. It is elongate elliptical, broadest in the
thoracic region. The lateral and caudal margins are convex, not sub-
transverse. The segments are well defined. The caudal end is not
inflated and the cephalic and lateral margins are not provided with a
keel. The antennae are prominent, consist of eight to ten segments, and
are articulated to the ventral aspect of the head near each lateral
margin. The eyes are present. There is a single ocellana located on
each side of the head near the articulation of an antenna. The mouth-
parts, rostrum and rostralis, are always wanting. The legs are present.
They are subequal in length and normal in form. The profemora are
never enlarged and the prothoracic legs are never fitted for digging. The
trochanto-femoral and the tibio-tarsal sutures of all the legs are always
distinct. The spiracles of the mesothorax and metathorax are normal
in form and ventral in position. The mesothoracic spiracles are never
located near the anus. The abdominal spiracles are located on the
ventral or subventral aspects, are variable in number, and are never
large in size. The spiracular tracheae of the abdominal spiracles are
provided with one or more rings of cerores. The stigmatic clefts, spinae,
and canellae are wanting. The abdomen is about as long as or longer
than the head and thorax together. It is convex on the dorsal and
ventral aspects, never depressed. The lateral margins are convex or
subconvex and the caudal end is convex, never subtruncated. All the
segments are fully exposed and similar in appearance with distinct
coriae between them, none are retracted to form a marsupium. The
abdomen is never provided with an anal cleft and opercula, an anal ring
and anal ring setae, or anal lobes and anal setae. The caudal abdominal
segment is never short and narrow and extending beyond the other seg-
ments. The body is provided with cerores, none of which are octacerores
or pilacerores, and is never provided with ceratubae. The body is never
enclosed in a resinous cell with three adjacent openings or covered by
a scale with or without exuviae. It may be covered more or less or
associated with white m^ealy or powdery wax or loose filaments of wax.
The rectum is never provided with a long glassy tube of wax or with a
chiiinized rectal tube bearing rings of anacerores. The body is never
naked and gall-like in form.
The female nymphs of all stages are provided v/ith a rostrum of two
or three segments and a long rostralis. The three pairs of legs are
always present in first nymphal stage, so far as known, wanting in
all others. The profemora are never greatly enlarged and the
76
SUBFAMILY KUWANIINAE 77
prothoracic legs are never fitted for digging. The body is never provided
with .an anal cleft and opercula, an anal ring and anal ring setae, anal
lobes and anal setae, octacerores, pilacerores, or ceratubae. The caudal
end of the rectum is not chitinized, forming a rectal tube provided with
one or more rings of anacerores excreting a long glassy tube of wax.
The caudal abdominal segment is not short, narrow, and projecting.
The male has compound eyes. The abdomen is never provided with
long lateral filaments, but may be provided with tufts of long wax fila-
ments. The stylus may be large and prominent.
This subfamily contains six genera and six species, two
species from Japan, one from Ceylon, one from Russia and Eng-
land and two from California. The complete life cycle of only
one species, one of the American species, is known.
The most striking difference found in the different genera is
in the number of abdominal spiracles. Paragreenia according to
Ferris has eight pairs, while the spiracles of segments seven and
eight are greatly reduced and are but little more than points for
the attachment of their spiracular tracheae. The presence of this
number of abdominal spiracles is of interest, because eight pairs
of abdominal spiracles on segments one to eight is the maximum
number found in the abdomen of any embryonic, immature, or
adult insect. Steingelia, which has six pairs of abdominal spiracles
according to the description and figures of Green, has lost those
of segments seven and eight. Kuwania has four pairs of abdominal
spiracles on the four cephalic segments and has lost those of
segments five to eight. It might be assumed since no abdominal
spiracles are figured or described in Matsucoccus that there
existed a beautiful series from the maximum number of Para-
greenia to their entire absence in Matsucoccus. Since no thoracic
spiracles are described or figured in Matsucoccus and since the
spiracles are frequently considered of little systematic value and
are frequently omitted from figures and descriptions, it is more
likely that when specimens of this genus are examined, not only
thoracic but abdominal spiracles will be found. The other char-
acters of Matsucoccus are of sufficient importance to make it worthy
of consideration as a distinct genus.
GENERA OF KUWANIINAE
a. Antennae with ten segments; abdomen apparently without spiracles;
female nymph of first stage with femur of each leg enlarged and
elongated, distinctly longer than combined tibia and tarsus and claw,
so that legs are crab-like in appearance. Matsucoccus Ckll.
aa. Antennae with less than ten segments; abdomen always with spira-
cles.
78 THE COCCIDAE
b. Antennae with nine segments.
c. Abdomen with eight pairs of spiracles. Paragreenia MacG.
cc. Abdomen with less than eight pairs of spiracles,
d. Abdomen with seven pairs of spiracles; all nymphal stages
known, normal in form. A7nericoccus MacG.
dd. Abdomen with four cephalic segments with pairs of spiracles.
Kuwania CklL
bb. Antennae with less than nine segments.
c. Antennae with eight segments; abdomen with spiracles on seg-
ments one to six; nymphal female of first stage not with femur
of each leg enlarged and elongated Steingelia Nassanow.
cc. Antennae with seven segments; abdomen with spiracles on
segments one to six; adult female with rostrum and rostralis
present. Stomacoccus Ferris.
Matsucoccus Ckll. — A single species matsumurae Kuwana from
Japan on pine-tree (matsu), is known. The adult female has a swollen
body which is ovate in outline. When ready to deposit her eggs, the
female seeks a crack or crevice in the bark, v/here she forms a cottony
cushion of wax upon which she rests and covers her body with a con-
siderable amount of white cottony wax. The nymphs of the first stage
have the body similar in form to that of the adult female, but the legs
have a peculiar crab-like appearance due to the swollen and elongate
femora and the short and slender tibiae and tarsi. The antennae of this
stage, which consist of seven segments, are peculiar in form. Segments
two, four, and six are subequal in length as are segments three, five, and
seven, but the segments of the last group are only about one-third the
length of the segments of the first group.
Paragreenia MacG. — Green has described a single species, zeylanica,
from Ceylon on Antidesma. The complete description of this species has
not been published. The body of the adult female is long and narrow and
the female nymphs of the second stage are found in small cavities in the
living tissue of the plant. The adult female according to Ferris has well
developed mouth-parts with a distinct mentum and six to eight pairs of
abdominal spiracles. If this species has mouth-parts, rostrum and ros-
tralis, in the adult female, this genus should be referred to the Monophle-
binae.
Americoccus MacG. — There is a single described species, fascic-
uleusis Herbert, from California on Pinus. The adult female has trans-
versely striated antennae, the proximal segments are large. The claws
are simple with two ungual digitules. The thoracic spiracles are large
and distinct. The dorsum is provided with transverse rows of simple
cerores and the dorsum and venter with cerores similar to the octa-
cerores. The ocellanae, one on each side of the head, are present. The
mouth-parts are sometimes present. The distance between the prothoracic
and mesothoracic legs in the first nymphal female is nearly three times
that between the mesothoracic and metathoracic legs. The rostrum is
shown between the latter pairs of legs.
Kuwania CklL — Kuwana has recorded a single species, quercus
Kuwana, from Japan on Quercus. This genus, as here limited, contains
SUBFAMILY KUWANHNAE 79
only the type species. The adult female has an elongate oval body with
all the surfaces convex. The antennae consist of nine segments and are
articulated to the head near each other. The tarsal claws are without
digitules. The nymphs of the first stage do not have crab like legs, Llieir
femora and tibiae and tarsi are normal in form. The antennae in this
stage consist of six segments. The distal segment is greatly enlarged,
clavate, and longer than all the other segments together. In the other
nymphal stages the legs are wanting, the antennae are mere chitinized
points, and each of the four cephalic abdominal segments is provided
with a pair of spiracles.
Steingelia Nassanow. — The type of this genus, gorodetskia
Nassanow, was described from Russia. It was described a little later
by Green from England on birch as Kuwania britannica. Green considers
the two species as synonymous. The adult female of Steingelia is very
different in general appearance from either of those previously described.
The body is long and slender, three or four times as long as broad, and
the lateral margins are parallel or nearly so. The antennae consist of
eight or nine segments and are articulated to the cephalic portion of the
head and near to each other. The body of the adult female is greatly
shriveled during ovoposition and covered with a woolly secretion of wax,
forming an ovisac for holding the eggs. The ovisac may be placed in
crevices in the stems of old birch trees, but are found more abundantly
among the decayed leaves and debris at the base of the trees. The nymphs
of the first stage do not have crab-like legs. Their antennae consist of
six segments. The sixth segment is the longest, the second, third, and
fourth are the smallest, and the fifth is broadly dilated.
Stomacoccus Ferris. — There is a single species, platani Ferris,
from California on Platanus. Its most distinctive feature is the reten-
tion of mouth-parts in the adult female. The body is elongate as in the
other species of the subfamily. The thoracic legs are wanting in all
nymphal stages but the first. This character will have to serve for
differentiating the genera of this subfamily from the Monophlebinae.
The single species differs from Steingelia in having a single row of
setae on each side of the ventro-meson of the abdomen, while there are
two rows on each side in Steingelia. There are six ungual digitules in
Stomacoccus, eight to ten in Steingelia, and none in Kuwania but its
claws are toothed. Kuwania has a group of about eight tenent setae on
each coxa which are wanting in Stomacoccus. The insect apparently
passes the winter upon the bark, migrating onto the leaves in spring.
They live exposed upon the underside of the leaves. The males are very
active. This genus was overlooked until after this chapter was in type.
CHAPTER VII
SUBFAMILY XYL.OCOCCINAE
The Melratuhan Coccids
The body of the adult female is rotund or convex on the dorsal and
flat on the ventral aspect. It is elongate elliptical or rotund and glob-
ular, broadest in the thoracic region. The lateral margins are convex
and the caudal margin is subtransverse or uniformly convex. The
segments are well defined. The caudal end of the body is not inflated
and the cephalic and lateral margins are not provided with a keel. The
antennae are prominent, consisting of five to eleven segments and are
articulated to the ventral aspect of the head near each lateral margin.
The eyes are present. There is a single ocellana located on each side of
the head near the articulation of an antenna. The mouth-parts, rostrum
and rostralis, are always wanting. The legs are usually present. They
are subequal in length and normal in form. The profemora are never
enlarged and the prothoracic legs are never fitted for digging. The tro-
chanto-femoral and the tibio-tarsal sutures of all the legs are always
distinct. The spiracles of the mesothorax and metathorax are small and
inconspicuous and ventral in position or are large and conspicuous.
The mesothoracic spiracles are never located adjacent to the anus. The
abdominal spiracles are located on the lateral or ventral aspects, six or
eight pairs, and are large in size. The spiracular tracheae of the
abdominal spiracles are frequently provided with one or more rings of
cerores. The stigmatic clefts, spinae, and canellae are wanting. The
abdomen is about as long as the head and thorax together. It is convex
on the dorsal and ventral aspects, never depressed. The lateral margins
are convex and the caudal end is subtruncated or convex. All of the
segments are fully exposed and similar in appearance with distinct coriae
between them, none are retracted to form a marsupium. The abdomen
is never provided with an anal cleft and opercula, an anal ring and anal
ring setae, or anal lobes and anal setae. The caudal abdominal segment
is short and narrow, small and inconspicuous, and extends beyond the
other segments or is normal in form and not protruding. The segments
are not campanulate. The body is provided with cerores, none of which
are octacerores or pilacerores, and is never provided with ceratubae. The
body is never enclosed in a resinous cell with three adjacent openings or
covered by a scale with or without exuviae, but may be more or less
covered or associated with white, mealy, powdery, or filamentous wax.
The rectum is rarely if ever provided with a long glassy tube of wax
or with a chitinized rectal tube bearing rings of anacerores. The body
is never naked and gall-like in form.
The female nymphs of all stages are provided with a rostrum of two
segments and a rostralis. The three pairs of legs are present in the first
80
SUBFAMILY XYLOCOCCINAE 81
nymphal stage and wanting or greatly reduced in size and inconspicuous
in the second, third, and fourth; and when present are similar in form.
The profemora are never greatly enlarged and the prothoracic legs are
never fitted for digging. The body is never provided with an anal cleft
and opercula, an anal ring and anal ring setae, anal lobes and anal setae,
octacerores, pilacerores, or ceratubae. The caudal end of the rectum is
strongly chitinized and provided with one or more rings of anacerores
excreting a long glassy tube of wax. The caudal abdominal segment is
short, narrow, and projects beyond the other segments; the segments are
not campanulate.
The male has large coarsely facetted compound eyes and are not
provided with ocelli. The abdomen is not provided with long lateral
filaments, but bears tufts of long filaments of wax. The stylus is short
and conical.
The typical genus, Xylococcus, of the subfamily Xylococcinae
was based upon the species filiferous Loew, described in 1882 as
infesting Tilia europaea and T. grandifolia in Austria. Four
species have been described from North America, betulae Pergande
from birch and poplar from the Lake Superior region, and three
species from California, quercus Ehrhorn from live oak, macrocar-
pae Coleman from the Monterey cypress, and alni Florence from
alder. Two species have been described from Japan by Kuwana,
napiformis from oak and matsumurae. The first species has been
made the type of a new genus, Matsucoccus by Cockerell. It has been
placed in the subfamily Kuwaniinae. The second species, judging
from the figures and descriptions of the species, the presence of
mouth-parts in the adult female, and the presence of anal setae and
apparently anal ring setae in the first nymphal stage, has no
relation with the genus Xylococcus as here defined and probably
belongs to the Eriococcinae.
The most complete account of the life history of any of the
species of Xylococcus is that given by H. G. Hubbard and Th.
Pergande of the birch xylococcid. The account of Florence based
upon the study of the three species found upon the Pacific Coast
confirms for the most part and amplifies the excellent account of
these observers. The following descriptions are based upon the
work of these writers.
The adult female is ordinarily buried in a cavity in the bark.
The eggs are deposited in a depression formed by the collapse of
the ventral portion of the abdomen of the female and the sides of
the opening in the bark.
The young nymphs, when they emerge from the Qgg, seek a
place like other coccids of the first nymphal stage, where they can
82 THE COCCIDAE
attach themselves. The young nymph works its way in between
the layers of bark or enters a lenticel, or breathing pore. The
insect by its growth and the excretion of wax causes the layers of
bark to separate and curl, producing areas where the bark is
roughened and covered with splits and curls. The bark also in its
growth surrounds the body of the coccid forming a cavity in which
it rests.
The body of the first stage nymph is more or less depressed,
without tubercles, without an anal ring, but with the caudal
abdominal segment, the ninth, a short transverse tubercle-like
projection. The caudal abdominal segments are never campanulate
as is characteristic of these segments of many coccids. There is a
row of large cerores, varying in number in the different species,
on the ventro-meson. The females and males are similar during
the first nymphal stage but quite different during the other
nymphal stages and for this reason they will be considered sep-
arately. The nymphs are provided with prominent legs, rostrum,
rostralis, and antennae of six segments each.
The females at the first molt loose their legs and antennae,
but retain their mouth-parts. The body is somewhat depressed
and sack-like. There are two other molts producing two other
nymphal stages, four in all. The nymphs differ in general appear-
ance only in being successively larger. The insect, since it has no
legs and also because of its location, is stationary during these
stages. The fourth molt produces an adult female which is pro-
vided with distinct nine-segmented antennae and legs. The
rostrum and rostralis are lost at this molt so that the insect is
unable to feed. Ferris calls attention to the fact that in certain
individuals of macrocarpae a complete series can be shown of
practically every stage from a complete absence of mouth-parts to
mouth-parts which are apparently functional. He does not specify
whether it is only the rostrum or labium that is present or both
rostralis and rostrum. There is a striking difference in this con-
dition as the rostralis must be present in order that the insect can
procure food. The body of the adult female is distinctly segmented
and the two ends are more or less truncated. The insect is now
able, if it desires, to leave its cavity in the bark and to wander
about which it sometimes does. But it is ordinarily so deeply
buried in the bark that it is unable to leave its cell and in fact
does not entirely free itself from the cast cuticle of the fourth
nym^phal stage. In such cases the caudal end of the molted cuticle
SUBFAMILY XYLOCOCCINAE 83
is ruptured and the female presents the end of her body at the
crevice in the bark for the reception of the male.
The nymphs which develop into males never form pits like
those of the female nymphs. They are apt to assemble in the
vicinity of some older females and to establish themselves under the
protection of her accumulation of wa^. This makes its possible
for them to escape later.
The males of the first nymphal stage are, as noted, similar
to the females. They differ from the females in that at the first
molt they do not lose their antennae and legs. They also retain
their rostrum and rostralis and feed like the females.
According to Pergande the males molt five times and pass
through five nymphal stages. Florence in the species which she
examined only found four nymphal stages. The male nymphs at
the second molt, the third nymphal stage, lose their legs and
antennae but retain their rostrum and rostralis. They closely
resemble the female nymphs of the third stage. At the third molt
the rostrum and rostralis are lost and the insect regains its legs
and antennae. During this stage, the fourth nymphal stage, the
insect leaves its w^axen cyst and wanders about. The body is soon
covered with little tessellations of cottony wax which forms a loose
flocculent follicle. The insect rests in this follicle until ready for
the fourth molt. Neither of the accounts make clear whether these
legs and antennae are new nymphal appendages or are the ap-
pendages that are passed on later to the adult. At the fourth molt,
which is usually passed under some sheltering fragment of bark,
the male appears with wing-pads and protuberances surrounding
the genital organs. The fifth stage nymph wanders about freely
for a time, but eventually covers itself with a cylindrical cocoon of
threads of wax. It later breaks its way out of this follicle, molts
its cuticle for the fifth time, and transforms to a winged adult.
The adult male is provided with coarsely faceted compound
eyes and is said to be without ocelli. The abdomen bears a short
style, a long penis, and the sixth and seventh abdominal segments
bear a dense brush of delicate thread-like filaments of wax. The
metathoracic wings are of interest in that they contain veins,
proving that the halteres of the males of the specialized families
are modified wings. The males emerge during the fall and winter
and females with eggs are found during the winter and spring.
The eggs of a single female hatch over a long period of time, five
to six weeks.
84 THE COCCIDAE
One of the striking features of the anatomy of the species of
Xylococcus is the chitinization of the caudal end of the rectum,
forming the so-called anal tube. This chitinized portion is pro-
vided with one to three rings of anacerores. This structure is
present in all the female nymphal stages. It becomes more and
more highly organized with each successive stage. Pergande does
not mention its presence in the adult female but states: "The
anal opening is simple, situated on the underside of the abdomen
close to the concavity, and with difficulty to be seen." Florence
in her description of the adult female of each species, states :
**Anal tube represented by an infolding of the body- wall and not
strongly chitinized." It is apparent that the chitinization of the
rectum is peculiar to the nymphal stages as in Coelostomidea and
Icerya. The function of the rings of anacerores is to excrete the
long hair-like filament of glassy wax which is protruded through
the anus. These filaments are often two inches or more in length
and contain at the free end a viscid droplet of honeydew. The
male is provided with a chitinized rectum in the first three nymphal
stages and excretes a long glassy thread-like tube of wax like that
of the female. This tube is lost in the male at the third molt.
Where these insects are abundant, the large number of waxen
threads give the bark a hoary appearance.
The body of the female and male nymphs is covered with
eerores which are more abundant along the lateral and caudal
portions. In this region there is excreted thick masses of wax
which undoubtedly aids in the curling and splitting of the bark.
The wax is in the form of curls which are consolidated into thick
lamellae on the sides of the body.
The development of the respiratory system as described and
figured by Pergande is unique. He does not mention or figure
any thoracic spiracles but Hubbard states that ''there are no
spiracles anterior to the abdominal portion of the body in the
female. ' ' This is corrected by Florence. She makes the following
statement: "The insects have ten pairs of spiracles, two pairs
on the thorax and eight pairs on the abdomen. Those on the
abdomen have large simple openings, and within the body form
stigmatal tubes which are anterior constrictions in which there
are one or two rings of pores according to the instar. The thoracic
spiracles have no stigmatal tubes. In the active stages their
openings are marked by a small group of pores, and in the apodous
stages these pores are wanting." The thoracic spiracles are evi-
SUBFAMILY XYLOCOCCINAE 85
dently inconspicuous and were overlooked by Pergande. In the
female nymphs of the first stage the respiratory system consists of
a very short unconnected spiracular trachea near each spij:^cle.
Each of the tracheae is strongly constricted near the middle of its
length. These constrictions contain one or more rings of cerores.
With the growth and development of the nymph the spiracular
tracheae elongate and fork. The branches of the forks elongate,
the adjacent forks curve toward each other and finally fuse
forming a continuous system of longitudinal tracheae on each side
of the abdomen. Pergande in his description of the male nymph
of the third stage states: "pores and stigmatic tubes similar tc
those of the female in the second stage.'*
This subfamily contains in addition to the genus Xylococcus,
three other genera: Coelostomidia containing four Australian
species described by Maskell with zealandica as type; Ultra-
coelostoma containing a single Australian species; and Crypto-
kermes containing a single Brazilian species.
The species of the above named genera agree so far as known
with those of Xylococcus in that they are either provided with a
chitinized rectal tube, excrete a long slender glassy tube of wax,
or their spiracular tracheae are provided with rings of cerores.
They differ, however, in the general shape of the body, which is
not always deeply imbedded in the tissue of the host -plant but in
some species is encased in a test of hard or comparatively hard wax
and attached to the surface of the bark or only slightly imbedded.
The legs and the antennae, although greatly reduced in size and
inconspicuous, are present in certain species and the thoracic
spiracles are large and prominent and the number of pairs of
abdominal spiracles is usually less than in Xylococcus. The genera
referred to this subfamily can be separated as follows : —
GENERA OF XYLOCOCCINAE
a. Adult female with nine-segmented antennae and with well developed
legs; thoracic spiracles small and inconspicuous, at least in early
nymphal stages, smaller than abdominal spiracles; abdomen with
eight pairs of spiracles. Xylococcus Loew.
aa. Adult female with more or less than nine segments in antennae, but
not with nine; legs wanting or Avell developed; thoracic spiracles
large and conspicuous, always larger than abdominal spiracles; ab-
domen with six or seven pairs of spiracles,
b. Antennae of adult female always distinct, consisting of five or
more segments; abdomen, so far as known, with six pairs of
spiracles.
86 THE COCCIDAE
c. Antennae of adult female consisting of ten or eleven segments;
second nymphal female with rudimentary legs; adult female
with well developed legs; rostrum of nymphs consisting of two
or three segments. Coelostomidia Ckll.
CO. Antennae of adult female consisting of five or six segments;
second nymphal female and adult female without legs; rostrum
of nymphs consisting of three segments. .-Ultracoelostoma Ckll.
bb. Antennae of adult female wanting or represented by mere tu-
bercles; legs greatly reduced, represented by small tubercles with
large serrated claws; abdomen with seven pairs of spiracles;
antennae of second nymphal female short six-segmented tubercles,
legs tuberculate with minute claws; rostrum of nymphal females
consisting of two segments. Cryptokermes Hemp.
Xylococcus Loew. — Four American species of Xylococcus have
been described. These can be separated by the following table based
upon that of Florence. Two species only are recognized by Ferris. The
species on alnus is considered the same as the species on birch which
is also recorded from beech. He records betulae Pergande from Ontario,
Canada, New York, and Michigan.
SPECIES OF XYLOCOCCUS
a. Abdomen of first nymph with caudal portion of ventral aspect with
single mesal pore; rectal tube of apodous nymphal stages of female
with single ring of anacerores. — California on Cupressus
macrocarpae Coleman.
aa. Abdomen of first nymph with caudal portion of ventral aspect with
more than one mesal pore; rectal tube of apodous nymphal stages of
female with two groups of anacerores, cephalic and caudal group.
b. Rectal tube of apodous nymphal stages of female with only single
ring of anacerores in caudal group; abdomen in first nymphal
stage with caudal portion of ventral aspect with mesal row of
five pores. — Lake Superior Region on Betula 'betulae Pergande.
bb. Rectal tube of apodous nymphal stages of female with more than
one ring of anacerores in caudal group.
c. Rectal tube of apodous nymphal stages of female with two
rings of anacerores in caudal group; abdomen in first nymphal
stage with caudal portion of ventral aspect with mesal row of
two to four pores. — California on Quercus. --quercus Ehrhorn.
cc. Rectal tube of apodous nymphal stages of female with three
rings of anacerores in caudal group; abdomen in first nymphal
stage with caudal portion of ventral aspect with mesal row of
six or seven pores. — California on Alnus alni Florence.
Coelostomidea Ckll. — This genus contains four species, all from
New Zealand: — compresses Mask, on Podocarpus; pilosa Mask, on Podo-
carpus and Fagus; wairoensis Mask, on Phormium and Leptospermum;
and zealandica Mask, on Muhlenbeckia and Rhipogonum. The species of
this genus have a similar structure of the rectum and execrete a tube of
wax like the species of Xylococcus.
SUBFAMILY XYLOCOCCINAE 87
Ultracoelostoma Ckll. — A monotypic genus from New Zealand
on Fagus and Phyllocladus containing the species assimilis Mask. This
species execretes a slender white filament of wax from three rings of
anacerores.
Cryptokennes Hemp. — This genus was described from Brazil on
the limbs and trunks of Schinus. The adult female and second stage
nymph of the single species, brazilensis Hemp., has the rectum chitinized
so as to form a rectal tube from which in the second nymphal stage there
is excreted a stiff tube of wax through which honeydew is passed. The
mouth-parts are reported as wanting by Ferris in the adult female and
as present in all the nymphal stages. The abdoment is provided with
six instead of seven pairs of spiracles in all stages. The legs are slender
and typical of first stage monophleblds, short and stout, femur and tibia
nearly as broad as long, in last nymphal stage; and wanting in the adult
female. The caudal end of the first nymph bears four pairs of long anal
chalazae. The antennae are clavate and consist of six segments in the
first nymphal stage; are short and stout in the last nymphal stage, con-
sist of six segments, the segments three or four times as broad as long;
and wanting in the adult female. The exuvia of the last nymphal stage
serves as a puparium for enclosing the adult female.
CHAPTER VIII
SUBFAMILY MARGARODINAE
The Ground Pearls
The body of the adult female is rotund or strongly convex on the
dorsal and flat on the ventral aspect. The cephalic, caudal, and lateral
margins are broadly convex. The segementation is indicated. The cau-
dal end of the body is not inflated and the cephalic and lateral margins
are not provided with a keel. The antennae are small, consist of six to
nine segments, and are articulated to the ventral aspect of the head near*
each lateral margin. The eyes are wanting. The mouth-parts, rostrum
and rostralis, are wanting. The legs are present and always different
in form. The prothoracic legs are fitted for digging. The profemora
are greatly enlarged and swollen, their tibiae are short, their tarsi and
claws are fused and together are much longer than their tibiae. The
mesothoracic and metathoracic legs are normal in form, their femora
are not swollen and their tarsi and claws are not fused. The claws are
as long as or longer than their tibiae. The trochanto-femoral and the
tibio-tarsal sutures are always distinct. The spiracles of the mesothorax
and metathorax are normal in form and ventral in position. The meso-
thoracic spiracles are never located near the anus. The spiracles of
the first abdominal segment are ventral in position and those of segments
two to seven or eight are usually dorsal. The spiracular tracheae of the
abdominal spiracles are never provided with one or more rings of
cerores. The stigmatic clefts, spinae, and canellae are wanting. The
abdomen is about as long as the head and thorax together. It is convex
on the dorsal and ventral aspects, never depressed. The lateral mar-
gins are uniformly convex. All of the segments are fully exposed and
subsimilar in appearance, none are retracted to form a marsupium.
The abdomen is never provided with an anal cleft and opercula, an
anal ring and anal ring setae, or anal lobes and anal setae. The caudal
abdominal segment is never short and narrow and extending beyond
the other segments. The body is provided with cerores, none of which
are octacerores or pilacerores, and never provided with ceratubae. The
body is never enclosed in a resinous cell with three adjacent openings
or covered by a scale with or without exuviae, but may' be more or less
covered with a white mealy or powdery wax or loose woolly threads of
wax. The rectum is never provided with a long glassy tube of wax with
a chitinized rectal tube bearing rings of anacerores. The body is never
naked and gall-like in form.
The female nymphs, except the last nymphal stage, are provided
with a one-segmented rostrum. The three pairs of legs or at least the
prothoracic pair are present in the first nymphal stage and dissimilar in
form. All of the legs are wanting in one or probably more nymphal
stages. The profemora, when present, are enlarged and the prothoracic
SUBFAMILY MARGARODINAE 89
legs are fitted for digging. The body is never provided with an anal
cleft and opercula, an anal ring and anal ring setae, anal lobes and anal
setae, octacerores, pilacerores, or ceratubae. The caudal end of the rec-
tum is not chitinized, forming a rectal tube provided with one_or^more
rings of anacerores excreting a long glassy tube of wax. The caudal
abdominal segment is not short, narrow, and projecting.
The male has compound eyes and may have an ocellus along the
caudal margin of each compound eye. The abdomen is not provided with
lateral filaments. The caudal end of the dorsal aspect of the abdomen
bears two tufts of long glassy filaments of wax. The stylus is apparently
wanting.
This subfamily is best known from its typical genus,
Margarodes. This was erected in 1828 by Guilding to include some
insects found by him in the soil and in the nests of ants on the
island of Bahama. The species, because of its association with
ants, was given the name of formicarum. The cysts of the insects,
known as ground pearls, occur in the soil in prodigous quantities
and show in great number when the soil is plowed. Riley states
that in certain parts of Florida they compose over one-half of the
soil. In the Bahamas they were known according to Guilding as
ant-eggs and were strung into necklaces and manufactured into
various fancy articles and used for ornamental purposes long
before they were recognized as one of the stages of an insect. A
similar use for ground pearls has been reported by other writers,
who have seen ornaments made by the use of ground pearls as
beads from South Africa and Australia, two South African species
have been described since but the Australian species is still
unknown. Guilding mistook the filaments of wax excreted from
the openings in the cyst over the thoracic and abdominal spiracles
for special organs developed for obtaining moisture by capillary
attraction and named them siphones.
A complete detailed account of the serial development of any
species of Margarodes from the egg to the adult has not been
made. The number and form of the various nymphal stages of
the female has not been determined and practically nothing is
known as to the development of the male. The following account
is based upon that of Green which is the most complete published
thus far.
The eggs are deposited in the ground in a mass of filaments
of wax excreted by the adult female. The nymphs of the first
stage are typical in form and usually provided with three pairs of
legs. This stage in one species, mediterraneus, has only the pro-
thoracic legs present. The antennae in the first nymphal stage
90 THE COCCIDAE
normally consist of six segments, but there are only three present
in the species named. The antennae of this stage of this species
are peculiar in form for coccids. The proximal and the distal
segments are, so far as shown, greatly swollen, much larger than
the intermediate segments, and the distal segment is distinctly
truncate. The greatly swollen prothoracic legs, which are fitted
for digging, are characteristic not only of the first nymphal stage
but of all the stages where the legs are present. The comparative
size and length of each prof emur are always much greater than the
combined tibia and tarsus and claw. The nymph attaches itself
to the root of a plant by its rostralis and feeds upon the sap. It is
quite likely that the sexes are, as among other coccids, indis-
tinguishable in the first nymphal stage.
Green states that the nymphs apparently undergo several
stages. Since he describes the individuals of the first nymphal
stage as larvae, this would be in addition to these. The insects,
while growing, according to Mayet, excrete the envelope or cyst
that subsequently encloses the nymph. It is this so called cyst,
because of its iridescence and wonderful pearly lustre, that has
attracted the attention of observers and that has received the
common name of Ground Pearl. Writers frequently speak of this
stage of the insect as an encysted one, but this is not a true charac-
terization as this term is ordinarily used in zoology. The insect
is simply enclosed in a compact thin-walled case or scale which
bears the same relation to the nymph as the cocoon to the pupa or
as the wax or scale of other coccids does to the insect. This case
is formed either entirely from wax excreted by the insect, entirely
from the cuticle molted by the insect, or from a combination of the
wax excreted by the insect and its molted cuticle. The cysts,
when fully developed, are more or less irregular in form and
composed of closely applied small overlapping flakes or plates.
The overlapping of the plates gives the cyst the appearance of a
minute pine-cone or swollen bud. The form and structure of the
cyst would suggest that it is formed entirely of wax. The cysts
vary considerable in color. What are apparently the young cysts
are pale yellow, becoming darker with age until they are deep
bronze brown in the oldest individuals. The amount of lustre
also varies with the age of the cyst and as to whether the outer
plates, which are usually dull, are wanting or not.
The various accounts would seem to carry the idea that the
cyst was begun during the first nymphal stage. The female nymph
at the first molt loses its legs and eyes. The antennae are repre-
SUBFAMILY MARGARODINAE 91
sented by ringed disks bearing one or two curved setae. There is
some disagreement as to the disappearance of the rostrum and
rostralis. Giard maintains that they exist and are functional in
all the nymphal stages but are small and inconspicuous and that
they are not lost until the assumption of the adult condition.
The later observations of Green would seem to confirm this conten-
tion of Giard.
Green has figured and described a stage of niger, which was
obtained from small scales, which differs from all of the stages
previously described. It is oval in outline and lacks legs. The
antennae are represented by small tubercles with a pit bearing two
curved setae. The mouth-parts are well developed. The abdo-
men bears seven pairs of spiracles instead of eight as in the adult.
The sterna of the abdominal segments bear ^Ye longitudinal rows
of small circular glandular pits or cerores. A still earlier stage
was observed. These may represent stages in the development of
the male which is unknown.
The female Margarodes may remain apparently for a consid-
erable time, several years, within their cyst if the conditions for
their emergence are unfavorable. She eventually breaks through
the wall of the scale and emerges as an insect that is normal in
form at least for a coccid. The body is globular. The antennae
are short, gradually tapering toward the distal end, varying with
the species from seven to nine segments. The mouth-parts, including
the rostrum and rostralis, are wanting. The prothoracic legs are
greatly enlarged, the profemora are thickened and fitted for dig-
ging. The other two pairs of legs are normal in form. The body
may be shaggy with fine short setae or nearly glabrous. The
dorsal and ventral aspects may bear short conical spines, the
dermal spines of Green. The lateral margins of the body and the
dorsal and ventral aspects may bear transverse groups of cerores
with intermixed setae. The body becomes more or less covered
with a white mealy or powdery wax or a loose woolly wax. The
thoracic spiracles and those of the first abdominal segment open on
the ventral aspect, while the remaining abdominal spiracles open
on the dorsal aspect or on the lateral aspect as in mediterraneus.
The thoracic spiracles are much larger than the abdominal. There
are cerores usually associated with both the thoracic and the
abdominal spiracles. The anus and vulva are located on the
ventral aspect. The former is small and surrounded by a
chitinous plate. The adult females vary greatly in size. Green
reports in the case of papillosus a variation of 1.14 mm. to 4 mm.
92 THE COCCIDAE
and in the case of niger 2 to 8 mm. and quotes the following from
Giard: *' according to the conditions of nutrition in which the
nymph finds itself, the latter is able to transform itself into an
adult female of very different sizes. Certain adult females of
M. vitium are no more than 2 mm., while others attain a length of
5 or even 8 mm. As a result one often finds adult females smaller
than some of the larvae."
The development of the male is unknown. Green assumed
from collectors' notes that the males become encysted like the
females, but their cysts have not been identified. The collectors
reported the cysts "were fairly numerous and were obtained five
to seven inches beneath the soil. The males were also enclosed in
shells but emerged soon after excavating and were observed
copulating toward evening." The adult males and females
emerged in June in papilliosus and the cysts .were reddish in
color. In the case of niger the collector makes the following note :
"Egg-masses of this Ground Pearl were observed in the soil at a
depth of 2-3 inches. The cells in which the eggs were found were
long-oval, lined inside with a coating of mealy wax. Hundreds
of eggs were found filling up the interior of each cell. The dead
mother scale — shrivelled and rotten — was to be found at one pole
of the cell."
The legs of the adult male are similar to those of the female,
the prothoracic legs which are used for digging into the soil in
search of the female, are enlarged. The tibio-tarsal sutures are
usually present but may be wanting. The claws of the mesotho-
racic and metathoracic legs are simple, rarely trifid. The antennae
are long and slender and consist of ten to thirteen segments.
They are ordinarily simple but may be branched or flabellate.
The wings with numerous conspicuous veins show their affinity tc
the wings of the generalized Gularostria. The halteres are small
and bear a single curved hook.
The two genera of Margarodinae, of which one is known only
in the male sex, can be separated by means of the following table : —
GENERA OF MARGARODINAE
a. Male with antennae simple and with tibio-tarsal sutures of meso-
thoracic and metathoracic legs present; adult female with abdomi-
nal spiracles, those of first abdominal segment located on ventral
aspect and those of other segments on dorsal aspect, rarely lateral
aspect Margarodes Gnilding.
aa. Male with antennae distinctly flabbellate and Avith tibio-tarsal
sutures of m-esothoracic and metathoracic legs obsolete; adult female
unknown. Ncomargarocles Green.
SUBFAMILY MiVRGARODINAE 93
Neomargarodes Green. — The genus Neomargarodes contains a
single species, erythrocepliala Green, based upon males collected in Al-
geria.
Margarodes Guilding. — A part of the species of Margarodes have
been based upon characters drawn from the scale alone. Such charac-
ters have been shown to be worthless and consequently the characters
necessary for the differentiation of the fourteen recognized species have
not been indicated. The following characters drawn in the main from
the papers of Green, who figures a prothoracic leg of six of the species,
may be found useful in separating the species. There are four European
species, gallica Signoret from France with nine segmented antennae,
polonicus Linnaeus from southern Europe on Scleranthus with eight
segmented antennae, perrisii Signoret from France with seven segment-
ed antennae, and mediterraneus Silvestri from Italy on Cynodon with
seven segmented antennae. This latter species has the profemur as long
as its tibia and tarsus and claw together and the ventral surface of the
profemur and the surface of the protibia bears several large setae. A
species, hameli Brandt, from the borderland between Europe and Asia,
in Armenia on Poa, has nine segmented antennae. The two South
African species, trimeni Giard and capensis Giard, from Cape Colony,
probably from the nests of termites, are easily separated. The pro-
femur is shorter than the tibia and tarsus and claw together in trimenii
and the ventral surface of the profemur bears only a few small scattered
setae while the protibia bears a few more arranged in a single trans-
serse row. In capensis the profemur is longer than its tibia and tarsus
and claw together and the ventral surface of the profemur bears sev-
eral minute setae while the protibia is glabrous. There are three species
described from India by Green, indicus of which the antenna is unknown
and in which the profemur is longer than its tibia and tarsus and claw
together and in which the profemur bears four or five small setae on
the ventral aspect and its protibia bears two setae on each side; papil-
losus has seven segmented antennae and its profemur is subequal in
length to the combined tibia, tarsus, and claw and bears several small
setae on the ventral aspect of the profemur and four or five setae on the
protibia; and niger which differs from all of the other species so far
as known in having the profemur three or four times as long as its
tibia and tarsus and claw together, this latter combined structure is in
the form of a strongly chitinized toothed claw. The South American
species, vitium Giard from Chili on Vitus, has eight segmented antennae.
The same number is present in hiemalis Cockerell from New Mexico.
Of the two West Indian species, formicarum Guilding from Antigua and
Bahama Islands from the nests of ants, has seven segmented antennae
and the profemur is much longer than its combined tibia and tarsus
and claw and bears numerous long large setae on the ventral aspect
of the profemur and one on each side of the protibia, while the form
of the prothoracic legs and the antennae of rileyi Giard from West
Indies and Florida Keys is unknown. The figures of Green suggest that
good diagnostic characters are to be found in the prothoracic legs and
it is to be hoped that those having authentic specimens of the other
species will publish figures of these structures.
CHAPTER IX
SUBFAMILY CALLIPAPPINAE
The Marsupial Coccids
The body of the adult female, as seen from the lateral aspect, is
strongly inflated. The cephalic portion is thin and bluntly pointed, the
caudal portion is much inflated or swollen and truncated. As viewed
from above, the body is more or less triangular in outline, with a de-
cided keel extending around the margin of the body, except across the
truncated caudal portion. The segmentation is distinctly indicated by
notches in the lateral rim or keel. The antennae are prominent, usu-
ally consist of ten segments, rarely eleven, and are articulated to the
ventral aspect of the head near each lateral margin. The eyes are pres-
ent. There is a single globular ocellana located on each lateral margin
caudad of the articulation of an antenna. The mouth-parts, rostrum
and rostralis, are always wanting. The legs are present. They are
long and stout, subequal in length, and normal in form. The profemora
are never enlarged and the prothoracic legs are never fitted for digging.
The trochanto-femoral and tibio-tarsal sutures of all the legs are always
distinct. The thoracic and abdominal spiracles are not definitely de-
scribed and are not figured. The stigmatic clefts, spinae, and canellae
are wanting. The abdomen is deeply invaginated or retracted and only
the cephalic two or three segments are exposed, so that the head and
thorax together are several times as long as the abdomen. It is strongly
convex on the dorsal and ventral aspects and is never depressed. The
lateral margins bear a rim and the caudal aspect is broadly truncated
and bears a transverse slit, the entrance to the invaginated marsupium.
The abdomen is never provided with an anal cleft and opercula, an anal
ring and anal ring setae, or anal lobes and anal setae. The caudal
abdominal segment is not short and narrow and extending beyond the
other segments but is concealed in the marsupium. The body is sparse-
ly provided with cerores none of which are octacerores or pilacerores
and is never provided with ceratubae. The body is never enclosed in a
resinous cell with three adjacent openings or covered by a scale with or
without exuviae, but may be covered with a thin coating of powdery
wax and sometimes with a pad of wax between the sternum of the
thorax and the host-plant. The rectum is never provided with a long
glassy tube of wax or with a chitinized tube bearing rings of anacerores.
The body is never naked and gall-like in form.
The female nymphs at least of the first stage are provided with a
rostrum and rostralis. The three pairs of legs are always present in
all nymphal stages so far as known and are similar in form. The profem-
ora are never greatly enlarged and the prothoracic legs are never fitted
94
SUBFAMILY CALLIPAPPINAE 95
for digging. The body is never provided with an anal cleft and opercula,
an anal ring and anal ring setae, anal lobes and anal setae, octacerores,
pilacerores, or ceratubae. The caudal end of the rectum is not chitin-
ized forming a rectal tube provided with one or more rings of anacerores
excreting a long glassy tube of wax. The caudal abdominal segmenf is
not short, narrow, and projecting.
The male has large finely faceted compound eyes. The ocelli are
wanting. The abdomen is not provided with long lateral filaments but
bears tufts of long wax filaments. The stylus is minute or concealed.
The name of the single genus, Callipappus, belonging to this
subfamily, is derived from the two Greek words, kallos meaning
beauty and pappos meaning grandfather, down, pappus, a floating
seed. This undoubtedly refers to the resemblance of the male
when on the wing to the floating cottony thistle-like seeds of
certain plants. The beauty of the male is further shown in the
vernacular name of one of the common species. The Bird of
Paradise Coccid.
The females of this subfamily are among the largest of all the
coccids. While specimens half an inch long are of common
occurrence, others three times this size or an inch and a half or
about thirty-five millimeters in length are not unusual. Individ-
uals an inch in length are normal in size for certain species.
The most striking peculiarity in the female is the invaginatiop
or intussusception of the caudal portion of the abdomen. In the
adult female all the segments but the first and second and in
some individuals also the third are invaginated to form a pocket,
pouch, or marsupium. This pouch extends almost to the cephalic
end of the body. Since the caudal end of the body is simply
pushed in, as it were, the vulva and anus retain their position in
the invaginated part and are located on the inside of the cephalic
end of the marsupium. The eggs are deposited within the
marsupium and their development takes place there. The young,
when they hatch, escape through a transverse slit in the caudal
end of the pouch, the mouth of the invagination. This slit is
guarded by lips which are apparently different in form in the
different species. The genus Callipappus has been variously asso-
ciated with Monophlebus, Icerya, Margarodes, and Xylococcus.
from all of which it is structurally very different through the
development of its marsupium and is regarded as worthy of
consideration as a distinct subfamily.
The only condition allying Callipappus with Margarodes and
Xylococcus is the lack of a rostrum and rostralis in the adult
96 THE COCCIDAE
female. The following quotation from Tapper shows how easily
it is for one to be misled by appearances and how unwise it is tc
base statements regarding anatomical structures in this family on
other than prepared specimens. ''Mr. Maskell placed the species
(immanis Maskell) in the genus Coelostoma 'on account of the
absence of the rostrum' (op. cit. p. 51) and says 'there is no doubt
of its being a monophlebid, ' for both Monophlebus crawfordi and
Coelostoma australe are known to possess a rostrum (sucking
tube) of great length, which during life is deeply buried in the
tissues of the bark of the twigs on which the insect is located,
when, however, the organ is withdrawn it is at the same time
retracted into the body; its extreme slenderness and the minute-
ness of the otherwise unmarked orifice rendering it next to
impossible to detect its presence. Mr. A. Zietz informs me that
he has frequently detached living Monophlebi from the bark and
seen their rostrum retracted, and I have noticed in the present
instance that the living insect (Coelostoma) had its body closely
and immovably fixed to the twig by its rostrum, but later, when
dead, had its forepart raised and the rostrum completely retracted.
The absence of a rostrum, as a distinction, will, therefore, have to be
omitted in future." It is true that living Icerya purchasii are
able to conceal the rostrum under folds of the ventral aspect of the
head and thorax when removed from the host-plant, but an
examination of a specimen clarified in caustic potash will show
that anatomically it is impossible for the insect to retract its
mouth-parts and the only absolutely sure method of determining
whether the mouth-parts are present or absent in a given species
or specimen is the study of a properly prepared specimen.
The adult female is triangular in outline as viewed from above.
The cuticle of the body contains circular areas with minute central
projections, these will probably prove to be cerores. The
cephalic end is pointed and the long tapering antennae of ten
or eleven segments are articulated one on each side near the
cephalic end of the pointed portion. The caudal end is broad and
greatly swollen. The insect as viewed from either side is pointed
at the cephalic end and gradually swollen until it reaches its
greatest expansion at the caudal end. There is a distinct rim or
margin which extends around the pointed cephalic end and along
the middle of each lateral aspect to the large transverse truncated
caudal end, where the rim is interrupted by the transverse slit
leading into the marsupium. The legs are long and slender and
SUBFAMILY CALLIPAPPINAE 97
normal in form. They are articulated to the three large thoracic
segments about equidistant from each other and the two ends of
the body. The segmentation of the thorax and exposed abdominal
segments is distinctly marked by indentations in the lateral rim.
The female, after depositing her eggs in the marsupium, is very
different in appearance and it is this form with the swollen
abdomen that is usually described. There is a pad of wax excreted
from the ventral aspect of the thorax. This wax attaches the
female firmly to the bark of the tree upon which she is resting.
Her legs are spread out around the twig or leaf in such a way
that she has the appearance as Maskell states of apparently being
afraid that she will fall off from the twig. The surface of the
body is covered with white mealy wax with which is mixed some
flakes of wax and some short cottony threads of wax.
The young female is figured by Signoret. The antennae
consist, according to this author and Fuller, of six segments. The
body has the lateral margin broadly convex and the two ends are
bluntly pointed. The abdomen is represented as containing the
full number of segments. The insect is figured as if in this stage
it was distinctly coccid-like in form, which the adult females are
not. The thoracic spiracles are figured by Signoret. The only
reference to the spiracles that has been found is the following
statement of Maskell dealing with a nymph of immanis. ''In this
specimen there is a very small quantity of whitish cotton under the
anal region, and along the margin ; just below what I have called a
seam on the edge, are ten small circular specks of white, (five on
each side), which may mark the position of the spiracles." It is
probable that these represent the thoracic spiracles and a pair for
each of the three exposed abdominal segments. It is striking that
with insects so large in size, so bazarre in their structure, and so
peculiar in their habits that no one has made a careful detailed
study of one or all of the species. Even when and how the
marsupium is formed has not been described. The invagination
apparently takes place after the last molt.
The adult male, which is about a quarter of an inch long, has
antennae of ten or eleven segments. The eyes are compound.
The ocelli are not mentioned in descriptions but are probably
present. The abdomen is distinctly segmented and without lateral
projections and the caudal segment bears a large brush of long
glassy filamentous threads of wax which are often spread out fan-
shaped, in some individuals three times as long as the body,
98 TflE COCCIDAE
resembling the plumes of a bird of paradise. These wax filaments
do not appear for five or six days after the emergence of the male
and require two or three days before they reach their complete
development. The stylus is a short conical spike from which pro-
trudes the penis which is very long and flexible. The wings are
large with an expanse of two-thirds of an inch. The halteres are
large and wide and each is provided with a single curved hook.
The males transform in cocoons made of a mass of white
cottony threads of wax. They are placed on the ground among
leaves and under pieces of sticks and bark near the trunks of the
trees upon which the nymphs feed. The life cycle of the male or
female has not been described.
Callipappus Guerin. — This genus contains six species, all are from
Australia, the land of marsupial animals. The six described species are
as follows: — australis Maskell on Eucalyptus and Augophora; bufo
Fuller on Casuarina and Banksia; farinosus Fuller on Casuarina;
immanis Maskell on Acacia and Eucalyptus; rubiginosus Maskell on
Banksia; and westwcodii Guerin, the type of the genus, on Eucalyptus.
CHAPTER X
SUBFAMILY COCCINAE
The Cochineal Insects
The body of the adult female is broadly convex on all aspects with
the two ends bluntly rounded. It is elongate oval in outline. The seg-
mentation is distinct. The caudal end of the body is not inflated and
the cephalic and lateral margins are not provided with a keel. The
antennae are small but distinct, consist of five to seven imperfectly
separated segments, and are articulated to the ventral aspect of the head
near each lateral margin. The eyes are present. There is a single
ocellana located on each lateral margin caudad of the articulation of an
antenna. The mouth-parts, rostrum and rostralis, are always present
and distinct, the rostrum consists of two indistinct segments. The legs
are present. They are small, subequal in size and length, normal in
form, and do not project beyond the lateral margin of the body. The
profemora are never enlarged and the prothoracic legs are never fitted
for digging. The trochanto-femoral and the tibio-tarsal sutures of all
the legs are always distinct. The spiracles of the mesothorax and met-
athorax are small and ventral in position. The mesothoracic spiracles
are never located near the anus. The abdominal spiracles are always
wanting. The stigmatic clefts, spinae, and canellae are wanting. The
abdomen is not quite as long as the head and thorax together. It is
convex on the dorsal and fiat on the ventral aspect, is never depressed.
The lateral margins are convex or subparallel and the caudal end is
broadly rounded. All the segments are fully exposed and similar in
appearance with more or less distinct coriae between them, none are
retracted to form a marsupium. The abdomen is never provided with
an anal cleft and opercula, an anal ring and anal ring setae, or anal
lobes and anal setae. The caudal abdominal segment is short and nar-
row, inconspicuous, not extending beyond the other segments. The
body is provided with cerores none of which are octacerores or pila-
cerores and is never provided with ceratubae. The body is never en-
closed in a resinous cell with three adjacent openings or covered by a
scale with or without exuviae, but is either completely covered or rests
upon a thick mass of white cottony threads of wax. The rectum is not
provided with a long glassy tube of wax or with a chitinized tube bear-
ing rings of anacerores. The body is never naked and gall-like in form.
The female nymphs of all stages are provided with a rostrum and
rostralis. The three pairs of legs are present in all nymphal stages and
are similar in form. The profemora are never enlarged and the pro-
thoracic legs are never fitted for digging. The body is never provided
with an anal cleft and opercula, an anal ring and anal ring setae, anal
99
100 THE COCCIDAE
lobes and anal setae, octacerores, pilacerores, or ceratubae. The caudal
end of the rectum is not strongly chitlnized forming a rectal tube provided
with one or more rings of anacerores excreting a long glassy tube of wax.
The caudal abdominal segment is not short, narrow, and projecting.
The male has six ocellanae. The abdomen is not provided with long
lateral filaments or tufts of long glassy filaments, but bears two long,
slender, caudal filaments or threads of wax. The stylus is small and in-
conspicuous.
The most useful products for the production of permanent
shades of red, carmine, or scarlet are derived from the cochineal
insects, Coccus cacti. The dried bodies of these insects were used
by the native Mexicans when the Spaniards under Cortes arrived
in Mexico in 1518. There was much conjecture and discussion as
to the true nature of cochineal. It was considered by Acosta as
early as 1530 and later by others as an insect, but was generally
believed to be the seed of a plant and was not definitely proven
to be an insect until in 1694 by Hartsoeker and by Loeuwenhoek
in 1704, who examined specimens carefully with a microscope.
This coccid feeds upon various species of Cactaceae, more
often those of the genus Opuntia, but is said particularly to feed
upon the nopal plant, Opuntia coccinellifera, although this is
denied by W. J. Hooker. The insects and the plants upon which
they feed have been introduced and cultivated in many parts of
the world. They were introduced by the French and Spaniards
into Spain, Italy, and northern Africa, and various of the tropical
islands of the Atlantic where the Cactaceae thrive. It warj also
introduced at an early date into India. Cochineal is stated to be
produced commercially at the present time in Mexico, Peru.
Algiers, and southern Spain. The discovery and introduction of
the much cheaper but inferior and less permanent coal tar colors
has supplanted in great part the use of cochineal. According tc
Humboldt and Westwood 800,000 pounds of cochineal were annually
imported into Europe before the use of the cheaper dyes, of which
150,000 pounds, worth $1,800,000, was used in England alone.
In collecting the insects they are carefully brushed from the
surface of the plant into bags or sheets. They are killed by
immersion in hot water or by exposure to the sun or the heat from
steam or an oven. The difference in the appearance of the
commercial product is due to the way in which it is treated. The
grains or dried bodies are longitudinally folded or wrinkled or
fluted. The best crop is secured early in the season by collecting
the unimpregnated females. The specimens collected later in the
SUBFAMILY COCCINAE 101
season consist in great part of young nymphs and molted skins
which are consequently not nearly so rich in coloring matter. A
pound of the dead females contains about 70,000 individuals.
There are three grades of cochineal. The silver cochineal, which
is greyish in color in which the furrows of the body are said to be
covered or filled with a whitish bloom of fine down, is the best
grade. This bloom is undoubtedly the wax excreted by the female
after her removal from the host-plant and before her death. The
other grades are black cochineal which is dark reddish brown and
is destitute of the bloom and granilla which is an inferior grade
collected from uncultivated or wild plants.
The living adult females of the genus Coccus are between twc
and three millimeters in length. The body is elongate oval in
outline. The antennae are short and inconspicuous, the six or
seven segments consisting of short chitinized rings connected by
coriae of about the same length. They do not project beyond the
lateral margin of the body. The legs are short and stout, similai
in form. The femur of each leg is as long as its tibia and tarsus
together. The rostrum consists of two segments. The thoracic
spiracles are four in number, small, and ventral in position. The
abdominal spiracles are wanting. The anus and vulva are incon-
spicuous. All portions of the external cuticle of the body bears
bluntly truncated setae or tubercles as they are called which are
very characteristic in form. The number of these setae present
and their length and breadth varies with the species. There are
a number of minute cerores arranged in groups of varying size
scattered among the truncated setae.
The females, which are dark crimson in color, when they reach
maturity, excrete a mass of white cottony threads of wax. The
insects are either completely covered by this mass of wax or rest
upon its surface. The eggs are deposited within the mass.
The nymphs of the first stage are similar in shape to the adult
females, elongate oval in outline, and dark crimson in color. They
have long legs that project beyond the sides of the body. The
antennae, which also project beyond the margins of the body,
consist of six segments with distinct constrictions between them.
The dorsal aspect bears six rows of truncated setae, a longitudinal
row along each lateral margin and four between them. There is a
single transverse row on each abdominal segment and probably twc
for each thoracic segment and the head. The truncated setae are
wanting on the ventral aspect. The females of the second nymphal
102 THE COCCIDAE
stage differ from those of the first nymphal stage not only in the
presence of a large number of truncated setae but in their larger
size. The individuals of this stage are very similar to the adult
females.
The nymphs of the males form cylindrical cocoons with equally
rounded ends of thick felted white wax. The adult male is dark
crimson in color. The antennae consist of ten segments, each
segment bearing short blunt tubercular setae. The caudal end of
the abdomen bears two long very slender filaments of wax. The
most striking feature is, as first pointed out by Newstead, the
presence of two segments in the tarsus of each leg. The distal or
second segment is as long as in the tarsi of all other coccids, but
in addition to this segment, there is a minute segment, longer on
the ventral aspect, between the second segment and the tibia.
There seems to be a difference of opinion as to what name
shall be applied to the cochineal insects. Comstock and his predi-
cessors called it Coccus cacti Linnaeus, but Cockerell states that
some of the works cited by Linnaeus, as Reaumur, relate to the
true cochineal insect, "but the whole of the Linnean description
pertains to a monophlebid. The specimens described were col-
lected in the island of St. Eustache by Daniel Rolander, and sent
alive to Upsala in 1856. At the same time Rolander sent a number
in alcohol to De Geer, who (Mem., vol. VI, p. 449) gave a full
account of them with a figure." There is nothing in the account
of De Geer to show that the specimens he received were a part of
the collection sent to Linnaeus, whether it was sent at the same
time and whether it was the same species or not is not clear. De
Geer makes the following statement showing the source of his
knowledge as to the Linnean specimens: "comme le rapporte M.
de Linne. " The evidence that the Coccus cacti of Linnaeus is not
one of the cochineal insects is not positive and so long as it can not
be proven that it is not, less confusion will result from the appli-
cation of the generic name Coccus to the cochineal insects and the
application of the name cacti to one of the species as has been
done on the following pages. There are two genera referred tc
this subfamily. It is considered that only one of these, Coccus, in
reality belongs here. The other genus is placed here tentatively.
They can be separated by means of the following table : —
GENERA OF COCCINAE
a. Body of adult female bearing varying number of short truncate
setae; adult female and nymphal females without any trace of anal
SUBFAMILY COCCINAE 103
lobes; antennae typically with seven segments, sometimes with six,
some of intermediate segments longer than proximal or distal seg-
ments; rostrum consisting of two inconspicuous segments. —
Coccus Linnaeiis.
aa. Body of adult female bearing pointed setae, but never bearing short
truncate setae; antennae always with six segments, always with
distal segment longer than others; adult female with small incon-
spicuous microscopic anal lobes; rostrum not described
Epicoccus Ckll.
Coccus liinnaeus. — The following table based upon the descriptions
and figures of Green will serve for the separation of the species: —
SPECIES OF coccus
a. Body with truncate setae few in number, long and slender in form,
width of their distal ends about one-fourth their length; cerores
large with thick rims, arranged in clusters, each cluster consisting
of two to twenty cerores, average twelve or thirteen; eyes cylin-
drical, longer than broad; body pruinose, without conspicuous to-
mentum. — Mexico, Canary Islands, Algeria, Spain, West Indies,
Maderia, Peru, Florida, California, Guatamala, Italy, Cape Colony,
India on Opuntia. cacti Linn.
aa. Body with truncate setae numerous and comparatively large, dis-
tinctly truncate, the width of the distal end about one-half their
length; cerores small and without thick rims; eyes round or broad-
er than long; body conspicuously covered with white tomentum.
b. Antennae with distal segment subglobular, as broad as long;
cerores small and inconspicuous, usually in clusters of three or
four, rarely five to six; truncate setae stout, with parallel sides
and distal end distinctly narrower than diverging sides of proxi-
mal half. — India, Ceylon, California, New York, Argentina on
Opuntia. indicus Green.
bb. Antennae with distal segment longer than broad; cerores ar-
ranged in comparatively large clusters; truncated setae not as
above.
c. Antennae and legs very small, trochanter and femur .12 to
.16 mm. in length; antennal segments much contracted, total
length .10 to .11 mm.; cerores small and inconspicuous, in
clusters of two to sixteen, average about sixteen; truncate
setae numerous and conspicuous, variable in form, ranging
from quite slender to short and broad and cylindrical. — Mis-
souri, Arizona, New Mexico, California, Mexico, England on
Opuntia. tomentosus Lamar.
cc. Antennae and legs larger, trochanter and femur .18 to .23 mm.
in length; antennal segments compartively broad and extend-
ed, total length .15 to .18 mm.
d. Truncate setae elongate, cylindrical, slightly expanded at
proximal end, diameter of proximal end one-half or slightly
more than total length of the seta.
104 THE COCCIDAE
e. Cerores small, in clusters of three to thirty-six, average
fifteen; length of body 3.5 to 4.75 mm. — Texas, Arizona,
New Mexico, Montana, Mexico on Opuntia confusus Ckll.
ee. Cerores small, in clusters of four to thirty, average thir-
teen; length of body 2.5 to 3.0 mm. — Colorado, Texas,
Arizona on Opuntia. newsteadii Ckll.
dd. Truncate setae stout, shorter, cylindrical, diameter of prox-
imal end usually more than one-half the total length of
seta; cerores in clusters of one to thirty, average fifteen;
length of body 2.5 to 3.5 mm. — Cape Colony, South Africa on
Opuntia. capensis Green.
Epicoccus Ckll. — This genus contains a single species, acaciae
Maskell from West Australia on Acacia. The species appears to be
similar to those of the genus Coccus but lacks the truncate setae. The
absence of an anal ring with anal ring setae in all stages excludes it
from the Eriococcinae and the presence of legs in the adult female from
the Cylindrococcinae. The highly colored naked body, according to
Fuller one of the most beautiful of the Australian coccids, the body-wall
bearing only a few cerores, and the six segmented antennae in the adult
female and first nymphal stage are characteristic.
CHAPTER XI
SUBFAMILY ORTHEZIINAE
The Ensign Coccids
The body of the adult female is rotund, convex on all aspects. It is
elongate oval in outline, broadest in the metathoracic region. The lat-
eral margins are convex and the cephalic and caudal margins are blunt-
ly rounded. The segments are well defined. The caudal end of the body
is not inflated and the cephalic and lateral margins are not provided
with a keel. The antennae are prominent, consist of four to nine seg-
ments, and are articulated to the ventral aspect of the head near each
lateral margin. The eyes are present. There is a single ocellana located
on a tubercle on each lateral margin caudad of the articulation of an
antenna. The mouth-parts, rostrum and rostralis, are always present,
the rostrum consisting of two distinct segments. The legs are present.
They are subequal in length and normal in form. The profemora are
never enlarged and the prothoracic legs are never fitted for digging.
The trochanto-femoral and the tibio-tarsal sutures are frequently want-
ing. The spiracles of the mesothorax and metathorax are normal in
size and ventral in position. The mesothoracic spiracles are never lo-
cated near the anus. The abdominal spiracles are frequently distinct
and located on segments one to seven. They are smaller than the tho-
racic spiracles, ventral or subventral in position, and their spiracular
tracheae are not provided with rings of cerores. The stigmatic clefts,
spinae, and canellae are wanting. The abdomen is distinctly shorter
than the head and thorax together. It is convex on the dorsal and fiat
on the ventral aspect, never depressed. The lateral margins are convex
and the caudal end is broadly rounded. All of the segments are fully
exposed and similar in appearance with distinct coreae between them,
none are retracted to form a marsupium. The abdomen is pever pro-
vided with an anal cleft and opercula or anal lobes and anal setae. The
anal ring is always distinct and provided with six anal ring setae. The
caudal abdominal segment is short and narrow, small and inconspicuous,
but normal in form and not extending beyond the other segments. The
body is provided with cerores none of which are octacerores, but the
great majority are pilacerores. Ceratubae are always wanting. The
body is never enclosed in a resinous cell with three adjacent openings
or covered by a scale with or without exuviae, but is always in great
part covered by a mass of hard white wax which is fused together and
forms distinct plates or lamellae, a part forming a marsupium which is
carried by the female. The rectum is never provided with a long glassy
tube of wax or with a chitinized rectal tube bearing rings of anacerores.
The body is never naked and gall-like in form.
105
106 THE COCCIDAE
The female nymphs of all stages are provided with a rostrum and a
rostralis. The three pairs of legs are always present in all nymphal
stages and are similar in form. The profemora are never greatly en-
larged and the prothoracic legs are never fitted for digging. The body
is never provided with an anal cleft and opercula, anal lobes and anal
setae, octacerores or ceratubae. The anal ring is distinct and bears six
anal ring setae. The caudal end of the rectum is not chitinized, form-
ing a rectal tube provided with one or more rings of anacerores excret-
ing a long glassy tube of wax. The cuticle bears numerous pilacerores.
The caudal abdominal segment is short and narrow but not projecting.
The male is provided with compound eyes and sometimes with three
ocelli. The abdomen is not provided with long lateral filaments, but
bears a delicate pencil of long slender threads of wax. The stylus is
small.
The species of Orthezia are among if not the most beautiful
of all coceids. Their beauty depends upon the size and form of
the marsupium of the female and upon the size, complexity, and
arrangement of the plates or lamellae of wax associated with the
marsupium and attached to the dorsal aspect. The marsupium is
constructed of symmetrically arranged glistening white plates or
lamellae of wax, which are usually fused into a single tubular
structure which is carried by the caudal part of the abdomen of
the female. This tubular structure is attached to the abdomen
and is held in many species in a more or less elevated position, the
amount of elevation varying with the species. From this fact there
has been derived their common name of the Ensign Coceids, since
the movement of the marsupium when the insect is in motion
resembles the waving of an ensign. The marsupium is formed by
special groups of pilacerores located on the ventral aspect of the
abdomen.
The pilacerores are peculiar to this subfamily. They are
ordinarily designated as spines or glandiferous spines, but spines
are large setae and all setae are provided with calices by which
they are articulated to the cuticle. The calices are always wanting
in these projections or pilacerores. There are generally a few
minute setae with calices distributed among and between the
groups of pilacerores. Each pilaceroris is a simple slender pro-
jection of the cuticle of the body-wall with a single opening, the
oraceroris, at its distal end through which the wax is poured. That
this is their true structure is beautifully shown by the figures of
List. The pilacerores serve the double purpose of outlets for the
wax and as threads for holding the plates of wax to the body after
they have been excreted and become hard.
There are in addition to the pilacerores of the ventral aspect,
SUBFAMILY ORTHEZHNAE 107
which excrete the wax forming the marsupium, other groups of
pilacerores on the dorsal and lateral aspects of the head, thorax,
and abdomen. The number and shape of the dorsal plates of wax
is dependent upon the size and shape of these groups of pilacerores.
The tubular marsupium, which may be from twice to four or
five times as long as the body, surrounds the vulva. The eggs as
they are extruded pass into the marsupium where there is mixed
with them a few long curly threads of wax which are formed by
the ordinary cerores located on the ventral aspect among and
between the pilacerores. The adult female is active and carries
her marsupium about with her wherever she goes. The young
nymphs when they hatch, crawl out through the open end of the
marsupium and drop from this end onto the host-plant or crawl
down its side and over the body of the female onto the host-plant.
The beauty of the marsupium is due not only to its glossy white
color, but also to its longitudinal flutings, which are produced by
the irregularities or scallops in the row of pilacerores which form
it. The flutings are usually wanting on the ventral aspect of the
marsupium and vary in distinctness with the species on the dorsal
and lateral surfaces. They are sometimes only slightly indicated
The nymphs of the first stage have the body oval in outline
with the cephalic and caudal ends broadly rounded. The legs are
normal in form but are comparatively large aind the sutures
between the trochanter and femur and often also between the
tibia and the tarsus of each of the legs are frequently wanting, oi
if indicated, the segments are fused and immovable. The dorsal
and ventral surfaces of the body are provided with numerous
pilacerores and a slight covering of wax is generally formed before
the nymphs issue from the marsupium. The antennae consist oi
four to six distinctly separated segments. The anus is surrounded
by a prominent anal ring which bears six large anal ring setae.
There are numerous cerores, not pilacerores, located on the anal
ring. These cerores differ from the cerores of the anal ring of
many other coccids in not having a definite arrangement.
There are a number of nymphal stages in the female, four in
insignis, in all of which the insect is active at all times. The
nymphs resemble the adult female very closely so far as the
number and disposition of their pilacerores are concerned, but
they can be recognized always by the fact that the plates of wax
are never as large as those of the adult because they are shed at
each molt. The legs in the older nymphal stages, because of the
108 THE COCCIDAE
increase in the size of the body, do not appear so large and so out
of proportion with the remainder of the body. There is up
difference in the anal ring, the anal ring setae, and the anal ring
cerores between those of a nymph and those of the adult. The
adult antennae contains from four to nine segments, the number
varying with the genus.
While abdominal spiracles have not been found in all the
species of Orthezia, they have been found in all the species exam-
ined and it is probable that when all the species have been inspected
for abdominal spiracles they will be found to be present. The
evidence seems to point to the presence of seven pairs of abdominal
spiracles in all species. Newstead was unable to find abdominal
spiracles in Newsteadia and Ferris did not find them in Nipponor-
thezia.
The life cycle of the male has not been described. The females
of insignis frequently occur upon plants in such numbers that the
plants are white with them, but the number of males is always
very small in comparison to the number of females. It is likely,
judging from the published accounts, that the nymphs of the male
are active during the first nymphal stage, but become more elongate
during the second nymphal stage when they excrete loose woolly
fibres of wax or dense masses of white wax which are used to form
a cocoon. The insect is quiescent during the remaining nymphal
stages. The wing-pads and developing legs of the adult soon
become distinct.
The males in maensariensis have large compound eyes with
three ocelli placed on the vertex between the compound eyes.
There are probably two in the other species which are located
cephalad of the compound eyes. They are wanting in Newsteadia,
The caudal end of the abdomen bears a pencil of delicate glassy
threads of wax.
The species and the genera in part are based upon the number,
size, and form of the lamellae or plates attached to the dorsal and
lateral aspects of the body. It is unfortunate that the number
and form and arrangement of the various groupes of pilacerores
which form these lamellae have been neglected and structures that
are extremely fragile and easily broken have been used instead.
It is difficult when the various descriptions are compared tc
determine just what is meant. No one has attempted to stand-
ardize the nomenclature. The following names are suggested for
the plates. Care has been used in selecting these names so that
SUBFAMILY ORTHEZIINAE 109
they could be applied to the groups of pilacerores when these are
described and used for separating the species in place of the wax
excreted from them.
The ovisac is formed by the continuous band of pilacerores
which are located on the lateral portions of the ventral aspect.
The two sides converge toward the caudal end of the abdomen
until they meet and fuse. These plates may be known as the
ventral plates and the pilocerores that excrete them as the ventral
pilocerores. The cephalic end of the ovisac is formed by a trans-
verse band of pilacerores located on the first abdominal sternum.
Its cephalic margin is usually straight. This is why the ventral
surface of the ovisac is generally smooth and not fluted. The
transverse cephalic portion of the ovisac may be known as the
transverse plate and the band of pilocerores excreting it as the
transverse pilacerores. The two rows of plates located on the
dorsal aspect adjacent to each other and the meson are the dorsal
plates and the pilacerores that excrete them as the dorsal
pilacerores. They are apparently the subdorsal keels of Tinsley,
the subdorsal plates of Cockerell, and the dorsal or median lamellae
of other writers. There are typically eleven pairs of dorsal plates,
a row of eleven dorsal plates on each side of the meson. The
cephalic or first pair of dorsal plates is formed by groups of
pilacerores located upon the head and these may be known as the
head plates or the head pilacerores. They vary greatly in size and
shape and, when the two groups of pilacerores are continuous on the
head, are sometimes represented by a single plate. The tenth and
eleventh dorsal plates are usually very different in shape from those
of the other segments. They frequently form semicircular plates
which are continuous on the meson. The fusion of adjacent plates
can be indicated by the use of the word plate instead of plates as
when they are separate. The eleventh pair of dorsal plates is
located on the seventh abdominal segment cephalad of the anal
ring and they may be known as the preanal plates. The tenth
pair of dorsal plates are located on the sixth abdominal segment
and may be known as the paranal plates. The tube of wax sur-
rounding the anus is not excreted by pilacerores but by the cerores
of the anal ring. An examination of this wax will show that it is
different in structure and appearance from that execreted from the
pilacerores, as the wax of the dorsal plates. This tube of wax may
be known as the anal plate.
A few species have plates of wax located on the meson between
the dorsal plates. They are generally designated as the wedge-
110 THE COCCIDAE
shaped plates. These plates are apparently limited to the
mesothorax, metathorax, and the first abdominal segment and may
be known as the mesal plates.
There is a second series of plates which extend around each
margin of the body and form the lateral plates of the test. These
are the lateral plates or lateral pilacerores and are sometimes
designated as the marginal plates, the circumferential lamellae,
the lateral plates, or the lateral keels. There are ten lateral
plates located on each side of the test. The first lateral plate of
each side is formed by a group of pilacerores located on the pro-
thorax. The other groups are located on the other thoracic
segments and abdominal segments one to eight. The pilacerores
forming the lateral plates are located on each lateral portion of
their respective segment adjacent to the band forming the ovisac
or the ventral plate. The abdominal spiracles of segments one to
seven are located among the lateral pilacerores. The lateral
pilacerores of the eighth segment are frequently continuous on the
meson so that the pair of plates of this segment is usually repre-
sented by a single plate and in such cases it is frequently designated
as the telson. It is located caudad of the anal plate and for this
reason has been named the postanal plate. The plates formed by
the groups of lateral pilacerores on segments five, six, and seven
are frequently much larger and longer than any of the others and
have been designated the caudal plates. They are also apparently
the posterior lateral plates of some writers.
In a few species there is a longitudinal row of plates on each
side between the mesal plates and the lateral plates. These have
been named the intercalary plates. Newstead designated them the
supplementary scales. In some descriptions the so called sub-
dorsal keels or plates may be homologous with the intercalary plates.
The subfamily Ortheziinae contains six genera which can be
separated by means of the following table : —
GENERA OF ORTHEZIINAE
a. Antennae of adult female with seven to nine segments and of
nymphal female with six segments; tibio-tarsal sutures of each leg
distinct,
b. Antennae of adult female with nine segments; head of adult
male with three ocelli, Douglasia MacG.
bb. Antennae of adult female with seven or eight segments and of
nymphal female with six segments; head of adult female with
two ocelli or ocelli wanting,
c. Adult female without mesal plates. Orthezia Bosc,
SUBFAMILY ORTHEZHNAB 111
cc. Adult female with three or more mesal plates. Arctorthezia Ckll.
aa. Antennae of the adult female with less than nine segments and of
nymphal female of less than six segments.
I). Antennae of adult female with four or more segments ^nd^ of
nymphal female with four segments; tibio-tarsal suture of each
leg wanting, so that claw appears to be continuous with tibia.
c. Antennae of adult female with six or seven segments and of
nymphal females with four segments. Newsteadia Green.
cc. Antennae of adult female and of nymphal female with four
segments. Otheziola Sulc.
bb. Antennae of adult female and of nymphal female with three seg-
ments; tibio-tarsal suture of each leg wanting, but claws distinct-
ly separated from tarsus. Nipponorthezia Kuw.
Douglasia MacG. — This genus includes a single species, maen-
ariensis Douglas, from the Island of Montecristo, Italy on Erica. The
head plates are small and not strongly projecting or divergent. The
lateral plates are less than the typical number. Douglas figures the
ovisac with ten longitudinal ridges.
Orthezia Bosc. — The great majority of the described species of
this subfamily belong to the typical genus Orthezia. The species are
difficult to recognize because of the different nomenclature used and the
different value assigned the same characters by different authors. It is
not unlikely that some of the species included in the following table
have been misplaced for this reason. The table includes most of the
American species. It is unsatisfactory, but may prove of value in the
identification of specimens. In every case the specimens should be com-
pared with the original description.
SPECIES OF ORTHEZIA
a. Dorsal surface of body of the adult female exposed between dorsal
and lateral plates or caudad of dorsal plates or mesal and dorsal
plates wanting; uncovered portion sometimes covered with mealy
wax.
b. Portion of dorsal surface of adult female not covered by plates of
wax much greater in extent than covered area,
c. Dorsal surface of body of adult female completely exposed be-
tween lateral plates, dorsal plates entirely wanting; ovisac
slightly longer than body; antennae consisting of eight seg-
ments.— Arizona on Quercus. nuda Ferris.
cc. Dorsal surface of body of adult female always with two rows
of dorsal plates.
d. body of adult female exposed as black mesal band between
two rows of dorsal plates.
e. Dorsal plates much farther apart, more divergent on tho-
rax and cephalic abdominal segments; dorsal plates sub-
equal in length and thickness. — United States, Mexico,
West Indies, British Guiana, Brazil, England, South Af-
rica, Mauritius, Ceylon, China on Coleus, Lantana,
\
^
112 THE COCCIDAE
Ipomoea, Thunbergia, Strobilanthus, Verbena, Achillea,
Salvia, Cuphea, Capsicum, Ageratum, Veronia, Gardenia,
Chrysanthemum, Lonicera, Citrus, tea, strawbdfry, to-
mato, etc. insignis Dougl.
ee. Dorsal plates arranged in practically straight rows, paral-
lel, not divergent or farther apart on thorax, cephalic
dorsal plates thicker and longer man caudal; dorsal sur-
face of body naked except for two median rows of small
dorsal plates and a little mealy powder. — New Mexico
monticola Ckll.
dd. Body not exposed between two rows of dorsal plates; cephal-
ic dorsal plates thicker and longer than caudal. — New Mex-
ico on Gutierrezia. nigrocincta Ckll.
bb. Portion of dorsal surface of adult female not covered by plates of
wax much less in extent than covered area.
c. Portion of dorsal surface of body of adult female exposed
limited to small portion caudad of dorsal plates; four large
caudal plates project caudad over ovisac; postanal plate much
abbreviated. — Mexico on Hymenoclea. sonorensis Ckll.
cc. Portions of dorsal surface of body of adult female exposed
limited to narrow band-lie areas between dorsal and lateral
plates.
d. Dorsal plates forming high, double, mesal crest of long erect
white plates; two caudal plates of each side very long and
curving down over sides of ovisac; body pale pea-green. —
New Mexico on Garrya. garryae Ckll.
dd. Dorsal plates not forming high white mesal crest.
e. Caudal plates of each side two or three times as long as
other lateral plates and strongly divergent; head plates
fused and projecting over head as strongly divergent pro-
jections.— Jamaica, Trinidad, British Guiana, Brazil on
Capsicum, Croton, Citrus, Sanchezia. praelonga Dougl.
ee. Caudal plates of each side not greatly elongated and di-
vergent; head plates not strongly divergent projections,
f. Dorsal surface with deep mesal furrow between two
rows of dorsal plates; head plates prolonged over head,
not divergent or greatly produced; caudal plates sub-
equal in length. — Argentine Republic on Compositae(?).
ultima Ckll.
ff. Dorsal surface not with deep mesal furrow between two
rows of dorsal plates; head plates not prolonged over
head; caudal plates longest; caudal dorsal plates widest;
ovisac very long; antennae with third segment longest
or third and eighth subequal. — New Mexico, California
on grass. graminis Ckll.
aa. Dorsal surface of body not exposed, completely concealed by plates
of wax.
b. Dorsal surface not with row of intercalary plates on each side.
SUBFAMILY ORTHEZIINAE 113
c. Ovisac with eight longitudinal ridges,
d. Antennae of adult female with eight segments.
e. Antennae with first segment subequal to secoHTi tmd-first
and second each shorter than third.
f. Dorsum with furrow; antennae with third segment
slightly longer than second, first and second subequal;
antennae and legs dark brown. — New Mexico, Arizona,
Colorado on Chenopodium, Atriplex, Kochia. annae CkU.
ft. Dorsum not with furrow; dorsal plates two to five di-
rected cephalad, remaining dorsal plates successively-
smaller and directed caudad; antennae with third seg-
ment longest; antennae and legs dark red-brown. — New
Mexico on Artemisia. artemisiae Ckll.
ee. Antennae with first segment distinctly longer than second,
f. Antennae with segments one and three subequal and
second segment about one-half their length, eighth seg-
ment longest; legs and antennae brown; wax compact. —
Organ Mountains, New Mexico on Cheilanthus
cheilanthi Tins,
ff. Antennae with second and third segments subequal and
each shorter than first. — California on Bahia
californica Ehr.
dd. Antennae of adult female with seven segments, living in
nests of ants.
e. Caudal plates very long, about two-thirds length of body,
curving over ovisac but not attached to it; body orange
color; antennae with seven segments. — New Mexico from
the nests of Lasius americanus. lasiorum Ckll.
ee. Caudal plates longer than other lateral plates, not long and
curving as long projections over ovisac. — Colorado from
the nests of Lasius. olivacea Ckll.
cc. Ovisac with ten longitudinal ridges; dorsal plates one to six
successively longer and stouter, seven and eight much shorter.
— Ohio, Illinois on Solidago. solidaginis Sands.
bb. Dorsal surface with row of intercalary plates on each side.
c. Ovisac short, postanal plates not fused and nearly as long as
ovisac; caudal plates shorter than postanal plates. — Canada,
New York, Iowa on Solidago, Arctium. americana Walker.
cc. Ovisac long, five or more times as long as fused postanal
plates; caudal plates nearly twice as long as postanal plates. —
Europe on Matricaria, Caltha, Teucrium, Achillaea, Dodder
urticae Linn.
Arctorthezia CklL — This name was proposed by Cockerell for the
two species of Orthezia provided with mesal plates. An examination of
the mesal groups of pilacerores should offer distinctive characters for
the recognition of the genus. The two species can be separated as
follows :
114 THE COCCIDAE
SPECIES OF ARCTORTHEZIA
a. Mesal plates small and not extending beyond caudal margin of their
segment and not overlapping mesal plates of next segment. — Green-
land, Lapland, Norway, Scotland, England, Ireland from Saxifraga,
grass. cataphracta Shaw.
aa. Mesal plates large and extending beyond caudal margin of their
segment and overlapping mesal plate of next segment. — Colorado, New
Mexico from nests of ants. occidentalis Dougl.
Newsteadia Green. — This genus contains a single species, floccosa
De Geer, found in British Isles, Germany, Bohemia, and Australia on
grasses and sedges. There are seven mesal plates, of which the inter-
mediate are the longest, and four lateral plates. The caudal lateral
plates are fused to the ovisac. The Australian specimens were found
upon some wet timbers in a mine at a depth of three hundred feet and
were identified by Green. This species in common with some others is
often collected among moss, lichens, and dead leaves and it is possible
that they may feed upon the two former. Whether the species was intro-
duced into Australia on the timbers is not known.
Ortheziola Sulc. — This genus contains three species, fodiens
Giard from Guadeloupe on the roots of the coffee tree; signoreti Haller
from France; and vejdovskyi Sulc from Prague from under leaves and
moss. This latter species, the type, has the dorsal aspect completely
covered with plates of wax.
Nipponorthezia Kuwana. — A single species, ardisiae Kuwana,
from Japan on Ardisia is included. The dorsum is almost naked due to
the small size of the dorsal plates. The third antennal segment is as
long as the other two together. Each ocellana is located distant from the
articulation of an antenna which is unusual.
CHAPTER XII
SUBFAMILY PHENACOLEACHIINAE
The Bingeyed Coccids
The body of the adult female is convex on the dorsal aspect and flat
on the ventral. It is elongate elliptical in outline with the cephalic end
bluntly rounded and the caudal end bluntly pointed. The caudal end
of the body is not inflated and the cephalic and lateral margins are not
provided with a keel. The segments are well defined. The antennae
are prominent, consisting of eleven segments, and are articulated to the
ventral aspect of the head near each lateral margin. The eyes are not
described. The mouth-parts, rostrum and rostralis, are always present,
the rostrum consists of three distinct segments. The legs are present.
They are subequal in length and normal in form. The profemora are
never enlarged and the prothoracic legs are never fitted for digging.
The trochanto-femoral and tibio-tarsal sutures of all the legs are proba-
bly distinct. The thoracic and abdominal spiracles are not definitely
described and not figured. The stigmatic clefts, spinae, and canellae are
wanting. The abdomen is about as long as the head and thorax togeth-
er. It is convex on the dorsal aspect, never depressed. The lateral mar-
gins converge slightly caudad and the caudal end is bluntly pointed. All
the segments are fully exposed and similar in appearance with distinct
coriae between them, none are retracted to form a marsupium. The
abdomen is never provided with an anal cleft and opercula or anal lobes
and anal setae. The anal ring is always distinct and provided with six
prominent anal ring setae. The caudal abdominal segment is never
short and narrow, extending beyond the other segments. The body is
provided with cerores, none of which are octacerores or pilacerores and
is never provided with ceratubae. The body is never enclosed in a res-
inous cell with three adjacent openings or covered by a scale with or
without exuviae, but is covered with white mealy wax among which is
mixed some curling cottony threads of wax. The rectum is never pro-
vided with a long glassy tube of wax or with a chitinized rectal tube
bearing one or more rings of anacerores. The body is never naked and
gall-like in form. '
The female nymphs of all stages, so far as known, are provided with
a rostrum and rostralis. The three pairs of legs are present in all
nymphal stages and are similar in form. The profemora are never
greatly enlarged and the prothoracic legs are never fitted for digging.
The body is never provided with an anal cleft and opercula, anal lobes
and anal setae, octacerores, pilacerores, or ceratubae. The anal ring
is distinct and bears six anal ring setae. The caudal end of the rectum
is not chitinized, forming a rectal tube provided with one or more rings
115
116 THE COCCIDAE
of anacerores excreting a long glassy tube. The caudal abdominal seg-
ment is not short, narrow, and projecting.
The male has eight ocellanae on each side of the head, forming an
almost complete circle, and two distinct ocelli. The abdomen is not pro-
vided with long lateral filaments, but the caudal end bears on each side
of the meson a bundle of four to six rather long setae and a very long
thick thread-like filament of wax. The wings are sometimes greatly
reduced. The stylus is inconspicuous.
This subfamily contains a single genus, Phenaeoleachia, based
upon a species described by Maskell from New Zealand. There is
but little known regarding the life-history of the species. Maskell
describes the adult female, the first nymphal stage, and the adult
male. The species resembles closely many species of Pseudococcus
in its general habits and general appearance.
The adult female is eight millimeters or less in length, the
two ends of the body are rounded, and the caudal end lacks all
indication of anal lobes and anal setae. These structures are
characteristic of most species provided with a prominent anal ring
and anal ring setae but are not characteristic features of this one
The antennae are long and consist of eleven segments. The body
is covered with mealy and cottony wax, some of the latter form
long curling coarse yellowish threads. Some of the caudal
abdominal segments bear small porrect pencils of wax which give
the insect the general appearance of a mealy-bug.
The nymphs are similar to the adults but smaller. They are
similarly covered with wax and bear about the same number, five
or six, pencils or lateral projections of wax. The antennae consist
of seven segments. The rostrum is elongate and, as in the adult,
consists of three segments. Those species provided with an anal
ring and lacking anal lobes in the adult, generally have large
prominent anal lobes in the nymphal stages, particularly the first,
but there are no indication of such structures in the nymphs of
this species.
The adult males are about two millimeters long and rather
densely covered with white mealy wax. The eyes are represented
by eight ocellanae on each side and, except for a small space on
the meson of the dorsal and ventral aspects, form almost a complete
ring about the head. There is on each side of the meson of the
dorsal aspect caudad of the ocellanae two other simple eyes which
are considered as ocelli. The antennae consist of ten segments.
The legs are long and slender. The caudal end of the abdomen
bears two long, slender, thread-like filaments of wax. The wings,
while normally large and prominent and held in place by halteres
SUBFAMILY PHENACOLEACHIINAE 117
with two hooks, are sometimes so rudimentary that the insect is
almost apterous.
This insect is of great interest. The eleven segmented
antennae of the adult female and the lack of anal lobes and anal
setae marks it as a generalized species while the well developed
anal ring shows a relationship with the anal ring conservers. The
number of simple eyes in the male, their arrangement, and the
presence of ocelli would suggest that this is a stage in the trans-
formation of compound eyes to groups of simple eyes. The males
of all the preceeding subfamilies but one, the Coccinae, have com-
pound eyes. Most of these also have a well developed ocellus
caudad of each compound eye. It may be assumed that all the
facets of each compound eye of Phenacoleachia have been lost
except those forming a half of the ring of each side. The facets
of the compound eyes where they are present are usually large
and not closely associated. It is easy to understand how the
various types with a varying number of ocellanae of the other sub-
families have been developed from a condition such as is found in
Phenacoleachia. The reduction has proceeded in all females until
there is only a single ocellana present on each side of the head.
The fact that in certain if not all wingless males, there is only a
single ocellana on each side, might lead one to assume that a
larger number was not needed in order to locate the females.
Phenacoleachia Ckll. — There is a single species in this subfamily,
Phenacoleachia zealandica Maskell from New Zealand, where it occurs
upon Podocarpus, Cupressus, and Fagus. This species was referred to
the genus Leachia by Maskell and later transferred to Palaeococcus which
is another name for the same group by Cockerell who erected in 1899
the present generic name, Phenacoleachia.
CHAPTER XIII
SUBFAMILY ERIOCOCCINAE
The Mealy Bugs
The body of the adult female varies from convex to suhrotund, the
sides from convex to subparallel. It is elongate or short oval in outline
with the two ends bluntly pointed or bluntly rounded. The segmentation
is usually well defined. The caudal end of the body is not inflated and
the cephalic and lateral margins are not provided with a keel. The
antennae are prominent, consist of five to nine segments, and are articu-
lated to the ventral aspect of the head near each lateral margin or they
are rudimentary or wanting. The eyes are usually present, when pres-
ent, they are inconspicuous fuscous spots, a single ocellana located on
each lateral margin near the articulation of an antenna. Tha mouth-
parts, rostrum and rostraiis, are always present, the rostrum usually
consists of two distinct segments, rarely only one. The legs are typi-
cally present. They may be rudimentary or wanting, if present, they
are subequal in length and normal in form. The profemora are never
enlarged, if enlarged, all the other femora are enlarged. The protho-
racic legs are never fitted for digging. The trcchanto-femoral and the
tibio-tarsal sutures of all normally formed legs are always distinct. The
spiracles of the mesothorax and metathorax are normal in size and
form and ventral in position. The mesothoracic spiracles are never
located near the anus. The abdominal spiracles are always wanting.
The stigmatic clefts, spinae, and canellae are wanting. The abdomen is
normally about as long as the head and thorax together. It is subconvex
on the dorsal aspect and flat on the ventral aspect, sometimes thin and
disk-like, approaching a depressed condition. The lateral margins are
nearly straight and subparallel or strongly convergent and convex, the
caudal end is bluntly pointed or bluntly rounded or almost truncate. All
©f the segments are fully exposed and similar in appearance v^^ith dis-
tinct coriae between them, none are retracted to form a marsupium.
The abdomen is never provided with an anal cleft and opercula. There
is typically a distinct anal ring provided with four, six, eight, or more
anal ring setae, anal ring and anal setae sometimes wanting. The anal
lobes are usually distinct and bear anal setae. The caudal abdominal
segment is never short and narrow and extending beyond the other seg-
ments, the caudal segments at least are campanulate. The body is pro-
vided with cerores none of which are octacerores or pilacerores and is
never provided with ceratubae. The body is never enclosed in a resinous
cell with three adjacent openings or covered by a scale with or without
exuviae, but is generally completely covered with powdery or mealy
wax, sometimes with masses of cottony or woolly threads of wax, often
found on or within a mass of rather compact cottony threads of wax
lis
SUBFAMILY ERIOCOCCINAE 119
among which the eggs are placed. The rectum is never provided with
a long glassy tube of wax or with a chitinized rectal tube bearing rings
of anacerores. The body is never naked and gall-like in form.
The female nymphs of all stages are provided with a rostrum and
rostralis. The three pairs of legs are present in the first nymphal stage
and generally in the other nymphal stages, sometimes greatly reduced or
wanting, when present, are usually similar in form. The profemora are
never greatly enlarged and the prothoracic legs are never fitted for dig-
ging. The caudal end of the rectum is not chitinized, forming a rectal
tube provided with one or more rings of anacerores excreting a long
glassy tube of wax. The body is never provided with an anal cleft and
opercula or pilacerores and rarely with octacerores or ceratubae. The
anal ring is distinct and provided with four to eight or more anal ring
setae, usually six, and frequently different in number from that of the
adult. The anal lobes are usually distinct and provided with anal setae.
The caudal abdominal segment is not short, narrow, and projecting.
The male has ocellanae. The abdomen is not provided with lateral
filaments or with tufts of long slender glassy threads of wax, but is
usually provided with two or four long slender caudal threads of wax.
Wings are sometimes wanting. The stylus is short and inconspicuous.
This subfamily contains some of the best known species of the
family, the species of Pseudococcus, generally known as Mealy
Bugs. This name comes from the fact that their bodies, dorsal
and ventral aspects, are generally densely covered with a fine
granular wax which gives them the appearance of having been
rolled in coarse flour. Several of the species are of great economic
importance in subtropical and tropical regions where they are
able to pass the winter out of doors. In temperate regions where
the winters are generally rigorous, they are found in the main
only in glass houses where they are often serious pests unless held
in check by frequent fumigations.
The females are active throughout their entire or almost their
entire life and in all nymphal stages, cover their bodies with
mealy wax, so that they appear conspicuous against the green
surface of the plant. They are prone to congregate together, so
that they assemble, usually from two to six, in the angle between
two branches or in the angle between a leaf and a branch. The
adult females form the mass of wax, the so called ovisac, in which
the eggs, 150 to 200 in the case of Pseudococcus citri, are depos-
ited. These are usually placed in the angles between the stems or
a leaf. The eggs hatch in ten to eighteen days, but of the eggs
deposited only a small number produce males. The young nymphs
remain in the ovisac for a short time, but on leaving spread rapidly
over various parts of the plant and settle along the midrib and in
the angles in the stems.
120 THE COCCIDAE
The segmentation of the body is distinct and is usually made
clearer by the covering of mealy wax. The antennae are articulated
to the ventral aspect of the head and the single ocellana on each
side shows prominently on the ventral surface through the presence
of black pigment. The female molts three times and passes through
three nymphal stages. The females are very similar in all the
stages, the only striking difference between them is that of size.
Even this is not constant for there is considerable variation in the
size of different individuals even of the same age or stage. There
is also a variation, as in other coccids, in the number of antennal
segments present in individuals of the various stages.
It is impossible to differentiate the males from the females
during the first nymphal stage, except in mounted specimens that
have almost completed the first nymphal stage. Berlese has
pointed out that in such individuals, the mandibles and maxillae
of the second stage of the female can be identified before those
of the first stage are shed. They can be observed coiled spirally
on each side of the meson beneath the cuticle of the body. Since
the males lose their rostralis at the first molt in most if not all
species of this subfamily, there is no such spirally coiled rostralis
to be observed under the cuticle in this sex.
The nymphs of the male molt four times in Pseudococcus
citri and pass through four nymphal stages. The male nymphs
are sluggish and seek some secluded spot where they spin a cocoon
of felted threads of wax. The second molt occurs soon after the
completion of the cocoon and the cast skins are pushed out of the
end of the cocoon. The eyes of the adult male are simple, six
ocellanae, a pair of comparatively large ones on the dorsal aspect.
a similar pair on the ventral aspect, and one on each lateral aspect
which are about one-half the size of the others. They are some-
times designated as the primitive eyes and are considered as
representing the ocellanae present in the first nymphal stage.
The supposition being that they have been retained through all the
nymphal stages although not identifiable in the latter ones. They
are also considered as ocelli and the large facets in such cases are
designated as eyes. The number of ocellanae varies in the males
of other genera than Pseudococcus, that is, there is more than six.
Writers are careless about stating the number in most cases. The
winsrless males of Fonscolombia have retained only the lateral
ocellanae. Whether the wingless males of other groups have only
the lateral ocellanae is not known. The wing-pads and developing
SUBFAMILY ERIOCOCCINAE 121
legs and antennae of the male appear after the second molt. The
antennae and legs of the nymph are lost at the first molt. The
adult males are minute fragile creatures with delicate wings.
The halteres are small and provided with hooks. The males of
Fonscolombia are always wingless and the males of a few other
genera are sometimes wingless.
The wax excreted by the female on different parts of the body
appears from a cursory examination to be identical. A careful
examination will show, however, that there are several different
types. An unpublished study of the cerores of Pseudococcus citri
and the wax they excrete was made in 1907 under my directions
by Robert Matheson, he found that there were several distinct
types, four or more, and that the wax excreted from each of these
types was of a different kind, but the difference could be determined
only in most cases after a careful microscopical examination of the
wax. The predominant type of ceroris in this species is the
diamond-shaped cerores with numerous openings. These have been
named the tricerores. While they are of general occurrence on all
parts of the dorsal and lateral aspects, they are usually more
numerous upon the anal lobes and form the pencils or projections
of wax attached to the anal lobes. These pencils are not confined
to the anal lobes but in many species there is a row of them along
each lateral margin of the body. The lateral pencils are excreted
by groups of tricerores like those of the anal lobes. The porrect
form of the pencils is due to the presence of one or more
short conical setae. The number varies not only with the segment
but with the species. The setae are typical in form and frequently
are as broad as long. They are usually associated with the tri-
cerores and support the pencils of wax. Berlese named these
groups of tricerores and conical setae filiere. They are also known
as cerari, cerarii, and filuri. There are eighteen pairs of cerari on
each side of the body in the common greenhouse Pseudococcus,
which must take the name of destructor Comstock, while there are
only seventeen in citri according to Berlese, Marchal, and Ferris.
The cerari of destructor are arranged, four on the head, only three
in citri, two on each thoracic segment and one on each of the eight
abdominal segments. The cerari of the anal lobes, the eighteenth
pair, as they are generally counted, are larger than those on the
other segments of the body. These cerari are known as the anal
cerari and all the others as the lateral cerari. If necessary those
of the head, thorax, and abdomen can be distinguished as the head
122 THE COCCIDAE
cerari, thoracic cerari, and abdominal cerari. There may be twc
rows of cerari on the dorsal aspect between the lateral cerari.
the dorsal cerari, and if a single dorsal row should be present, they
could be known as the mesal cerari. The setae of the cerari are
not always short and conical, but may be long and slender like
the other setae in certain species. There are often ordinary setae
associated with the cerari, named the auxiliary setae by Ferris to
distinguish them from the conical setae of the cerari, the cerarari
setae. The ordinary setae of the dorsal aspect are known as the
dorsal setae. The number of ceraran setae varies from two to
twenty, the largest number so far observed. If there is a vari-
ation in number between the various cerari, there is likely to be a
larger number in the first six cerari or in the anal cerari. The
cerari are sometimes obscure in the adult female, probably due to
the swelling of the body by the eggs, but are generally very dis-
tinct in the half grown nymphs. In certain genera, as Antonia,
the cerari are wanting ; are represented by only a few in others, as
Trionymus, where only the anal cerari are present in some species,
while others may have four pairs ; or there may be a considerable
number of pairs as in many species of Pseudococcus. The number
varies from a single pair to as many as twenty-four pairs in
Macrocerococcus as figured by Leonardi.
Marchal and Ferris have described some minute cylinders
which are connected with the external cuticle of the body-wall.
Each is represented as having one side of the distal end continued
as a much smaller cylinder. These cuticular cylinders are con-
sidered as the outlets of wax cells, which are circular in outline.
The number in destructor seems to be variable. I have found
them more often along the margin of the body near the anal lobes.
Some species seem to show them in considerable number. They
are regarded as ceratubae.
The ventral aspect of the abdomen of the female bears the
numerous cerores from which the cottony threads of wax in which
the female conceals herself and her eggs. These cerores are
peculiar to the adult female, are typical in form, and like the
genacerores do not appear until after the last molt.
Berlese has called attention to the presence of four peculiar
structures located on the dorsal aspect which he designates as the
foveola labiate. There is two of them on the head and two in the
coria between the sixth and seventh abdominal segments. These
SUBFAMILY ERIOCOCCINAE 123
structures may be known as the labiae, those on the head as the
cephalabiae and those on the abdomen as the caudalabiae. They
are also known as the eye-like glands, cicatrices, f ossette ostioloform
labiate, and dorsal ostioles. The labiae were considered by Ber-
lese as places where the cuticle had been invaginated for the
attachment of muscles. They are lip-like structures. The sides
or infolded surfaces of the lips or labiae bear setae and cerores.
The caudalabiae are generally longer than the cephalabiae, the
latter are sometimes greatly reduced and wanting. Between the lips
of the labiae there appears to be the mouth of invaginated pockets.
There can frequently be observed on living specimens a small
globule of a clear fluid over the mouth of each labia, more
frequently the caudalabiae than the cephalabiae, so that they are
probably also glandular in structure as suggested by Comstock.
For, as he suggested, when the specimens are stroked with a pencil
or dissecting needle, the insect will hump up its back and extrude
a globule of liquid. The insect is unable to repeat this operation
until the pocket is again filled with the clear fluid. Specimens
have been observed to extrude globules from all four labiae at the
same time. The labiae undoubtedly have a glandular function
which is probably of later origin than their earlier function, a
parademe for the attachment of muscles. Sulc, who has made a
histological study of these structures, believes that the globules
of sticky matter rolled in wax are for defensive purposes. "When
they come into contact with antennae or palpi of the enemy, as he
states, these appendages are all stuck up and unserviceable until
they have been cleaned. This fact together with the belief that
the excretion is derived from the fat tissue, has led Sulc to name
these structures the ' ' adiopugnatorische organ."
There is another labia on the ventral aspect of the abdomen
between the second and third segments, the ventralabia. This has
not been observed to extrude any globules. Sulc has also made a
careful study of this structure. He considers this labia as entirely
different in origin and structure from the cephalabiae and
caudalabiae. His published figures support his contention as to
their structure. He considers, I believe erroneously, this structure
as homologous with the stink glands of the adult heteropterous
Hemiptera. He also calls attention to a rudimentary structure in
the first abdominal segment which does not show in specimens
124 THE COCCIDAE
cleared in caustic potash. The ventralabia is not an invaginated
pocket like the other labiae.
The ventral aspect of the head frequently bears about eight
setae located between and caudad of the articulation of the
antennae. These setae are known as the interantennal setae.
They can usually be identified from the other setae of this region
by their greater size and the regularity of their arrangement.
The mesothoracic and metathoracic spiracles are located in
the transverse furrows marking the position of the mesocoria and
metacoria. They appear in mounted specimens as hour-glass-
shaped structures. One end of each spiracle is smaller than the
other, is located in the cuticle of the body-wall, and contains a
small opening. This opening is the spiracle. The round plate of
cuticle which surrounds the spiracle is the peritreme. The re-
mainder of the hour-glass-shaped or funnel-shaped structure is a
modification of the spiracular trachea, although the entire structure
is generally figured and described as a spiracle. The flaring
structure or modification of the spiracular trachea is a cavera.
The abdominal spiracles are always wanting.
The claws, while generally simple, in a few genera bear a
single tooth near the middle of their ventral aspect. This is the
"denticle of the face of the claw" of Ferris. There is also some
variation in the number and form of the digitules borne by the
claws.
The anal ring and the anal ring setae are typically present
in the adult female. When absent, they are present in the nymphal
stages or at least in the first nymphal stage. The normal number
of anal ring setae is six judging from the large number of genera
that have this number and also from the fact that those genera
\yithout, with less than, or with more than six in the adult female
often have six in the first nymphal stage. The number varies
from two to fifty or more, but is never apparently an odd number.
The anal ring setae can be differentiated as the cephalic, median,
and caudal when there are only six present. The cephalic anal
Ting setae are those on the cephalic or dorsal portion of the anal
ring, depending upon the position of the ring. The anal ring is
usually more or less vertical in position, but may be dorsal, as
is frequently the case with the anus in the coccids. The closed
vend of the anal ring in Pseudococcus which is the dorsal side, if
SUBFAMILY ERIOCOCCINAE 125
the ring is vertical, is considered as the cephalic end and the
opposite end as the caudal. The anal ring bears cerores, which
are arranged in two rows in Pseudococcus, the orbacerores, and
judging from their staining reaction are different in structure.
The outer and shorter row is the lateral orbacerores, the inner and
mesal row is the mesal orbacerores. This latter is frequently
continuous between the cephalic anal ring setae. Some of the
cerores of the mesal row in this genus are elongated and produce
extensions of or irregularities in the membrane surrounding the
anus. These cerores may be known as the dentacerores. They
were called denticulate pores by Smith, who was the first to call
attention to them. The number of dentacerores on each side of
the anal ring is not large, so far as observed two to four. It is
difficult to make good flat preparations showing the characteristic
number and arrangement of the orbacerores, dentacerores, and
anal ring setae, but my observations lead me to believe that some
of the best characters for the differentiation of species are to be
found in these structures. The importance of these characters
was first emphasized by P. E. Smith. Flat preparations for the
study of these structures can be secured only by dissecting out
the anal ring and mounting it separately. The anal lobes should
be removed and the dissecting should be done on a slide after the
specimens have been stained and cleared.
There are in certain genera four prominent setae located
caudad or ventrad or cephalad of the caudal end of the anal ring,
depending upon whether the anal ring is dorsal, caudal or vertical,
or partially ventral. These setae are known as the cisanal and
obanal setae. They mark the four corners of a quadrangular area.
The two setae nearest to the anal ring are the longest and are
known as the cisanal setae. These are the cephalic postanal setae
of Smith. The two setae farthest from the anal ring are generally
smaller than the cisanal setae and are frequently located farther
apart. They are obanal setae, the caudal postanal setae of Smith.
The species of this subfamily are frequently based upon the
comparative length of the antennal segments. Kellogg and Bell
and later P. E. Smith made careful studies and measurements of
antennae of specimens of unquestioned identity and showed that
characters of this sort were worthless for the separation of species.
Their studies went to show that there was enough variation in the
length of the segments of the two antennae of the same individual
126 THE COCCIDAE
to make different species of them. While Smith has shown of how
little value most of the comparative characters used in separating
species are, he has called attention to the use of several new struc-
tural characters, as the cephalabiae and caudalabiae which were
first figured by Berlese, the cerari and the number of conical setae
contained in each which had been previously used by Marchal
unbeknown to Smith, the cisanal and obanal setae, the cerores of
the anal ring and their arrangement in rows, the difference in the
form and extent of the two rows, and the dentacerores of the mesal
row. It has also been noted by Matheson that in the case of
Pseudococcus citri that there is a great variation in the size of
the adult females when they commence laying their eggs, as he
states, "often what I, judging from the size only, considered were
nymphs in the third stage would prove to be mature females and
commence egg laying." This is undoubtedly true of many if not
most other species of coccids.
The subfamily Eriococcinae includes two groups of genera
which are easily distinguished in the first nymphal stage. In one
group the dorsum bears rows of blunt conical setae and in the
other the dorsum bears fine pointed setae. This latter group
clearly includes the generalized genera and has been designated
as a tribe by Cockerell under the name Dactylopiini. The group
with the rows of blunt dorsal setae is clearly the specialized end
of the series, genera in which the ordinary pointed setae have
become changed in form and acquired a symmetrical arrangement
and have been designated as a tribe by the same author under the
name of Ericoccini. "Whether these two groups of genera repre-
sent distinct subfamilies as the form and arrangement of the setae
of the body might suggest, is not clear. Our knowledge of the
anatomy of the adult female is very imperfect. What is needed
in this subfamily is not the description of more new species and
genera, but a careful study of the anatomy of the described forms
and an extended search for additional characters upon which their
classification and their phylogeny can be based.
The following table to the genera is based in great part upon
descriptions and is faulty not only in construction but undoubtedly
in the characters used. Where a difference in number of antennal
segments is shown, specimens should be traced through both sides
of the table. I have egg laying females of what I believe are
undoubtedly species of Phenacoccus with less than nine segments
SUBFAMILY ERIOCOCCINAE 127
in the antennae. It is hoped that it will save at least the repeated
reading of a large number of descriptions. The arrangement of
the genera placed after the table is that of the table. It^ is not
intended to show anything as to the relationship of the genera.
GENERA OF ERIOCOCCINAE
a. Thorax of adult female with three pairs of normal legs.
b. Insects not producing or living in galls upon plants.
c. Anal ring bearing six anal ring setae in at least some stage of
development of insect.
d. Anal ring bearing six anal ring seitae in all stages, nymphs
and adult.
e. Antennae of adult female with less than nine segments.
f. Antennae consisting of eight segments.
g. Body of insect oval in outline, subdepressed, en-
closed in cottony wax but not in cottony sac of wax;
body not bearing numerous short conical setae.
h. Body with lateral margins not produced into
series of blunt tubercles, each bearing cerarus.
1. Adult female with lateral margins of body con-
vex, body about twice as long as wide; nymphs
of male with antennae consisting of six seg-
ments; abdomen with labiae and usually with
more than four pairs of cerari.
Pseudococcus Westw.
ii. Adult female with lateral margins of body
straight and subparallel, about three times as
long as wide; nymphs of male with antennae
consisting of eight segments; abdomen with
labiae and never with more than four pairs of
cerari; antennae of adult female sometimes
consisting of seven segments. Trionymus Berg.
hh. Body with lateral margins produced into series
of prominent blunt tubercles, each bearing
cerarus; cephalabiae and caudalabiae present.
Tylococcus Newst.
gg. Body of insect circular in outline, globular or sub-
globular, without lateral cottony tufts and enclosed
wholly or in part in cottony sac of wax; body bear-
ing short conical setae and with only anal cerari
present. Erium Maskell.
ff. Antennae consisting of less than eight segments.
g. Adult female not enclosed in shining black scale
with caudal orifice.
h. Anal lobes not strongly chitinized with blunt
spine-like projections at distal end.
128 THE COCCIDAE
i. Cuticle of adult female not with numerous con-
ical setae that are as broad as long,
j. Body not circular in outline; legs not great-
ly swollen and enlarged,
k. Anal ring retracted and surrounded by ir-
regular outer chitinous ring or plate which
supports on each side prominent tubercle
bearing several stout setae; antenna of
adult female consisting of seven segments.
Lefroyia Green,
kk. Anal ring not retracted and surrounded by
outer chitinous ring supporting tubercles
with prominent setae; antennae of adult fe-
male usually with less than seven seg-
ments.
1. Antennae of adult female always with
more than three segments.
m. Distance on the ventral aspect of head
between antacoriae distinctly less than
length of two proximal segments of
antennae.
n. Antennae of the adult female with
five or six segments, not genicul-
ate; distal segment nearly as long
as other segments together
Ripersiella Ckll.
nn. Antennae of adult female with five
segments; antennae geniculate
Rhizoecus Kunck.
mm. Distance on ventral aspect of head
between antacoriae equal to or great-
er than length of two proximal seg-
ments of antennae.
n. Cephalic pair of anal ring setae of
adult female nearer together or at
most only slightly farther apart
than caudal pair; cuticle of female
nymphs not provided with rows of
blunt setae.
o. Antennae of adult female with
six segments, rarely five or seven;
cerari wanting or only anal
cerari present; ceraran setae seti-
form, not conical.
Ripersia Sign.
00. Antennae of adult female with
six segments; anal cerari only
present; ceraran setae setiform.
Gryptoripersia Ckll.
SUBFAMILY ERIOCOCCINAE 129
nn. Cephalic pair of anal ring setae of
adult female twice as far apart as
caudal pair; cuticle of female
nymphs provided with rows of
blunt setae; antennae of adult fe-
male consisting of six segments;
body without labiae
FonscolomMa Fonsc.
11. Antennae of adult female consisting of
three segments; femora with distal por-
tion dilated; tarsal claws large, entire,
scarcely arcuate, armed; presence of six
anal ring setae not certain
Termitococcus Silv.
jj. Body circular in outline; legs short and stout,
femora greatly swollen, radiating from peri-
phery of the body; abdomen shorter than dis-
tance between two caudal pairs of legs; an-
tennae of adult female consisting of six seg-
ments. Pseudoripersia Ckll.
ii. Cuticle of adult female with numerous conical
setae that are about as broad as long; anal
ring crescentic in outline with anal ring setae
located on convex side; antennae of adult fe-
male with seven segments. Gymnococcus Dougl.
hh. Anal lobes strongly chitinized with blunt spine-
like projection at distal end, making them appear
segmented; anal ring circular; antennae of adult
female with six segments; cuticle with conspicu-
ous irregularly arranged cefores in groups of
three. Geococcus Green.
gg. Adult female enclosed in shining black scale with
caudal orifice; antennae of adult female consisting of
six segments; anal lobes not much produced
Porococcus Ckll.
ee. Antennae of adult female consisting of nine segments.
f. Body of adult female covered with mealy or granular
wax, wax frequently forming pencils but never fused to
form plates or lamellae.
g. Abdomen with glistening discs on ventral aspect of
first three abdominal segments, probably marking
position of labiae. Coccura Sulc.
gg. Abdomen not with glistening discs on ventral aspect
of first three abdominal segments, even if with ven-
tralabiae, not with them on first three abdominal seg-
ments.
h. Adult female with simple claws, not toothed.
i. Body with twenty-four pairs of cerari and six
130 THE COCCIDAE
dorsal longitudinal rows of cerari-like struc-
tures, Macrocerococcus Leon.
ii. Body with fourteen pairs of cerari and with-
out dorsal longitudinal rows of cerari-like struc-
tures. Macrocepicoccus Morr.
hh. Adult female with claws distinctly toothed; body
at most only rarely with more than eighteen pairs
of cerari.
i. Adult female provided with large cerores sur-
rounded by three short spine-like setae, cerores
opening in short processes which are arranged
in longitudinal rows; each ceroris excreting a
long glassy thread of wax Heliococcus Sulc,
ii. Adult female not provided with large cerores
surrounded by three spine-like setae opening in
projections arranged in longitudinal rows.
j. Adult female with tricerores and round ceror-
es; antennae of adult female consisting of
eight or nine segments, usually nine; body
usually with eighteen pairs of cerores
Phenacoccus Ckll.
jj. Adult female with round cerores only; an-
tennae of adult female consisting of nine seg-
ments; body with only two pairs of cerores,
these located at the caudal end of abdomen.
Heterococcus Ferris,
ff. Body of adult female not covered with mealy or granu-
lar wax, but with wax fused to form plates or lamellae;
antennae of adult female consisting of nine segments;
claws of adult female with tooth; body usually with
eighteen pairs of cerari. Geroputo Sulc.
dd. Anal ring of first stage nymphs with six anal ring setae and
of adult female with more or less than six anal ring setae.
e. Anal ring of adult female with two minute anal ring setae.
Ehrhornia Ferris,
ee. Anal ring of adult female with twenty-four to thirty anal
ring setae. Tra'butina Marchal.
cc. Anal ring bearing eight or more anal ring setae,
d. Anal ring bearing eight anal ring setae.
e. Antennae of adult female with nine segments; tarsal
claws with tooth at middle; body of first stage nymphs not
with row of blunt dorsal setae. Puto Sign.
ee. Antennae of adult female with less than nine segments;
tarsal claws simple, not with tooth; body of first stage
nymphs with rows of blunt dorsal tubercles.
f. Body of adult female naked; antennae of adult female
consisting of seven segments. Rhizococcus Sign.
SUBFAMILY ERIOCOCCINAE 131
ff. Body of adult female not naked, covered at least in part
by wax.
g. Body of adult female uniformly covered witlt m«aly
or granular wax, never with lateral tufts of cottony
lamellae-like wax; antennae of adult female with six
or seven segments. Ericoccus Targ.
gg. Body of adult female with the dorsal aspect naked,
hut lateral aspects with prominent erect tufts of cot-
tony wax resembling lamellae or plates; antennae of
adult female with six segments. Gossyparia Sign.
dd. Anal ring always bearing more than eight anal ring setae,
e. Antennae of adult female consisting of seven segments;
anal ring of adult female with twenty to twenty-four anal
ring setae. Lachnodius Mask.
ee. Antennae of adult female always with less than seven seg-
ments.
f. Antennae of adult female consisting of six segments;
anal ring of adult female with fifty or more anal ring
setae. Lachnodiella Hemp.
ff. Antennae of adult female consisting of three segments;
anal ring of adult female with twelve anal ring setae.
Micrococcus Leon,
bb. Insects producing or living in galls upon plants.
c. Antennae and legs normal in size and form, not rudimentary.
d. Antennae of adult female consisting of seven segments; anal
ring of adult female with anal ring setae; galls subglobular
swellings at the base of leaf Atriplicia Ckll. & Rohw.
dd. Antennae of adult female consisting of six segments; anal
ring of adult female without anal ring setae; galls formed on
both sides of leaf; anus guarded by four sharp setae
Tectococcus Hempel.
cc. Antennae and legs rudimentary; antennae of nymphs of first
stage consisting of six segments; anal ring with long setae;
galls cup-shaped and growing upon branches
Cissococcus Ckll.
aa. Thorax of adult female not with three pairs of normal legs, some or
all of them greatly reduced or wanting.
b. Anal ring of adult female always with anal ring setae.
c. Anal ring of adult female and of first stage nymph always
with six or less anal ring setae.
d. Anal ring of adult female and first stage nymph with six
anal ring setae.
e. Abdomen never with ventral cribiform plate caudad of
each metaspiracle.
f. Nymphs of first stage not with rows of dorsal setae;
adult female with stub-like vestigial antennae, usually
132 THE COCCIDAE
consisting of three segments and without legs; rostrum
not located on prominent protuberance,
g. Abdomen of adult female sometimes with the caudal
end more or less invaginated, forming anal tube with
anal ring with six large anal ring setae; first stage
nymph with antennae consisting of six segments and
anal ring with six anal ring setae.
h. Adult female and first nymphal stage with cau-
dalabiae; adult female enclosed in tough cottony
ovisac from which projects small tube of wax
Antonia Sign,
hh. Adult female without caudalabiae and first nym-
phal stage with cephalabiae and caudalabiae; last
nymphal stage of female with legs; adult female
enclosed between small scales, clothing base of
long pointed gall Sphaer-ococcus Mask.
gg. Abdomen of adult female with caudal end not invagi-
nated to form anal tube; anal ring with six small
anal ring setae; first stage nymph with antennae
consisting of six segments and with anal ring with
six slender anal ring setae; body apparently without
cephalabiae and caudalabiae; adult female enclosed
under sheathing base of host plant
Paludicoccus Ferris,
ff. Nymphs of first stage with rows of dorsal setae; adult
female with antennae rudimentary, consisting of three
segments, and without legs; rostrum borne by promi-
nent cephalic prolongation; antennae of first stage
nymph consisting of six segments Nidularia Targ.
ee. Abdomen with circular cribiform plate on ventral aspect
caudad of each metaspiracle; antennae of adult female
consisting of one segment and of first stage nymph consist-
ing of three segments; adult female without legs
Kuwanina Ckll.
dd. Anal ring of adult female and of first stage nymph with four
short spine-like setae; antennae of adult female vestigial,
consisting of about four segments; metathoracic legs vesti-
gial, others wanting; antennae of first stage nymph consist-
ing of five segments; body without labiae.
Cryptococcus Dougl.
cc. Anal ring of adult female and of first stage nymph with seven-
teen anal ring setae; abdomen with prominent tubercle caudad
of each metaspiracle; antennae and legs of adult female want-
ing; nymphs of first stage with antennae consisting of seven
segments. Kermicus Newst.
lib. Anal ring of adult female described as without anal ring setae;
antennae of adult female vestigial, segmentation difficult to iden-
tify, about three segments, legs always wanting; antennae of first
nymphal stage consisting of six segments, anal tubercles small
SUBFAMILY ERIOCOCCINAE 133
and inconspicuous; anal ring of female nymphs with six anal
ring setae. Chaetococcus Mask.
Psendococcus Westw. — This is the largest genus of the subfamily.
Over one-half of the descrihed species, one hundred or more, have been
reported from America. The species are as a rule very poorly differenti-
ated and difficult to identify. The most of them have been based upon
the comparative length of the antennal segments, characters which have
been shown several times to be absolutely worthless. The only published
study of a considerable number of species of any value is that of Ferris.
He has made use of the cerari, characters first noted by Marchal and
Smith independently. It is unfortunate that Ferris did not make use
of the cerores of the anal ring and the setae associated with them.
Credit is due Smith for the accuracy of his observations on the anal
ring as all his work was based upon unstained specimens. The follow-
ing table, although differently arranged, is based upon the work of Ferris,
supplemented where possible by that of Smith. There are the following
American species not included in the table: — aphylonis Ckll. from Dis-
trict of Columbia on Aphyllon; azaleae Tins, from California on Azalea;
brevipes Ckll. from Jamaica on pineapple; bromeliae Bouche from India,
South Africa, Zanzibar, South America, Massachusetts on Mulberry,
Canna, Hibiscum, pineapple; claviger King & Tins, from Massachusetts
from the nests of Lasius; cockerelli King & Tins, from Massachusetts
from nests of ants; cualatensis Ckll. from Mexico; dasylirii Ckll. from
New Mexico on Dasylirion; filamentosus Ckll. from South Caicos Island,
Jamaica, Mauritius, Hawaiian Islands, Japan; formicarii Ehrh. from
Arizona on roots of Artemesia; grandis Hemp, from Brazil on Myrtaceae;
hymenocleae Ckll. from Arizona on Hymenoclea; iceryoides Mask, from
New Zealand, Australia, California on Fagus; ledi Ckll. from New York
on Ledum; liliacearum Bouche from South America on Liliaceae; mag-
nolicida King from Brazil; mamillariae Bouche from France, Central
America on Mamillaria, Cactus; mendozinus Leon, from Argentina on
Hyalis; missionum Ckll. from Argentina; neomexicanus Tins, from New
Mexico on Gutierrezia; olivaceus Ckll. from Mexico on Yucca; pseudoni-
piae Ckll. from Massachusetts, Michigan, California, Mexico on Cocoa-
nut-palm; percerorsus Leon, from Argentina on Gourliea; phoradendri
Ckll. from Arizona in hollow stems of Phoradendron attended by
Cremastogaster ; quaintancii Tins, from Florida on Rhus; roseotinctus
T. & W. Ckll. from New Mexico on roots of grass; sacchari Ckll. from
Trinidad, Barbadoes, Porto Rico, Mauritius, Mexico on Sugar-cane;
secretus Hemp, from Brazil on Solenaceae; segregatus Ckll. from Jamaica
on grass; setosus Hemp, from Brazil on Ficus; simplex Ckll. from Ja-
maica on Pancratum, "Liliaceous plants;" solani Ckll. from New Mex-
ico on potato; subterraneus Hemp, from Argentina on roots of grape;
texensis Tins, from Texas on Acacia; townsendi Ckll. from New Mexico;
tuliparum Bouche from South America; virgatus Ckll. from Jamaica,
Mexico, Texas, Mauritius, Hawaiian Islands on Acalypha, violets, cotton,
cocoanut-palm, cactus; and wheeleri King from Texas from nests of
Campanotus.
134 THE coccroAE
SPECIES OF PSEUDOCOCCUS
a. Anal cerari with ten or more ceraran setae.
b. Body with seventeen pairs of cerari, anal cerari with fifteen to
twenty ceraran setae, other cerari with three to six ceraran setae;
anal ring with single dentaceroris on each side; cisanal setae
adjacent to anal ring, distance between them subequal to distance
between obanal setae. — California on Ramona, Artemisia, Quercus,
Pasania, Adenostoma. crawii Coq.
bb. Body with not more than eleven pairs of cerari, anal cerari with
fifteen to twenty ceraran setae, first pair of cerari cephalad of anal
cerari with six to ten ceraran setae, other cerari with two to four
ceraran setae. — California on Ephedra. ephedrae Coq.
aa. Anal cerari never with more than two ceraran setae.
b. Cerari other than anal cerari never with more than two ceraran
setae.
c. Body always with eighteen pairs of cerari; ceraran setae us-
ually two in number, rarely with one or two of cephalic cerari
with three or four ceraran setae. — Eastern United States on
Coleus, ferns, various greenhouse plants. destructor Comst.
cc. Body always with less than eighteen cerari.
d. Body always with at least seventeen pairs of cerari.
e. Anal cerari never placed on well defined chitinized area,
f. Ceraran setae of usual conical form, never with distal
portion of shaft or with distal portion of shaft of dorsal
setae flagelliform; cerari without auxiliary setae,
ceraran setae large; dorsal setae small and scattered;
anal ring setae shorter than anal setae; anal ring with
two dentacerores on each side; the cisanal setae dis-
tinctly closer together than obanal setae. — Europe,
United States, Jamaica, Brazil, Hawaiian Islands, Mauri-
tius on orange, lemon, coffee, tobacco, ivy, peony, coleus,
and many other plants. citri Risso.
ff. Ceraran seate with distal portion of shaft flagelliform;
dorsal setae numerous, elongate, with dorsal portion
more or less flagelliform; cerari without auxiliary
setae; anal setae about one and one-half times as long
as anal ring setae. — Japan, California, New Jersey on
Krauhnia. krauhnicte Kwn.
ee. Anal cerari always placed on well chitinized area; ceraran
setae not with distal portion flagelliform; dorsal setae
numerous, long; anal cerari and adjacent one or two pairs
of lateral cerari with auxiliary setae, other cerari without;
anal setae and anal ring setae subequal. — California on
Distichlis. timberlakei Ckll.
dd. Body always with less than seventeen pairs of cerari.
e. Body with twelve to fifteen pairs of cerari; ceraran setae
short and stout, those of lateral cerari distant, more dis-
tant than those of anal cerari; auxiliary setae wanting,
SUBFAMILY ERIOCOCCINAE 135
also groups of tricerores; anal setae slightly shorter than
anal ring setae. This statement of characters is based
upon the work of Ferris. One of my students who has
examined specimens of nipae and, what we believe to be
pseudonipae has found characters showing these two spe-
cies to be not only well but easily differentiated species. — '
British Guiana, Mexico, Massachusetts, Indiana, Michigan,
California on Nipa, palms nipa Mask.
ee. Body with five to six pairs of cerari present on five or six
caudal segments; anal cerari on indefinite chitinized area
bearing two or three slender auxiliary setae and few tri-
cerores; dorsal setae small and few in number; anal setae
subequal to anal ring setae. — Australia, New Zealand,
Natal, California on Araucaria, Dammara
aurilanatus Mask,
bb. Cerari other than the anal cerari, at least cephalic pair, always
with more than two ceraran setae.
c. Body always with more than twelve pairs of cerari.
4. Body with cephalic six or seven pairs of cerari with each
cerarus with three or four ceraran setae; body with seven-
teen pairs of cerari; lateral cerari with three or four auxil-
iary setae; anal cerari borne on chitinized plate; dorsal setae
numerous and rather long; anal setae equal to or slightly
longer than anal ring setae. — Japan, New York, California on
Mulberry, maple, Pinus. comstocki Kwn.
i<t. Body with the cephalic three or four pairs and sixth pair of
cerari with each cerarus with three or four ceraran setae.
e. Anal cerari not borne by distinct more strongly chitinized
area,
f. Cerari without auxiliary setae.
g. Anal cerari with tricerores numerous, twenty-five or
more, immediately about their ceraran setae,
h. Body with fifteen to seventeen pairs of cerari,
cephalic two or three pairs of cerari with each
cerarus with three ceraran setae, other cerari
with two; tricerores about six in number about
lateral cerari, but few on body; anal setae and
anal ring setae subequal. — California on Cupres-
sus. Thuja, Libocedrus, Sequoia ryani Coq,
hh. Body with seventeen pairs of cerari, cephalic
four pairs of cerari with each cerarus with three
or four ceraran setae, other cerari with two; tri-
cerores about twelve to fifteen on cephalic and
caudal cerari, always more than six, fairly nu-
merous on the body; anal setae two-thirds length
of anal ring setae, latter distinctly longer than
diameter of anal ring; ventral surface of anal
lobes chitinized. — Illinois, Massachusetts on sor-
ghum, corn, nests of LABlvLB.^.sorghiclhu Forbea.
136 THE COCCIDAE
gg. Anal cerari with tricerores always few in number,
twelve or less, about their ceraran setae.
h. Body with seventeen pairs of cerari, cephalic
three or four pairs of cerari with each cerarus
with three ceraran setae, other cerari with two;
anal lobes with three or four slender setae; dorsal
setae few, very small; anal setae somewhat
shorter than anal ring setae. — California on
Sequoia, Cupressus. sequoiae Coleman.
hh. Body with less than seventeen pairs of cerari.
i. Body with sixteen pairs of cerari, cephalic
three or four pairs of cerari with each cerarus
with three or four ceraran setae, other cerari
with two.
j. Ceraran setae of cephalic pair of head cerari
located close together, less than their own
length apart; dorsal setae few, small and
slender; anal ring setae scarcely longer than
width of anal ring and about one-half length
of anal setae. — ^Arizona on Covillea
irishi Ckll.
jj. Ceraran setae of cephalic pair of head cerari
distant, located their own length or more
apart; dorsal setae few, small, and slender;
anal ring setae scarcely longer than width
of anal ring and about one-half length of anal
setae. — New Mexico on Covillea
steelii Ckll. & Towns,
ii. Body with fifteen pairs of cerari, cephalic two
or three pairs of cerari with each cerarus with
four or five ceraran setae, other cerari with
two; ceraran setae small and widely separated;
dorsal setae few, extremely few and slender;
anal ring setae somewhat longer than width of
anal ring and subequal in length to anal setae.
— Arizona on Juniperus juniperi Ehrh.
ff. Cerari with auxiliary setae.
g. Anal cerari with tricerores always few in number,
not forming crowded area about their ceraran setae;
body with fifteen to seventeen pairs of cerari, typi-
cally seventeen; cephalic three or four pairs of
cerari with each cerarus with three conical setae;
other cerari with two; all cerari with prominent
auxiliary setae; dorsal setae numerous and usually
long; anal setae and anal ring setae subequal. — Cali-
fornia on Castillea, Oribanche, Armeria
longisetosus Ferris.
gg. Anal cerari with numerous tricerores crowded into
a circular area about the ceraran setae; body with
SUBFAMILY ERIOCOCCINAE 137
seventeen pairs of cerari; cephalic two or three and
sixth pairs of cerari with each cerarus with three or
four ceraran setae, other cerari with two; all cerari
with numerous auxiliary setae; dorsal setae fey and
slender; anal setae shorter and more slender than
anal ring setae; anal ring with single dentaceroris
on each side; mesal orbacerores forming irregular
double row between cephalic anal ring setae; cisanal
setae farther apart than obanal setae. — California,
Oregon, Missouri, New York on tomato, carnation,
passion flower, maple, clover, cherry, Eriogonum,
Quercus, Magnolia, Crataegus, Pinus
maritimus Ehrh.
Anal cerari borne by distinct more strongly chitinized area.
f. Penultimate pair of cerari never borne on more strong-
ly chitinized areas; body with seventeen pairs of cerari,
cephalic three or four pairs of cerari with each cerarus
with three or four ceraran setae, other cerari with two;
lateral cerari with auxiliary setae; dorsal setae few
and slender; anal setae and anal ring setae subequal;
anal ring with single dentaceroris on each side; lateral
orbacerores forming irregular double row between me-
dian and caudal anal ring setae; cisanal setae distant
from anal ring and closer together than obanal setae. —
California on Passania, Quercus agrifoliae Essig.
ff. Penultimate pair of cerari always borne on more strong-
ly chitinized areas,
g. Anal cerari with their tricerores arranged in compact
group about anal ceraran setae; body with seventeen
pairs of cerari; cephalic three or four pairs of cerari
with each cerarus with three or four ceraran setae,
other cerari with two; all cerari with auxiliary setae;
dorsal setae few and slender; anal setae shorter than
anal ring setae; anal ring with two or three denta-
cerores on each side; cisanal setae adjacent to anal
ring and farther apart than obanal setae. — Pseudo-
coccus longispinus Targ. — United States, Jamaica,
Chili, Europe, Mai^itius, New Zealand on ferns,
mango, guava, fig, plum, Citrus, Croton, Cycas, Fla-
courtia, Nephrodium, Stangeria adonidum Linn.
gg. Anal cerari with their tricerores scattered, not ar-
ranged in compact group about anal ceraran setae;
body with seventeen pairs of cerari; cephalic three
or four pairs of cerari with each cerarus with three
ceraran setae, other cerari with two; dorsal setae
few and scattered; lateral cerari with auxiliary
setae; anal setae and anal ring setae subequal. — Cal-
ifornia on Citrus, Azalea, Rubus, Hedera, Ficus,
138 THE COCCIDAE
Malva, Brassica, Solanum, Shinus, Chenopodium,
Rheum, Rosa, Melilotus, Helianthus, Juglans, etc
citrophilus Claussen.
cc. Body always with less than twelve pairs of cerari.
d. Body with eleven pairs of cerari.
e. Cerari arranged as follows: pair cephalad of eyes, pair
near eyes, pair opposite prothoracic legs, pair opposite
mesothoracic spiracles, and seven pairs on abdomen, ce-
phalic and thoracic pairs sometimes wanting; each cerarus
usually with two ceraran setae; anal setae and anal ring
setae subequal. — California on Cupressus
cupressicolus Ferris,
ee. Cerari arranged as follows: four pairs on head and seven
pairs on abdomen; two cephalic pairs of cerari with each
cerarus with three ceraran setae, other cerari with two;
auxiliary setae associated with ultimate and penultimate
pairs of cerari only; dorsal setae few and small; anal ring
slightly chitinized; anal setae about one-half length of
anal ring setae. — California on Distichlis salinus Ckll.
dd. Body always with less than eleven pairs of cerari.
e. Body with eight or nine pairs of cerari counting cephalad
from anal cerari and usually with pair cephalad of eyes;
cephalic pair on head with several ceraran setae, other
cerari with two; anal cerari with auxiliary setae, lateral
cerari without; dorsal setae numerous, slender; anal setae
about twice as long as anal ring setae. — California on
Eriogonum, Viola, Stachys, Eriodictyon, Ceanothus, Erig-
eron. eriogoni Ehrh.
ee. Body with ten pairs of cerari.
f. Anal ring setae about equal in length to anal setae and
about one and one-half times as long as width of anal
ring; head with only ocular cerari, which contain three
or four ceraran setae, other cerari with two; auxiliary
setae present only among the anal cerari; dorsal setae
few and small. — New Mexico, Utah, California on
Gutierrezia. gutierreziae Ckll.
ff. Anal ring setae somewhat shorter than anal setae; anal
setae about one and one-half times as long as width of
anal ring; head with only ocular cerari, which contain
three or four ceraran setae, other cerari with two; anal
cerari only with auxiliary setae; dorsal setae small and
few in number. — New Mexico, Arizona on Prosopis
prosopidis Ckll.
Trionymus Berg. — This genus contains only two species according
to Fernald's Catalogue, perrisii Sign, from France, the type, and a second
species, americanus Ckll. from the District of Columbia. Ferris has pre-
. pared a synopsis of the Californian species. It is possible that several
of the species now referred to Pseudococcus belong here. The follow-
ing table is based upon the work of Ferris: —
Q
SUBFAMILY ERIOCOCCINAE 139
SPECIE'S OF TRIONYMUS
. Body never with more than single pair of cerari, anal cerari.
b. Anal cerari with their ceraran setae elongate and longer than
dorsal setae, never short and conical; antennae of adult female
consisting of seven segments; anal cerari with two or three auxil-
iary setae; anal ring not strongly chitinized; anal setae slightly
shorter than anal ring setae. — California on Distichlis
distichlii Ferris.
bb. Anal cerari with their cereran setae short and conical, usually
shorter than body setae.
c. Anal cerari with numerous tricerores grouped about their
ceraran setae, distance between them subequal to or less than
length of diameter of calices of ceraran setae.
d. Setae varying in size from small and slender to quite large
and stout, those on each lateral portion of eighth segment
subequal in diameter at proximal end to ceraran setae;
antennae of adult female consisting of seven segments; anal
cerari with two or three ceraran setae and several slender
auxiliary setae; anal setae somewhat more slender and
smaller than anal ring setae. — California on Grindelia
grindeliae Ferris
dd. Setae not varying in size, all very short and slender, sub-
equal, those on lateral portions of eighth segment never sub-
equal in diameter at proximal end to ceraran setae, always
slender; antennae of adult female consisting of seven seg-
ments; anal cerari with two ceraran setae and five to eight
auxiliary setae; anal setae slightly shorter than anal ring
setae. — Eastern United States on clover trifolii Forbes.
cc. Anal cerari with only few tricerores and always distant, dis-
tance between them always twice or more than length of diame-
ter of calices of ceraran setae,
d. Anal ring setae scarcely more than one-half length of anal
setae; antennae of adult female consisting of seven seg-
ments.
e. Body with dorsum with relatively few cerores and cylin-
drical ducts, none of ducts short, broad, and tubular; ab-
domen with two caudal segments with many short, slender,
curved setae, other segments with few. — California on
Elymus. smithii Es-.ig.
ee. Body with dorsum with few, scattered, small, slender
setae and numerous tricerores and short, broad, tubular
ducts with a narrow raised rim; abdomen with two caudal
segments not with many curved setae. — California on
Quercus. villosa Ehrh.
dd. Anal ring setae subequal in length to anal setae; antennae
of adult female consisting of eight segments; anal cerari
with two ceraran setae and two or three auxiliary setae;
dorsal setae few, short, slender, curved; anal ring small and
140 THE COCCIDAE
heavily chitinized. — California on Bromus, Ammophila,
Ericameria. bromii Ferris.
aa. Body always with more than one pair of cerari.
b. Anal cerari always situated upon distinct chitinized area bearing
numerous tricerores and auxiliary setae; antennae of adult female
consisting of eight segments; abdomen with two to four cerari at
caudal end, ceraran setae of anal cerari large and stout, of other
segments smaller; anal cerari surrounded by numerous tricer-
ores and eight to ten auxiliary setae; anal ring setae slightly long-
er than anal setae. — California on Elymus. califomicus Ehrh.
bb. Anal cerari never situated on chitinized area.
c. Body with three pairs of cerari, located at caudal end; ceraran
setae all short and cylindrical, those of anal cerari largest;
antennae of adult female consisting of eight segments; anal
setae distinctly longer than anal ring setae. — France on Cala-
magrostis. Sphagnum, Agropyrum, Festuca perrisii Sign.
cc. Body with two pairs of cerari located at caudal end of body,
d. Antennae of adult female consisting of eight segments;
ceraran setae short, stout, conical, not accompanied by
grouped tricerores or auxiliary setae; dorsal setae numerous,
short, subequal; anal ring large and heavily chitinized; anal
: setae slightly longer than anal ring setae. — Australia, Cali-
fornia on Calceolaria, Danthonia, Phormium, Cordyline,
sugar-cane. calceolariae Mask.
dd. Antennae of adult female consisting of seven segments;
ceraran setae small, conical, two in each cerarus; cerari not
with grouped tricerores or auxiliary setae; dorsal setae short,
more numerous on caudal part of body; anal ring small and
strongly chitinized; anal setae slightly shorter than anal
ring setae. — California on Festuca. festucae Kwn.
Tylococciis Newst. — This genus was based upon a single species,
madagascariensis Newst., from Madagascar from the nests of Cremas-
togaster. The body is provided with cephalabiae and caudalabiae and
with fifteen pairs of cerari. It is questionable if the other species re-
ferred to this genus by Brain belong to it.
Eriiuu Mask.— A genus containing five species. The type,
globosum Mask., is from Australia on Acacia. All the other species are
from America, they are: armatum Hemp, from Brazil; eriogoni Ehrh.
from California on Eriogonum; lichtensioides Ckll. from Colorado, Cali-
fornia on Artemesia; and zapotlanum Ckll. from Mexico on "Huele de
Noche." Ferris has referred irishii Ckll. and steelii Ckll. & Towns, to
Pseudococcus. He describes lichtensioides Ckll. as follows: antennae
consisting of seven segments, anal cerari only present, consisting of ten
ceraran setae borne on a chitinized area without auxiliary setae; dorsum
of thorax and abdomen with numerous short conical setae, similar to the
ceraran setae.
Lefroyia Green. — A single species, castaneae Green from Assam,
India on Castanea, is included.
Ripersiella Ckll. — This generic name was first published by Cock-
SUBFAMILY ERIOCOCCINAE 141
erell and should be credited to him. The type species, rumicis Maskell
is from New Zealand on Rumex. The other species are kelloggi Ehrh.
& Ckll. from California on roots of bunch grass; leucosoma Ckll. from
New Mexico from the nests of Lasius; and maritima Ckll. from Long
Island, New York on roots of Spartina.
Natalensia Brain. — This genus is based upon a single species,
fulleri Brain from Natal from roots of grass from nests of ants. By the
table this genus will run to Ripersiella Ckll. The circular form of the
body, the antennae consisting of six segments, and the anal ring bear-
ing six anal ring setae in nymphal and adult females, are characteristic.
The adult female is enclosed in a double-walled cyst without openings.
Bhizoecus Kunck. — An European genus of four species: eloti
Giard from roots of coffee-tree; falcifer Kiinck. from roots of palms and
vines; (?)targionii Ckll. on Mammillaria; and (?)terrestris News, on
roots of Stephanotis.
Bipersia Sign. — This genus is of indefinite limits. The following
American species have been referred to it: aurantia Ckll. from New
Mexico from nests of Lasius; blanchardii King & Ckll. from Massachu-
setts from nests of Lasius; cockerellae King from New Mexico from
nests of Lasius; fimbriatula Ckll. & King from New Mexico from nests
of Lasius; flaveola Ckll. from Massachusetts, New Mexico from nests
of Lasius; kingii Ckll. from Massachusette from the nests of ants; lasii
Ckll. from Canada, Massachusetts on roots of asters and from the nests
of ants; magna T. & W. Ckll. from New Mexico; minima Tins. & King
from Massachusetts from nests of Lasius; myrmecophila Mask, from
Michigan; porterae Ckll. from New Mexico on the roots of grass; salmon-
acea Ckll, from New Mexico on roots of grass; serrata Tins, from Trini-
dad; sporoboli Ckll. from New Mexico on Sporobolus; tenuipes Ckll.
from New Mexico on the roots of grass; trichura Ckll. from New Mexico
from the roots of grass; trivittata Ckll. from New Mexico from nests of
Lasius; villosa Ehrh. from California on Quercus; and viridula Ckll.
from New Mexico from nests of Lasius.
Cryptoripersia Ckll. — Two species have been referred to this
genus, Eriococcus salinus Ehrh. from California on Distichlis and Riper-
sia arizonensis Ehrh. from Arizona, California on Elymus. Ferris has
examined the types of both species and has pronounced them identical.
Fonscolombia Fonsc. — There are two European species: fraxint
Kalt. on Fraxinus and radicum-graminia Fonsc, the type, on roots of
grass and grain and two American species: braggi Ckll. & Robs, from
Colorado on Berberis and yuccae Ferris from New Mexico on Yucca. In
yuccae there are numerous cerores in the region of the vulva which are
wanting in braggi.
Termitieoccus Silv. — Two species, aster Silv. from Paraguay and
bicornis Silv. from Brazil, collected in the nests of termites, not from
ant's nests as recorded in Fernald's Catalogue, belong to this genus.
Pseudoripersia Ckll. — A genus including a single species, turgipes
Mask, from Australia on Casuarina.
Gymnococcus Dougl. — The type of this genus is agavium Dougl.
from Europe. The genus contains two American species found upon
grass, nativus Parr, from Kansas and ruber Parr. & Ckll. from New
142 THE COCCIDAE
Mexico. These species can be separated by the following table prepared
by Parrott. It should be noted that the American species will not run
to Gymnococcus in the table to the genera of Eriococcinae.
SPECIES OF GYMNOCOCCUS
a. Cuticle of adult female with numerous short conical setae. — Europe
on Agave. agavium Dougl.
aa. Cuticle of adult female not bearing conical setae.
b. Thoracic spiracles with cerores forming complete ring about
them. — New Mexico on grass. ruber Parr. & Ckll.
bb. Thoracic spiracles with cerores limited to one side, not forming
complete ring about spiracle. — Kansas on grass nativus Parr.
Geococcus Green. — A single species, radicum Green from Ceylon
upon the roots of grass, is included.
Porococcus Ckll. — There are two species from Mexico included,
occurring on mistletoe on oak; pergandei Ckll. and tinctorius Ckll.
Ooccura Sulc. — A genus erected for Phenacoccus comari Kiinow
from Germany on Comarum by Sulc. Ferris believes that the American
Phenacoccus stachyos Ehrh. is congeneric.
Macrocerococcus Leon. — This genus includes a single species,
superbus Leon, from Italy, Corsica, and France on Lepidum, Alyssum,
lotus, Fumaria, Cetarach, Antirrhinum, and undetermined grass.
Macrocepi coccus Morr. — A single species, loranthi Morr., the type
of the genus, described from British Guiana. The cerari consist of a
chitinized area bearing from one to four ceraran setae.
Tetrura Licht. — This genus is omitted from the table. The
description of the genus and species is too indefinite to permit of its
location. It would run to Phenacoccus by the table. There is a single
species, rubi Licht. from Europe on Rubus.
Heliococcus Sulc. — A single species included, bohemicus Sulc
from Bohemia and Moravia on Robinia.
Phenacoccus Ckll. — There are thirty-four species referred to this
genus in Fernald's Catalogue. Of this number sixteen are recorded from
America, some of these have since the publication of the Catalogue been
referred to the synonomy. The following species, other than those in-
cluded in the table, should be noted. They are: americanae King &
Ckll. from Massachusetts from the nests of Lasius; cevalliae Ckll. from
New Mexico on Cevallia; cockerelli King from Colorado on Amelanchier;
gossypii Towns. & Ckll. from Mexico on cotton, other Malvaceae, wild
guava; minimus Tins, from Colorado on Picea; rubivorum Ckll. from
New Mexico on Rubus; ripersioides W. & T. Ckll. from New Mexico from
nests of Lasius; simplex King from California on Atriplex; solenopsis
Tins, from New Mexico from nests of Solenopsis; spiniferus Hemp, from
Brazil; and wilmattae Ckll. from New Mexico on Viola.
SPECIES OF PHENACOCCUS.
a. Body with many short, stout, cylindrical ducts. — California on
lichens on Picea. kuwanae Coleman.
aa. Body not with many short, stout, cylindrical ducts,
b. Anal cerari with only two ceraran setae.
SUBFAMILY ERIOCOCCINAE 143
c. Head cerari and all other cerari of body never with more than
two ceraran setae.
d. Body with eighteen pairs of cerari; auxiliary setae want-
ing; tricerores few and scattered; anal lobes sometimes with
single seta, sometimes Mrith two or three small setae in addi-
tion to ceraran setae; dorsal setae small, sharply pointed;
anal setae about one and one-half times as long as the anal
ring setae; antennae consisting of eight segments. — Cali-
fornia, Utah on roots of Hemizoonia, potato, tomato, wild
radish, Malva, pansies, wild sunflower, aster, nightshades,
purslane. solani Ferris.
dd. Body with fifteen or sixteen pairs of cerari; cerari with
small but distinct clusters of tricerores and without auxil-
iary setae; ceraran setae large, subequal in size, stout, coni-
cal, with distal portion slightly flaggellate; antennae consist-
ing of six to eight segments; anal ring setae about twice
as long as diameter of anal ring and much longer than anal
setae. — Pseudococcus lycii Ferris. — ^Arizona on Lycium.
lycii Ferris.
CO. Head cerari, at least one pair, with three to four ceraran setae,
d. Anal setae slightly shorter than anal ring setae; cephalic
three or four pairs of cerari with three or four ceraran
setae, other cerari with two, all cerari obscure and without
auxiliary setae; dorsal setae minute and few in number;
tubular ducts few, without raised rim. — California oti
Artemisia. artemisiae Ehrh.
dd. Anal setae about one and one-half times as long as anal ring
setae; ocular cerari with three to four ceraran setae, other
cerari with two, all without auxiliary setae; anal lobes large
and prominent; dorsal setae few and extremely small; tubu-
lar ducts with their mouths borne at apical end of conical
projections, projections surrounded by one to four small
setae; tubular ducts arranged two to three near each cerar-
us, irregular double row near the meson, and indefinite
number on head. — Phenacoccus osborni Sanders. — Phenacoc-
cus pettiti Hollinger. — California, Ohio, Missouri on Stachys,
Monardella, Diplacus, Solanum, Toxicodendron, Catalpa,
Sycamore. stachyos Ehrh.
bb. Anal cerari always with more than two ceraran setae.
c. Anal cerari with three or four ceraran setae, also cephalic
three or four pairs of cerari, all other cerari with two ceraran
setae; cerari without auxiliary setae; body setae few, scat-
tered, and small; anal setae nearly twice as long as anal ring
setae. — California on Rubus, Fragaria, Symphoricarpus, Cas-
tilleia, Eriophyllum, Geranium. eolemani Ehrh.
cc. Anal cerari always with more than four ceraran setae.
d. Anal setae nearly twice as long as anal ring setae; anal
cerari with six to ten ceraran setae, second pair from caudal
end with six to ten, third with two and several other setae
144 THE COCCIDAE
scattered about, all other cerari with two; cerari with auxil-
iary setae; dorsal setae shaped like ceraran setae and as
large as smallest of them; anal ring setae longer than
diameter of anal ring. — Phenacoccus betheli Ckll. — Canada,
Michigan, Illinois, Arizona on Amelanchier, Crataegus.
dearnessi King.
dd. Anal setae and anal ring setae subequal, former never twice
as long as latter.
e. Anal cerari with about ten ceraran setae, all other cerari
with three to six, these setae slender and sharply pointed;
cerari without auxiliary setae; tricerores of cerari few,
scarcely exceeding setae in number; dorsal setae almost
lacking and extremely minute; anal setae equal to or trifle
shorter than anal ring setae. — California on Eriogonum
eriogoni Ferris,
ee. Anal cerari with ten or more ceraran setae and numerous
scattered tricerores; penultimate pair of cerari with five
to eight ceraran setae and numerous tricerores; cephalic
two or three pairs of cerari with four to six ceraran setae
and eight to ten tricerores; all other cerari with two
ceraran setae and cluster of eight to ten tricerores; cerari
without auxiliary setae; dorsal setae few, small, in part
slender and in part like ceraran setae; anal ring setae
somewhat longer than diameter of anal ring and about
equal in length to anal setae. — New Mexico, Arizona, Cal-
ifornia on Helianthus, Pluchea, Solanum, Baccharis,
Ptiloria. helianthi Ckll.
Heterococcus Ferris. — A genus containing a single species, arenae
Ferris from California on Poa.
Ceroputo Sulc. — The type of this genus is pilosellus Sulc from
Europe. The other species are from America, as follows: ambigua Full-
away from California on Salicornia; bahiae Ehrh. from California on
Bahia; barberi Ckll. from Antigua, New Mexico on Thunbergia, Alla-
manda, Coleus, Croton; calcitectus Ckll. from New Mexico on grass;
koebelet Ehrh. from Arizona on Quercus ; lassiorum Ckll. from New Mexico
from nests of Lasius; orthezioides Ckll. from Mexico on roots of dock-
weed; yuccae Coq. from Antigua, Mexico, California on Yucca, Lantana,
Mimulus, Ceanothus, banana, orange, lime, etc. Ferris writes as follows
on this genus: "I consider Ceroputo to be strictly a synonym of Puto.
The former was separated from the latter solely on the basis of the pres-
ence of but six hairs on the anal ring and of non-clubbed digitules."
Ehrhornia Ferris. — This genus contains two species, cupressi
Ehrh. on Cupressus and graminis Ferris on grass. Both species are from
California.
Trabutina Marchal. — A single species is included, elastica Marchal
from Algeria on Tamarix.
Puto Sign. — The type of the genus is antennata Sign, from France
on pine. There is a described American species, the Pseudococcus cu-
SUBFAMILY ERIOCOCCINAE 145
press! Coleman from California on Cupressus, Pinus, Torreya, and Se-
quoia.
Khizococcus Sign. — This genus includes about twelve described
species, one from France, gnidii Sign, on roots of Daphne, is the type;
one from Texas, texanus King from the nests of Cremastogaster ; and all
the other species are from the Australian region.
Eriococcns Targ. — This genus contains over sixty species. The
following have been recorded from America: adenostomae Ehrh. from
California on Adenostoma; araucariae Mask, from New Zealand, Ceylon,
Hawaiian Islands, South Africa, California on Araucaria, Kunzea;
arenosus Ckll. from New Mexico; armatus Hemp, from Brazil on Bac-
charis; artemisiae Knw. from California on Artemisia; aurescens Ckll.
from Mexico on "Guasima;" azaleae Comst, from Massachusettes, New
York, District of Columbia, Ohio, Indiana, Michigan on Azalea, Cratae-
gus, Rhododendron; bahiae Ehrh. from California on Bahia, Gutierrezia;
borealis Ckll. from Alaska on willow; brasiliensis Ckll. from Brazil on
Baccharis; catalinae Ehrh. from California on Artemisia; coccineus
Ckll. from Nebraska on Cactus; diversispinus Leon, from Argentina on
Zaccagnia; dubius Ckll. from Mexico; eriogoni Ehrh. from Arizona on
Eriogonum; gilletti Tins, from Colorado on Juniperus; howardi Ehrh.
from California on Quercus; kemptoni Parr, from Kansas on Andropo-
gon; larreae Parr. & Ckll. from New Mexico on Larrea; multispinosus
Kuhlgatz from South America on Cactus; neglectus Ckll. from New
Mexico on Atriplex; palmeri Ckll. from Carmen Island, Lower California
on Bourreria, Eriogonum; parcispinosus Leon, from Argentina on Atri-
plex; perplexus Hemp, from Brazil on Myrtaceae; quercus Comst. from
Massachusette, Florida, New Mexico, Arizona, Mexico on Quercus, gall-
berry, Vaccinium, Prosopis, grass; salinus Ehrh. from California on
Distichlis; and tinsleyi Ckll. from New Mexico on Atriplex, Malvastrum,
Gutierrezia.
Gossyparia Sign. — This genus contains five species, three from
the Australian region and two from Europe. One of these latter, spuria
Modeer on elm, has been introduced into the United States.
liachnodius Mask. — A genus containing three described species
from Australia on Eucalyptus and Acacia.
Iiachnodiella Hempel. — A single species, cecropiae Hempel from
Brazil on Cecropia, is included. The name of this genus and species
was first offered in print by von Ihering, but no description of either was
given by this author.
Micrococcus l»eon. — A single species, silvestri Leon, from Sardinia
from the nests of Tapinoma, is included.
Atriplicia Ckll. & Rohw. — A genus of gall making eriococeida
containing a single species, gallicola Ckll. & Rohw. from New Mexico and
Colorado on Atriplex.
Tectococcus Hemp. — There is a single species, ovatus Hempei
from Brazil on Myrtaceae.
Cissococcns Ckll. — A single species, fuller! Ckll. from Soatk
Africa on Cissus is included. Ehrhorn has referred oahuensis Ehrh. from
the Hawaiian Islands on Urera doubtfully to this genus.
Antonia Sign. — The American species of this genus can he se#«-
arated by means of the following table: —
146 THE COCCIDAE
SPECIES OF ANTONIA
a. Anal ring located at cephalic end of invaginated anal tube; an-
tennae of adult female consisting of three segments; caudal end of
abdomen usually strongly chitinized.
b. Abdomen with caudal end not strongly chitinized or not decided-
ly more strongly chitinized than adjacent parts; adult female
about three times as long as broad; metathoracic spiracles located
near middle of length of body, mesothoracic spiracles midway
between metathoracic spiracles and antennae, and rostrum mid-
way between mesothoracic spiracles and antennae; spiracles with
group of about twenty-five closely placed cerores adjacent to their
lateral margin; anal lobes wanting; caudal end of body on each
side of anal tube with three long setae. — England on recently im-
ported plants of Arundinaria japonica. socialis Newst.
bb. Abdomen with caudal end strongly chitinized on ventral or both
dorsal and ventral aspects.
e. Adult female, untreated, oval in form and cream-coloted ;
caudal portion of abdomen strongly chitinized on ventral as-
pect and dark in color; abdomen on caudal segments bearing
numerous small slender setae; spiracles surrounded on lateral
margin by crescentic-shaped group of cerores; first stage
nymph with transverse row of short conical setae on middle
of each segment. — Kansas on Bouteloua houtelouae Parr.
ee. Adult female, untreated, oval or circular in outline and deep
purplish-red in color; caudal portion of abdomen quite heavily
chitinized and bearing numerous short, spine-like setae, other
portions of body with only few setae at margin, first stage
nymphs with g.roup of two or three short setae at lateral mar-
gin of each of three last abdominal segments. — Japan, Californ-
ia on bamboo crawii Ckll.
aa. Anal ring not located at cephalic end of invaginated anal tube;
antennae consisting of three to five segments; caudal end of abdomen
»ot strongly chitinized.
li. Antennae short, with four or five segments, the two proximal seg-
ments are usually about the same width and distal segments
taper sharply; prothoracic legs sometimes present; body with few
short setae, more numerous on caudal portion; spiracles with
erescentric group . of about forty closely placed cerores, situated
»ear lateral margin of spiracle. — Kansas on Eragrostis, Bulbilis,
Paspalum. parrotti Ckll.
IWi. Antennae aborted, never with more than three segments.
«. Body, exclusive of caudal segment, bearing large number of
setae; spiracles colored and accompanied by abundant punctu-
ation.— France on stubble and roots of certain Graminaceae,
principally Milium, Agropyrum. purpurea Sign.
t«. Body not bearing large number of setae; adult female oval,
plump, cream-colored, with slight tinge of brown on margin;
spiracles on lateral margin with crescentic group of closely-
placed rather large cerores; anal orifice situated in »llgkt d«-
SUBFAMILY ERIOCOCCINAE 147
pression; anal area with many slender setae, much smaller than,
anal ring setae. — Kansas on Bouteloua nortoni. Parr & Ckll.
Sphaerococcus Mask. — Maskell erected his group Idiococcidae for
the inclusion of this genus and the genus Cylindrococcus. The anal ring
has been poorly studied in most of the species of Sphaerococcus and in
the descriptions of several is not even mentioned. Some of them have
already been removed and made the types of other genera. The type
species of the genus Sphaerococcus, casuarinae Mask., has according to
the describer an anal ring with anal ring setae in the second nymphal
stage and probably also in the first nymphal stage although the anal
ring is not mentioned in the description of this stage. Ferris, who has
studied this species, reports the presence of an anal ring with six anal
ring setae in the adult female and first nymphal stage. In the adult
female the anal ring is located at the cephalic end of an invaginated
anal tube. The cephalabiae and caudalabiae are present in the first
nymphal stage. The presence of an anal ring with anal ring setate
would debar this genus from the Idiococcidae, the Cylindrococcinae of
the present work. A single species, Sphaerococcus sylvestris Ckll. &
King from Massachusetts on white oak, has been recorded from America.
The species of Sphaerococcus fall into two groups, those forming true
galls, apparently always bud galls, or blisters on their host-plants and
those not forming abnormal growths but in which the body of the adult
female is covered with white cottony or woolly wax or are enclosed in a
hard test of wax. The nineteen species listed in Fernald's Catalogue
represent several generic types. Ferris has made disticlium Knw. the
type of Paludicoccus, pirogallis Mask, the type of Eremicoccus, and
pulchellus Mask, the type of Callococcus. He has referred obscuratus
Mask, to Kuwanina and leptospermi Mask, to Amorphococcus and states
"that Sphaerococcus sylvestris Ckll. & King is probably nothing more
than an immature stage of some species of Kermes."
Paludiicoccus Ferris. — A single species, Sphaerococcus disticlinm
Knw. from California on Distichlis, is the type and sole species of the
genus.
Nidularia Targ. — An European genus containing a single spescles,
pulvinata Planch. The cephalic end of the cyst bears two transverse
rows of small conical protuberances. The body of the adult female rests
upon an ovisac similar to that of Pulvinaria.
Kuwanina Ckll. — The type of this genus, Sphaerococcus parms
Mask, from Japan on cherry, was the sole species until the reference of
obscuratus Maskell to this genus.
Cryptococcus Dougl. — This genus contains a single species, fagi
Baer. from Great Britain, France, Belgium, Germany, Bohemia, and
Switzerland on birch.
Kermiciis Newst. — An Indian monotyplcal genus inel»dlBg
wroughtoni Newst. from the nests of an ant, Oecophylla.
Chaetococcus Mask. — A single species, bambusae Mask, oetmr^ ^
the Hawaiian Islands, Mauritius, Ceylon, and Braail on Bambufw..
CHAPTER XIV
SUBFAMHiY TACHARDIINAE
The Lac Insects
The body of the adult female Is typically conical or subconical or
subglobose with the mouth-parts located at the pointed end of the cone
and the anus at the other. Body always irregular in form, bag-like.
It is impossible to identify the lateral margins and the aspects are not
well defined. The caudal end is never inflated with the cephalic and
lateral margins keeled. The antennae are minute, vestigial, consist of
three or four segments, and are articulated to the ventral aspect of the
head. The eyes are wanting. The mouth-parts, rostrum and rostralis,
are always present, the rostrum consists of two segments. The legs are
wanting. The metathoracic spiracles are normal in size and ventral in
position. The mesothoracic spiracles are large and distinct, always larg-
er than the metathoracic spiracles, and are located near prominent stig-
matic processes which are always located near the anus, the distance
from the anus varying with the species. The abdominal spiracles are
always wanting. The stigmatic clefts and spinae are wanting. The
lateral surface of each stigmatic process bears a canella and spiracerores.
The anus is located at the end of a prominent anal process. There is
usually an anal spine associated with the stigmatic and anal processes.
The anus is surrounded by a prominent anal ring bearing about ten anal
ring setae. The body through the migration of the mesothoracic spiracles
is greatly distorted, so that the abdomen appears to be two or more times
as long as the head and thorax together. The abdomen is bag-like or
Jug-like with three necks. The lateral margins are wanting. All of the
segments are fully exposed, none of the coriae between them can be
identified, and none of the segments are retracted to form a marsupium.
The abdomen is never provided with an anal cleft and opercula or anal
lobes and anal setae. The anal ring is always distinct, located on the
anal process, and provided with ten anal ring setae located on six plates.
The caudal abdominal segment can not be identified. The body, except
the stigmatic and anal processes, is not provided with cerores and none
of these are octacerores or pilacerores. The body is never provided with
ceratubae. The body is always enclosed in a resinous cell with three
adjacent openings, small threads of wax often protruding from the three
openings. The rectum is never provided with a long glassy tube of wax
or with a chitinized rectal tube bearing rings of anacerores. The body is
never naked and gall-like in form. It is never covered or associated with
mealy or powdery or cottony wax.
The female nymphs of all stages are provided with a rostrum and
rostralis. The three pairs of legs are present in the first nymphal stage
148
SUBFAMILY txVCHARDIINAE 149
iand Wanting in all others, and when present are similar in form. The
profemora are never greatly enlarged and the prothoracic legs are never
fitted for digging. The body is never provided with an anal cleft and
opercula, anal lobes and anal setae, octacerores, pilacerores, or ceratiihae.
The anal ring is distinct, located on an anal process in the second
nymphal stage and bears six to ten anal ring setae. The caudal end of
the rectum is not chitinized, forming a rectal tube provided with one or
more rings of anacerores excreting a long glassy tube of wax. The
caudal abdominal segment is not short, narrow, and projecting.
The male has four ocellanae, two dorsal and two ventral. The ab-
domen is not provided with long lateral filaments and bears two long
slender caudal threads of wax. The wings are present and fully formed
or wanting. The stylus is distinct, about one-half the length of the
abdomen.
This subfamily includes the lac insects of commerce all of
which originally belonged to the genus Tachardia and previously
to the genus Carteria. There are probably few of the people who
are constantly using shellac who are aware as to how and where it
is obtained and that it is an excretion of a minute insect. The
wax or lac as it is usually called, from which is derived the shellac
of commerce, is excreted by the lac insect, Tachardia lacca. Its
chief home is the forests of India, Burma, and Assam. While the
best quality of lac comes from Burmah and Assam, lac is also
produced in Ceylon, Siam, China, and some of the islands of the
East Indies. There are about twenty-five thousand tons collected
annually in the central provinces of India. This lac has a valuation
of about a million and a quarter dollars. All of the lac of commerce
is produced on the forest trees in a wild state and no effort is made
to produce it artificially or to cultivate it as is done in the case
of the cochineal insect. The lac insect lives upon a large number
of species of trees, at least sixty according to Froggatt, the most
of which belong to the genus Ficus, fig.
The following account as to the preparation and use of lac
is taken from Froggatt. ''The different kinds of lac are known
in commerce under many names. Stick lac is the natural produc-
tion encrusting the twigs just as it is obtained from the forest;
seed lac is the stick lac after it has been ground up in water to
extract the coloring matter out of the insect, known as lac dye;
button and shell lac are prepared from the seed lac by melting the
latter. Other forms, such as garnet and liver lac, are produced
from different qualities of shellac, the color often differing consid-
erably in the various districts, while some of the fine bright orange
150 THE COCCIDAE
shellac is said to be artificially colored with orpiment. Lac is
extensively used for making the finer kinds of sealing wax, and
is the chief ingredient in most of the wood polishes, besides the
regular lacquer varnish used to coat the boxes, cabinets, and toys
known as lacquer ware used so much in China and India. The
lacquer ware from Japan, however, is polished with a varnish
made from the sap of one of the sumac trees (Rhus venix) and not
from the insect lac. ' ' The coloring matter secured from the bodies
of the lac insect is very similar to that obtained from the cochineal
insect. Species of the genus Tachardia are found in most parts
of the world. The following description is based upon the
admirable account by Green of the development of two species of
Tachardia, fici and albizziae. While it is quite likely that other
species will differ somewhat in minor details from this account,
yet the discrepancy is not likely to be great.
The nymphs of the first stage have a subdepressed body, are
elongate oval in outline with well developed legs and antennae and
an anal ring with six anal ring setae. They resemble somewhat
in general outline a young mealy-bug, but differ from these insects
in that their body is naked, not covered with mealy wax, and are
bright red or yellow in color. The antennae consist of six seg-
ments, of which the third is greatly elongated and enlarged on the
distal half, also the distal segment or sixth, as well as the fifth,
bears two very long slender setae. The rostrum is large and
conspicuous. The legs are distinct and normal in form. The anal
ring is large and each of the six anal ring setae is attached to a
small circular plate. The thoracic spiracles are small and incon-
spicuous. There is apparently a distinct canella with spiracerores
associated with each mesothoracic spiracle, but no such structures
are connected with the metathoracic spiracles.
The young nymphs soon begin to excrete wax. If one that is
two or three days old, is examined, it will be found that the entire
body is completely encased in a thin homogenous sheet of wax.
Green states that this wax first appears in the form of plates which
with increase in size coalesce. The form and general distribution
of the wax and the absence of cerores on the surface of the body
has led to the suggestion that the wax is the product of all the
hypodermal cells excreting together and not of certain cells con-
nected with cerores as in most coccids. A histological study of
these cells has not been made so far as I am aware.
SUBFAMILY TACHARDIINAE 151
The female nymphs at the first molt reduce their legs and
antennae to mere vestiges. The body is more swollen and
there is a prominent furrow on each side in line with the rostrum,
which gives the body a lobed appearance. There is a canella
extends obliquely from the former ventral position of each meso-
thoracic spiracle onto the dorsal aspect, where the mesothorac
spiracles are now located, due to the rearrangement of the folds of
the body. Each of these canellae bears numerous cerores or
spiracerores. The metathoracic spiracles are normal in position
on the ventral aspect and no canellae or spiracerores are associated
with them. The anal ring bears ten anal ring setae, each arising
from a cuticular plate with a tendency for eight of the plates to
fuse into pairs. The portion of the body bearing the anal ring is
prolonged and forms the beginning of the anal process.
The test or scale of the second stage female nymph is charac-
teristic in form. As viewed from above, it consists of lobes, three
on each side. There is at one end a prominent opening on the
meson of the dorsal aspect, this is the caudal opening and the one
in which the anal process fits. There is also a prominent opening
on each side of the dorsal aspect near the furrow marking the
caudal limit of the cephalic lobes. A stigmatic process fits into
each of these openings.
The adult females are easily recognized in most species, because
they bear a prominent anal spine. This is a pointed prolongation
of the cuticle which is located cephalad of the anus on the dorsal
aspect. The spine is sometimes placed on a fleshy tubercle, a
further prolongation of the cuticle of the body. The function of
the anal spine or dorsal spine as it is sometimes called, is unknown.
There are three stages of development through which the
early adult female passes, the early adult female or before gesta-
tion, the adult female during gestation, and the old adult female
or after gestation or after the eggs are laid.
The early adult female is small and similar in general form
to the female of the second stage. It differs in that the lobes are
more pronounced and the body is wider. The stigmatic processes,
the projections which bear the mesothoracic spiracles, are only
slightly elevated above the general surface of the body but the
canellae and spiracerores are distinct. The spiracles retain their
dorsal position and open dorso-laterad. The anal process is large
and prominent, the end is truncated, and the truncated portion
152 THE coccroAE
bears the anal ring and the ten anal ring setae located on six
cuticular plates. These plates bear numerous large cerores. The
cuticular periphery of the anal ring is bounded by projecting
spine-like processes.
The body of the adult female during gestation is larger and
more swollen from the numerous eggs which it contains. It becomes
so distorted that the position of the parts are considerably altered.
The metathoracic spiracles are small and retain their normal
position on the ventral aspect near the rostrum. The mesothoracic
spiracles are large and conspicuous and are located at one side of
the proximal end of the stigmatic processes. These processes with
the distortion and remoulding of the body are elongated and
brought nearer to the anal spine and the anal process. There is
a furrow, a canella, extends from the spiracle along one side of the
stigmatic process to its distal truncate end. Each canella bears
numerous spiracerores and its distal end which is continuous on one
side with the canella bears several large cerores with small cerores
or spiracerores mixed among them. The stigmatic processes were
originally known as the lac tubes from the mistaken notion that all
the wax of the test was excreted by them. The anal process also
becomes more prominent with the swelling of the body. The
stigmatic processes with the mesothoracic spiracles, the anal spine,
and the anal process are now all near together at one end of the
body. The migration in position of the spiracles is probably due
to the stretching of the cuticle of the body. The body of the adult
female during gestation completely fills the cell in the wax. As
the eggs are extruded, the body gradually shrinks and the empty
space in the cell about the body of the female is packed with eggs.
The young nymphs, when they hatch, emerge through the openings
in the cell in which the anus and mesothoracic spiracles fit. The
lobed appearance of the body is lost after the female deposits her
eggs, and, when it is expanded, is bag-like in form.
The waxy test formed by the female varies in size and form
with the species. In some the females are isolated, each female
forming a test; in others only a few females, three or four, are
found in the same globule of wax and contribute toward its forma-
tion; or a considerable number of females, twenty to thirty or a
large number, several hundred, form a large thick mass completely
encrusting a limb as in the stick-lac of India. The shape of the
ceU varies also somewhat with the species. The cell of the adult
SUBFAMILY TACHARDUNAE 153
female is always provided with three openings, one the anal
aperture, is slightly larger than the other two, the stigmatie
apertures. The apertures are arranged in the form of a triangle,
the anal aperture located at the apex of the triangle. The
stigmatie and anal processes fit into their respective apertures.
The position of the anal spine between these processes would
suggest that its function may be to hold the processes in place in
the apertures. The wax lining each aperture is different in
appearance and consistency and is undoubtedly excreted by the
cerores of the stigmatie and anal processes. The spiracerores
excrete a fine pulverulent wax or tufts of filaments of wax which
extrude from the stigmatie apertures sometimes almost closing
them. Green has suggested that this wax may function to prevent
the entrance of moisture.
The test of the male is oblong oval in outline, slightly
constricted near the oval caudal aperture and the dorsum is
supplied with a distinct median carina. The aperture is closed
during the quiescent stages with wax. The wax or lac of the male
is similar in appearance to the lac formed by the female. The
male tests are always formed by single individuals.
The nymphal stages of the male have not been described.
The adult male is ordinary in form, some individuals are winged
while others even in the same species may be wingless. The caudal
end of the abdomen bears a pair of long opaque filaments, which
are probably formed of wax. The head is provided with four
ocellanae, two dorsal and two ventral. The antennae of the
winged forms are provided with ten segments and of the wingless
forms with nine. **The genital sheath is elongate, slender, and
sharply pointed, rather more than half as long as the abdomen. ' '
This subfamily contains three genera, two of which were
originally described as subgenera, which can be separated by
means of the following table : —
GENERA OF TACHARDIINAE
a. Insects enclosed in masses of wax surrounding twigs, never in indi-
dividual tests; body of female elongate and vasiform; abdomen with
anal spine. Tachardia Sign.
aa. Insects enclosed in small masses of wax, not completely surround-
ing twigs, individuals may form separate test; body of adult female
more or less globular.
b. Abdomen of the adult female with anal spine; wax reddish or of
resin color. _ Tachardiella Ckll.
154 THE COCCIDAE
bb. Abdomen of adult female without anal spine; wax whitish, not
reddish. ■ Tachardina Ckll,
Tachardia Sign. — This generic name has been restricted to those
species forming a large mass of wax. The lac insect of India, Tachardia
lacca, is the type of the genus. It is found on fig, banyan, Rhamnus,
Mimosa, Anona, etc.
Tachardiella Ckll. — This genus contains the greater number of
the species, over thirty from all parts of the world. The following spe-
cies have been described from America: — caerulea Hemp, from Brazil;
cornuta Ckll. from New Mexico and Mexico on Parthenium; cydoniae
Hemp, from Brazil on Cydonia; cordaliae Leon, from Argentina on
Cordalia; argentina Dominguez from Argentina on Acacia; fulgens Ckll.
from Arizona and Mexico on Mimosa, Prosopis, Coursetia; fulvoradiata
Ckll. from Mexico on "Palo de gusano"; gemmifera Ckll. from Jamaica
on Chrysobalanus ; glomerella Ckll. from New Mexico on Gutierrezia;
ingae Hemp, from Brazil on Inga; larreae Comst. from southwestern
United States and Mexico on Larrea; lycii Leon, from Argentina on
Lycium; mexicana Comst. from Mexico on Mimosa; nigra Towns. & Ckll.
from Mexico on Acacia; parva Hemp, from Brazil on Myrtaceae; pustu-
lata Ckll. from Arizona; rosae Hemp, from Brazil on Rosa; rotundata
Towns. & Ckll. from Mexico on "Zicna, Guasima"; and rubra Hemp,
from Brazil on Cydonia and Croton. The Gascardia madagascariensis
Targ. is a species of Ceroplastes and the Colobopyga magnani Brethes
was evidently referred to this subfamily through an oversight.
Tachardina Ckll. — This genus contains a single species, albida
Ckll., which lacks an anal spine and has white lac. It was described
from Natal on Mimosa.
CHAPTER XV
SUBFAMILY LECANIINAE
The Tortoise Scales
The body of the adult female is variable in form, usually distinctly
longer than broad, sometimes almost circular, frequently convex on the
dorsum and concave on the venter, helmet-like in shape and outline.
The lateral margins are frequently thin and distinct, if so, usually
fringed with marginal setae, margin sometimes wanting or obscure.
The segmentation is usually obscure and the coriae are frequently want-
ing. The caudal end of the body is not inflated and the cephalic and
lateral margins are not provided with a keel. The antennae, when
present, are normal in form, usually small, and consist of seven or
eight segments. The number of segments and their size may be greatly
reduced, sometimes wanting. They are articulated to the ventral aspect
of the head, usually distant from each lateral margin. The eyes are
present. There is a single ocellana located on each lateral margin near
the articulation of an antennae. The mouth-parts, rostrum and rostralis,
are always present, the rostrum consists of a single segment. The legs
are usually present, they are sometimes rudimentary or wanting, when
present, they are subequal in length and normal in form. The profe-
mora are never enlarged, and the prothoracic legs are never fitted for
digging. The trochanto-femoral and tibio-tarsal sutures of all normally
formed legs are always distinct. The spiracles of the mesothorax and
metathorax are normal in form and size and ventral in position. The
mesothoracic spiracles are never located near the anus. The abdominal
spiracles are always wanting. The ventral aspect of the thorax is pro-
vided with spinae and canellae and usually with stigmatic clefts. The
abdomen is about as long as the head and the thorax together. It is fre-
quently convex on the dorsal and concave on the ventral aspect, some-
times convex on both aspects. The lateral margins are generally sharp-
ly defined, sometimes obscure or wanting. The caudal end is broadly
rounded or the caudal portion is produced and, whether rounded or
produced, is provided with a distinct mesal cleft. All of the segments
are exposed or at least no part of the abdomen is retracted to form a
marsupium. The abdomen is always provided with an anal cleft, some-
times apparently wanting, due to the fusion of the margins of the cleft.
The opercula are rarely if ever wanting. The anal ring is always pres-
ent, usually retracted into the anal tube, and provided with six, eight,
ten, or more anal ring setae. The anal lobes and anal setae are want-
ing. The caudal abdominal segment can not be identified. The body
is provided with cerores, but the number is frequently not large and is
often limited to the canellae and about the vulva. The body is never
155
156 THE_ COCCIDAE
provided with pilacerores, octacerores are rarely present. Ceratubae
are frequently present. The body is never enclosed in a resinous cell
with three adjacent openings or covered by a scale with or without
exuviae, is often naked or nearly so, sometimes enclosed in a compact
ovisac and sometimes excreting a mass of doughy wax from the ventral
cerores in which the eggs are laid. The rectum is never provided with
a long glassy tube of wax or with a chitinized rectal tube bearing rings
of anacerores. The body is frequently naked and sometimes gall-like in
form, but if so, always provided with normal antennae and legs and anal
ring and anal ring setae.
The nymphs of all stages are provided with a rostrum and rostralis.
The three pairs of legs are present in the first nymphal stage and usually
in all the others, particularly if they are present in the adult, and are
similar in form. The profemora are never greatly enlarged and the
prothoracic legs are never fitted for digging. The body is provided with
an anal cleft, rarely wanting, and with opercula in different stages of
migration from margin of body to cephalic end of anal cleft in nymphs
of different ages. The anal ring is always distinct and bears six to ten
or more anal ring setae. The anal lobes and anal setae, as such, are
present only in the first nymphal stage. Octacerores are rarely pres-
ent, pilacerores or ceratubae are never present. The caudal end of the
rectum may be strongly chitinized, but is never modified into a rectal
tube provided with one or more rings of anacerores excreting a long
glassy tube of wax. The caudal abdominal segment is never protuberant.
The male has ocellanae, the number varying with the genus. The
abdomen is not provided with long lateral filaments or tufts of long
wax filaments, but may bear two long thread-like caudal filaments of wax.
The stylus is short and conical.
The members of this subfamily are, as a rule, when denuded
of their wax, homogenous in appearance, not only as adults but as
nymphs. The most striking features in the adult female are the
mesal slit or cleft at the caudal end of the body which, except in a
few genera where the two sides of the slit have fused with an
obliteration of the line of fusion, is rarely absent. There is a
corresponding slit in the young nymphs, bounded on each side by
a distinct lobe or plate, an operculum, which bears a single large
anal seta and frequently one or more other smaller setae.
There is considerable variation among the numerous genera as
to the amount of wax excreted, which gives the body of the adult
female a very different look according as to whether the amount
of wax is large or small. The amount of wax formed by the nymph
except in certain genera is as a rule not large, so that the body of
the great majority of the species is generally stated to be naked.
The genera Lecanium, Saissetia, Physokermes and their allies are
such genera and are said to have the body naked. This is not
SUBFAMILY LBCANnNAB 157
absolutely true for the dorsal and more often the ventral aspect
is covered more or less with mealy wax. These species are,
however, truly naked when compared with representatives of such
genera as Ceroplastes which is completely encased in a mass of
wax or Pulvinaria which excretes a large white doughy mass of
wax several times as long as the body or as Takahashia where the
wax excreted is a tail several inches long. In Platinglisia
the waxy covering is glassy, in Signoretia woolly, and in Eriopeltis
a closely felted sac.
The body of the adult female in those genera where the body
is said to be naked, as Lecanium, is fairly uniform in appearance.
This form of body has been taken as the type for the following
description. The general outline is that of an elongate oval figure,
passing through the various stages to that of a rotund figure or one
that is almost or wholly globular. The two surfaces of the body are
near together, the dorsal convex and the ventral concave. This
gives the characteristic scoop-like figure of the shell of a tortoise
or of a steel war helmet and hence the vernacular name of the
group. Those species that are encased in a mass of wax, varying
in amount and thickness with the species and those species that
form large ovisacs depart more or less from this general form.
Those that form ovisacs have the body flat and both surfaces more
or less convex and approximate the condition of a typical eriocoecid
and so far as form is concerned represent the generalized end of
the series of genera. The person with an eye for judging form
will note, however, that even in these genera, there is a striking
resemblance between the wax excreting and the naked species. A
prominent feature of most of the naked species is the strongly con-
cave ventral aspect of the body of the older nymphal stages and of
the adult female.
A noticeable feature of the body is the entire absence of any
indication of segmentation in the adult female. The only mea;ns of
identifying the extent of the three primary regions is through the
presence and articulation of the antennae, rostrum, legs, and the
position of the thoracic spiracles.
The typical form of the body, shaped like a tortoise-shell,
makes the margin of the body sharp and prominent, forming a
distinct margin. This margin is generally fringed with setae
which vary considerable in number, size, and shape. They are
known as the marginal setae. While in many species these setae
158 THE COCCIDAE
are typical in form with a slender pointed flexible shaft, in others
the shaft is stiff and inflexible or spine-like. The distal portion of
the shaft is frequently distinctly enlarged or dilated forming a
clavate seta. Such setae may have the clavate distal portion of
the shaft indented or toothed forming frayed or furcate setae.
The marginal setae in certain genera have been modified into
flattened scales which are broad oval in outline or almost round,
broader than long. They differ from the lepidopterous scales in
their uniformly convex untoothed margin. They are designated
as flabellae by Green and as flabelliform marginal hairs by others.
There is on the dorsal aspect of certain species a varying
number of small tubercles, ten to twenty, which have been described
and figured by both Thro and Green. They were named sub-
marginal tubercles by the latter, who considered them as glandular
in nature. There are usually about five on each side of the body
near the lateral margin. Thro describes in one species a mesal row
of twenty. These minute structures rest upon the surface of the
cuticle and resemble in profile a small neckless flask with a central
dipper. Since they are likely to occur upon any part of the
dorsum, they have been named the dorsal tubercles.
The caudal extent of the head can not be determined. The
eyes are usually inconspicuous or wanting, when present they are
small, slightly convex areas which are sometimes darkened with
pigment and located on each lateral part of the ventral aspect of
the head.
The antennae usually consist of six to eight segments. This
number may be greatly reduced or the antennae modified into
mere tubercles, while in certain genera the antennae are always
wanting. In some genera the length of the segments is greatly
reduced and the antennae as a whole greatly shortened, while the
typical number of segments may be present. The comparative
length of the segments of the antennae, expressed in the form of
a formula, has been used by certain workers in differentiating
species. Such characters are no more reliable here than in the
other subfamilies. This fact and the general uniformity of the
number and arrangement of the antennal setae has been noted by
Thro, who found the number identical in the different species that
he studied and in cases noted where there was a difference and he
was able to find material for study, found that some of the setae
had been broken off and overlooked. The interantennal setae are
SUBFAMILY LECANIINAE 159
frequently present, but instead of being arranged to form the
comers of a four sided figure are frequently placed in a transverse
or approximately transverse line. _ _
The mouth-parts are normal in form and position. The rostrum
consists of a single segment. Its distal margin is rounded. There
is a prominent crumena connecting with the cavity extending
through the rostrum in which the loop of the rostralis is held.
The tendons attached to the proximal end of the mandibles and
the maxillae are prominent and the tendons attached to the in-
funda, the hypopharynx of Berlese and Green, are very large. The
ventral aspect of the endoskeleton of the head, the so called cradle
of Green, is bounded on the ventral side by a broad band, the
clypeus of Green. This latter structure, however, is always located
within the lumen of the head and since the clypeus is a sclerite of
the external skeleton, can not be considered as homologous with the
clypeus. This band has been called the clavus and the so called
cradle is derived from the tentorium, consisting in the main of the
arms of the metatentoria.
The legs, when present, are completely developed and all
parts are present, although they are frequently so small that their
tips project only slightly or not at all beyond the lateral margin
of the body. The tip of the tarsus frequently bears a pair of long
slender dorsal digitules and the claws a pair of short and stouter
ungual digitules.
The spiracles open on the ventral aspect, each mesothoracic
spiracle just laterad of the articulation of a prothoracic leg and
each metathoracic spiracle about in line with a mesothoracic
spiracle and midway between the mesothoracic and metathoracic
legs. The spiracles of this subfamily are frequently figured as
distinct trumpet-shaped or hourglass-shaped structures. All of
which except the opening, the spiracle, is a cavaera.
There is extending from each spiracle to the lateral margin
of the body in most species a more or less distinct furrow. Each
of these furrows is a canella, which bears a varying number of
cerores, the spiracerores. They are also known as the grouped
orifices of the thoracic spiracles, glandular pores or spinnerets,
spinnerets of the spiracles, glands of the spiracles, parastigmatic
glands, and parastigmatic pores. They were first named in the
Diaspidinae where the canellae and spinae are always wanting.
These cerores excrete a powdery wax which serves to keep the
caneUae open and permit the ingress of air to the spiracles. There
100 THE COCCIDAE
is frequently at the margin of the body a more or less distinct
indentation where the canellae terminate. These indentations are
the stigmatic clefts, also known as spiracular depressions or spirac-
ular grooves. The wax excreted by the spiracerores frequently
projects as a white powdery mass from the stigmatic clefts. The
lateral ends of the canellae are usually provided with large spine-
like setae which are located in the stigmatic clefts and serve to
keep the mouth of the canellae open. These setae, the spinae, also
known as stigmatic or spiracular spines or setae, vary greatly in
form and size from the marginal setae and are useful as characters
in identifying the species of this subfamily. The number of
spinae varies in the different species. There may be one, two, three,
four, five, or six in each stigmatic cleft. The great majority of
the species are provided with three spinae in each cleft. The
median one in such species is known as the median spina and the
others as lateral spinae. The median spina is usually much longer
than the lateral or all the others where there is more than three.
In some species the spinae are very short and broad, broader than
long, while in others they may be small pointed normal setae.
The abdomen is usually convex-concave like the thorax. There
is no indication of segmentation in the adult female. The marginal
setae and marginal scales, when these latter are present, extend
around the margin of the abdomen.
The most striking feature in the structure of the abdomen is
the deep mesal cleft or emargination, the anal cleft. The sides of
the cleft frequently overlap so that the cleft appears simply as
a longitudinal suture. In a few genera referred to this subfamily,
as Aclerda, the anal cleft is wanting. It is assumed that in such
cases the sides of the cleft have fused with a complete obsolescence
of the line of fusion. The length of the anal cleft varies somewhat
with the genus or species, it is generally about one-fourth to one-
fifth the length of the body, but in some cases, as Protopulvinaria,
it may be one-half or more of the total length of the body. In
those species deeply imbedded in wax as certain species of
Ceroplastes, the caudal portion of the body has been prolonged into
a prominent strongly chitinized horn or handle-like structure and
the anal cleft is located at the caudal end of the horn. This horn
is known as an opercularia. The length of the opercularia varies
not only with the species but with the depth of the wax excreted.
In those species where the amount of wax is considerable, the
opercularia is almost one-half the length of the body. It varies
SUBFAMILY LECANIINAE 161
also in position, in some species it is directed caudad and in others
dorsad.
A structural feature equally as striking as the anal cleft is
the presence in the adult female of a triangular plate on each side
of the cephalic end of the anal cleft. These plates are adjacent,
exposed on the dorsum, and each plate is known as an operculum.
They are also known as the anal plates, valve anali, while Green
considered the two plates as a single structure and designated the
two together as the operculum or anal operculum and each of its
parts as an anal plate or valve of the operculum. The shape of
the opercula varies with the species. The angle at the caudal end
of each operculum is known as the apex, the one at the cephalic
end as the inner angle, and the one on the lateral margin as the
outer angle. The margin connecting the inner angle and the apex
is known as the mesal or inner margin, the one connecting the apex
and the outer angle as the caudo-lateral or outer margin, and
the one connecting the inner and outer angles as the cephalo-lateral
margin or base. Each operculum is attached to the body along the
lateral margin. The outer angle is wanting in semicircular
opercula and the base and outer margin form a continuous curve.
The cuticle ventrad of the opercula in this subfamily is
invaginated and forms a barrel-shaped tubular structure or
eversible-sac continuous with the rectum. This is the '^organo
retrattile anali" of Berlese and the anal tube of Thro and other
writers. As to just how this tube originated is not clear. A
similar result would be obtained if the anus and anal ring were to
migrate away from the exterior of the body, pushing the rectum
into the body cavity and pulling the portion of the cuticle sur-
rounding the anal ring into the depression formed to bound the
tube extending from the anal ring to the exterior. The anus is
located inside the body. It is surrounded by a distinct anal ring
which bears a varying number of anal ring setae, six, eight, ten,
or more, which project through the anal tube and when longer than
the anal tube may be observed projecting from its caudal end.
The anal ring usually bears prominent orbacerores. There are
two projections on the ental surface of the anal tube adjacent to
the anal ring to which large muscles are attached. These muscles
extend and retract the anal tube. They function also for extrud-
ing the excrement or honeydew and for everting the anal ring.
The surface of the cuticle lining the lumen of the anal tube is
finely longitudinally striated. These striations are frequently
162 THE COCCIDAE
mistaken for setae and described as such. The anal tube is normally
retracted and must be everted in order to expose the anal ring and
the anal ring cerores or orbacerores. These cerores in some species
excrete a fine tube of wax which extends beyond the body and
serves to carry the honeydew still farther away from it. The
anal ring in some species is subdivided into two parts or nearly so
and frequently one-half is larger than the other. Berlese in his
''Le Cocciniglie Italiane vivente Sugli Agrumi" gives several
excellent figures of the anal tube and its associated parts. This
masterpiece of Berlese, so rarely noted in the literature of the
Coccidae that one might readily forget its existence, is the most
complete and comprehensive work that has ever been published
dealing with the morphology of coccids.
The caudal end of the anal tube on the ventral aspect may
bear two, four, six, or eight setae in a transverse or nearly trans-
verse row. The setae were named by Thro the fringe setae. The
seta at each lateral end of this row when there is more than two
fringe setae present, is usually larger than the mesal setae. When
there are only two fringe setae, each is located near a lateral
margin of the anal tube; when there is four, they are usually
equidistant from each other, two on each side of the meson or
adjacent near each lateral margin. While the fringe setae are
always arranged in a transverse row, it is not always a straight
line but usually more or less curved.
The ventral aspect of the outer wall of the anal tube in certain
species bears two longitudinal rows of setae, the hypopygial setae.
These rows usually converge cephalad or are parallel and appear
to be a continuation of the fringe setae.
The position of the anus and anal ring at the cephalic end of
the anal tube and forming the cephalic boundary of its lumen has
already been indicated. The position of the vulva is difficult to
identify. It is located on the ventral aspect cephalad of the
external opening of the anal tube and the fringe setae.
The lateral portion of the ventral aspect of each operculum
frequently contains a distinct longitudinal thickening, a ventral
thickening. It is also known as the ventral chitinous process of
the ventral plate. The ventral thickenings vary considerable in
length and breadth. The row of fringe setae appears to be limited
on each side by a ventral thickening.
The distal half of each operculum bears several small setae.
There is one group which appears to form a line continuous with the
SUBFAMILY LECANIINAE 163
caudal end of the ventral thickening. These setae are the sub-
apical setae. While there is frequently two subapical setae, there
may be as many as five or more. The small setae, generally if not
always four in number, located caudad of the subapical setae are
the apical setae. They may be located on the dorsal aspect or on
the ventral aspect or a part of them may be located on the dorsal
and the remainder on the ventral aspect. At times some may be
located on the margin of the operculum. The caudal half of the
dorsal surface in certain species bears a single large seta, the
discal seta. This seta is of about the same size as the subapical
setae and is located on the same portion of the dorsal aspect of the
operculum, so that in mounted transparent specimens, they appear
to be closely associated.
The ventral aspect of the body cephalad of the opercula bears
a series of three or more pairs of small setae, the abdominal setae
of Green. These setae are usually arranged in two longitudinal
rows. The ventral aspect of the abdomen cephalad and laterad of
the opercula also bears a few scattered cerores, the circumgenital
glands of Green.
The cuticle of the dorsal aspect of certain lecaniids, as the
species of the typical genus and its allies, when studied under
transmitted light, show a more or less distinct pattern of variously
shaped areas. These areas are usually round or oval or suboval
and appear as large cellular areas. They frequently consist of a
series of concentric lighter and darker colored bands or areas and
are known as the cells or as the dermal cells or pores or cellulae.
They are not cells in a histological sense but are probably thicken-
ings of the cuticle of different density. They vary greatly in
number, shape, and size in the different species. The cellulae are
more distinct in old individuals, they also become indistinct after
a long boiling in caustic potash. Those species, where the cellulae
form a series of disconnected rings or ovals, are said to have the
cuticle alveolate. When these areas are connected forming a
series of adjacent polygonal areas, these areas are designated as
tesserae.
The opening in each cellula is, so far is observed, always the
mouth of a ceratuba or an oraceratuba. The oraceratubae are
usually, if not always, eccentric in position. The ceratubae appear
to be characteristic in form but vary greatly in length. Each
generally consists of a proximal straight tube bearing a smaller
curved portion or tube. The change in diameter of the two tubes
164 THE coccroAE
is due to a sudden constriction which is limited to one side. The
distal end of the small tube bears a bouquet-like swelling.
The adult females of the Lecaniinae usually produce eggs,
which are in most cases deposited under the concave ventral aspect
of the body or in an ovisac. Many species instead of depositing
their eggs give birth to living young. In such cases the ruptured
egg-shell and the young nymph may be extruded from the vulva at
the same tim(<. Such ispecies differ from the true oviparous
species only in that the eggs are retained in the body of the female,
in the oviduct, until they hatch. Certain writers have made the
mistake of considering all species whose females give birth to living
young as reproducing parthenogenetically. While undoubtedly
many females are able to produce young without being fertilized,
it is quite likely a much larger number give birth to living young
from eggs that have been fertilized but are retained by the female
until after the eggs hatch. It is not impossible that many species
reproduce in both ways, that is, that certain females at certain
seasons reproduce pathenogenetically while other individuals re-
produce in the normal way. It is difficult to explain why there
should be an absence of males at certain times. The giving birth
to living young is not a proof of parthenogenesis.
The eggs are ovoid in shape and white, yellow, or red in color.
They are of common occurrence in those species which are naked
and where the ventral aspect is concave. In such cases the body
of the female serves as a plate for covering and protecting the
eggs. The edge of the body adheres to the host-plant and, when
the body is raised, the space under it is seen to be packed with
eggs. The cerores located on the ventral aspect of the body ex-
crete a powdery wax, the amount varying with the species, which
is mixed with the eggs.
The females in certain genera, as Pulvinaria, are naked until
they become adult and are ready to deposit their eggs. They then
begin to excrete a mass of doughy wax from the numerous cerores
located on the ventral aspect of the body. This wax is gradually
piled up as a pad or cushion until the caudal end of the body is
gradually raised from the host-plant and the body assumes a ver-
tical position. The insect is attached to the host-plant only by
the mouth-parts and the ovisac, which is often several times the
length of the body, and projects behind it. As a result the body
of the adult female after all the eggs are laid is often washed off
by the rain while the wax containing the eggs may remain for a
SUBFAMILY LECANIINAE 165
long time in contact with the bark of the plant. The adult female
in other genera forms a loosely or closely felted cylindrical sac of
wax in which the eggs are deposited. In Ceroplastes the body is
completely encased in wax, formed by the fusion of plates of vari-
ous sizes and shapes. The ventral aspect of the body contracts
away from the wax surrounding the body and forms a cavity in
which the eggs are deposited. A similar condition is found in
other genera where the body is completely encased in wax. The
formation of a definite ovisac is usually limited to the adult
female but in Eriochiton according to Maskell the covering of wax
is formed during the last nymphal stage. In Ceroplastes, if the
wax of the adult is considered as an ovisac, the formation of the
primary plates from which the wax of the adult is derived begins
in the first nymphal stage. The number of plates increases in
each stage and in many species the shape and symmetry of the
plates is lost by their fusion into a thick shapeless mass. But as
Green points out, the plates of wax formed by the first and later
nymphal stages can often be identified in the mass of wax encasing
the adult female. The number of molts undergone by the indi-
viduals of this genus and the relation of the exuviae to the wax
is not understood.
Where the body is deeply encased in wax, as in the genus
Ceroplastes, the volume of wax may be greater than the volume of
the body. While in this genus the wax and body together fre-
quently form a globular mass, this is not always true, for in some
species of Ceroplastes and in the species of such genera as Vinsonia,
Ctenochiton, and Inglesia the form of the wax is definite and char-
acteristic. The wax is generally designated as a test where the
body is encased in a mass that is hard, horny, or glassy. The term
test is used, however, by some writers to include the covering of
wax no matter what its form. The formation of a test is not
peculiar to the Lecaniinae, but it is found in genera of other
subfamilies.
The sexes of the lecaniids are not distinguishable in the first
nymphal stage. The individuals of this stage are oval in outline,
usually somewhat depressed with well developed legs and antennae.
The latter are usually provided with six segments. The young
nymphs are very active, searching out all parts of the plant and
with their ungainly appearing legs and antennae resemble mites
or acarids. The caudal end of the body is usually provided with
a distinct anal cleft. The most striking feature which seems to be
166 THE COCCIDAE
characteristic of practically all the first stage nymphs of species
of this subfamily, is the presence of a distinct operculum on each
side of the anal cleft. These instead of being at the cephalic end
of the anal cleft are at the caudal end and form a part of the
caudal margin. Each operculum bears a single large seta, an anal
seta, and often other much smaller setae. The opercula are con-
sidered as the homologues of the anal lobes and by some as the
homologues of the last abdominal segment, but more strictly should
be homologized with the penultimate segment. The thoracic
spiracles are always present, but there is some variation in the
presence and size of the canellae, stigmatic clefts, spinae, and
spiracerores. The anal ring and the anal ring setae are present
and situated at the cephalic end of the anal tube which opens at
the cephalic end of the anal cleft. The anal ring and anal tube
and their associated parts are frequently extruded as a bag-like
structure after treatment with caustic potash. The possession of
an anal ring and anal ring setae and especially of anal lobes and
anal setae in the first nymphal stage shows that there is a
phylogenetic relationship existing between the lecaniids and
eriococcids.
The female nymphs of the second stage have legs and antennae
that are smaller than those of the first stage. They resemble more
closely the adult females in general appearance. This is due to
the size in the naked species and to the increase in the amount of
wax in those genera excreting a covering of wax. The opercula,
while they are not yet located at the cephalic end of the anal cleft,
have migrated away from the caudal margin and at the first molt
lost their anal setae. A second molting of the cuticle transforms
them into adult females. It is not difficult to understand how the
naked species are able to molt, but in the species of such genera
as Ceroplastes which bear a test of wax of considerable thickness
which adheres closely to the body throughout life, it is not easy to
understand how this takes place. It has been suggested by Green,
since in this genus the plates of wax excreted by the dorsum and
latus of the larva can be recognized in the test of the adult, the
molted skin must be incorporated in the mass of wax of the adult.
The males are said to molt three times and to pass through
three nymphal stages before becoming winged adults. They are
difficult to separate from the females during the second nymphal
stage until they have formed their cocoons, which in the great
majority of the males of this subfamily is a subdepressed, thin,
SUBFAMILY LECANIINAE 167
glassy, test of wax. This test is formed toward the close of the
second nymphal stage. It is divided into a series of symmetrically
arranged areas by a series of raised or thickened lines. The num-
ber of these areas varies from six to nineteen. A common type of
the naked lecaniids is to have two elongate mesal or dorsal areas, a
cephalic marginal area, and about three lateral marginal areas on
each side, or nine areas in all. The males of many genera are
unknown. In the genus Cryptes the cocoon is felted, while in
Ericerus, a Chinese species, the males are gregarious and envelope
themselves in a mass of wax, which is of such amount that it was
collected and used for the making of candles before the introduc-
tion of paraffin candles. The third nymphal stage of the male is
passed under the cocoon. The developing legs and antennae of
the adult are prominent. The second and third exuviae are shed
while under the cocoon. They are pushed out, when cast, from
beneath the posterior extremity.
The adult males are similar to those described in previous
subfamilies so far as their general form is concerned. There is
considerable variation among the various species as to the number
of ocellanae present. Newstead figures the facets of twelve simple
eyes in Lichtensia, eight in Vinsonia, and six in Pulvinaria. The
caudal pair of the dorsal aspect are probably ocelli and the others
are ocellanae. The caudal end of the body may bear a pair of
conspicuous thread-like filaments of wax which are wanting in
some genera. The stylus is generally a long straight tube.
GENERA OF LECANIINAE
a. Body of adult female not naked, always forming or attached to or
encased in distinct mass of wax of varying form and consistency or
body covered by thin film of opaque or glassy wax; abdomen of adult
female sometimes with segmentation clearly indicated,
b. Wax excreted by adult female doughy or cottony, situated be-
neath or behind or entirely enclosing body, body never covered
with thin film of wax; abdomen of adult female sometimes with
segmentation clearly indicated.
c. Body of adult female more or less enclosed, frequently entirely
enclosed in cottony or felted ovisac, wax rarely limited in great
part to dorsal aspect; abdomen of adult female sometimes with
segmentation clearly indicated.
d. Antennae of adult female consisting of eight segments.
e. Mesothoracic and metathoracic legs much longer than
prothoracic, tarsi of latter secondarily divided into two
segments; antennae of first stage nymphs consisting of
168 THE COCCIDAE
Six segments; anal ring of adult female with eight and
of first nymphal stage of six anal ring setae
Exaeretopus Newst.
ee. Mesothoracic and metathoracic legs subequal in length
with prothoracic legs; tarsi of all legs consisting of single
segment.
f. Body of adult female greatly elongated, much longer
than broad.
g. Body of adult female elongate with parallel sides;
legs normal in form, slender and well developed;
anal ring with six anal ring setate
Luzulaspis Ckll.
gg. Body of adult female twice as long as wide; legs nor-
mal; antennae consisting of seven or eight segments;
antennae of first nymphal stage consisting of six seg-
ments. Spermococcus Giard.
ff. Body of adult female oval or suboval in outline, not
greatly elongated.
g. Body of adult female margined with distinctly thick-
ened marginal setae.
h. Spinae pointed, stout, subequal in length or small-
er than marginal setae; body of adult female ex-
cept a small cephalic portion, completely enclosed
in felted white ovisac; anal ring of adult female
with eight anal ring setae Lichtensia Sign.
hh. Spinae much larger than marginal setae; body
of adult female completely enclosed in compact
striated ovisac; anal ring of adult female with
eight anal ring setae Stotzia Marchal.
gg. Body of adult female margined with short, fine
pointed setae; body of adult female covered by ovisac
of curled threads of wax divided into four trapezoi-
dal areas, which in egg laying female completely
covers dorsal aspect; anal ring of first nymphal stage
with eight anal ring setae Euphillipia Berl. & Silv.
dd. Antennae of adult female with less than eight segments.
6. Antennae of adult female consisting of seven segments.
f. Body of adult female not enclosed in spherical felted
ovisac with elongated dorsal opening.
g. Body of adult female oval or elliptical, with loosely
felted excretion of wax, especially in second nymphal
stage, but absent or inconspicuous in adult female;
tarsi longer than tibiae; body with marginal fringe.
Eriochitin Mask.
gg. Body of female elliptical and convex, with closely
felted ovisac; cuticle with many large circular
cerores; anal ring with six anal ring setae; margin
with prominent marginal setae Mallococcus Mask.
SUBFAMILY LECANIINAE 169
ff. Body of adult female enclosed in spherical felted ovisac
with elongated dorsal opening; anal ring of adult fe-
male with six anal ring setae and their tibiae longer
than their tarsi AustroUchtensia Ckll.
ee. Antennae of adult female with less than seven segments.
f. Antennae of adult female consisting of six segments.
g. Wax excreted by adult female not limited for most
part to dorsal aspect; stigmatic clefts without spinae.
h. Body of adult female elongate, two or more times
as long as broad, antennae and legs short and
stout; ovisac cottony or felted; dorsum with
numerous truncated setae Eriopeltis Sign.
hh. Body of adult female oval or suboval, not twice
as long as broad.
i. Adult female with body enclosed in felted ovi-
sac; legs and antennae normal in size; insects
arboreal in habits; anal ring of first stage nymphs
with eight anal ring setae Fillipia Targ.
ii. Adult female with body enveloped in loose
white cottony wax or in thin shell of hard wax;
legs and antennae complete in form but greatly
reduced in size; insects subterranean in habits;
antennae of adult female sometimes consisting
of eight segments Lecanopsis Targ.
gg. Wax excreted by adult female limited for most part
to dorsal aspect, excreted from prominent lateral or
mesal rows of cerores with cuticular tubes; stigmatic
clefts with four spinae Geronema Mask.
ff. Antennae of adult female rudimentary, segmentation
indistinct, four segments or less; legs rudimentary,
stout conical claw-like protuberances
PseudophilUpia Ckll.
cc. Body of adult female excreting doughy or cottony ovisac situ-
ated beneath or behind body of insect; abdomen of adult female
never with segmentation indicated,
d. Adult female not excreting an ovisac that is elongated pos-
teriorly; dorsum of insect sometimes covered with wax,
e. Antennae consisting of eight segments; ovisac cone-
shaped and fluted; body of insect free from host-plant on
end of conical ovisac; anal ring with ten anal ring setae. __
Pulvinella Hemp,
ee. Antennae consisting of seven segments; ovisac not coni-
cal with fluted sides; dorsum of second nymphal stage
with lamellae of wax Pseudopulvinana Atk.
dd. Adult female excreting ovisac that is elongated posteriorly
behind body of the insect and that does not cover body at all.
e. Body of adult female triangular in outline; ovisac only
slightly developed, mere fringe around hind margin of
170 THE COCCIDAE
body; anal ring with six anal ring setae
Protopulvinaria Ckll,
ee. Body of adult female oval or suboval in outline; ovisac
usually well developed.
f. Ovisac at most never more than four or five times as
long as broad, attached to host-plant throughout its en-
tire length.
g. Body of adult female soft, pink in color in front,
greenish on dorsum with minute black specks, cephal-
ic part of dorsum with white area of wax, body with
irregular patches in three longitudinal rows; anten-
nae consisting of eight segments.- -Philephedra Ckll.
gg. Body of adult female hard and strongly chitinized,
dorsum usually naked, if provided with wax, not
arranged in longitudinal rows.
h. Body with dorsum entirely covered with felt-like
white secretion of wax in two parts, lateral por-
tion and mesal portion; antennae of adult female
with eight segments; anal ring with six anal ring
setae. Tectopulvinaria Hemp.
hh. Body with dorsum naked, at most only covered
with powdery wax; antennae of adult female
normally with eight segments, sometimes with
seven; anal ring with eight anal ring setae
Pulvinaria Targ.
ff. Ovisac greatly elongated, always much longer than
broad, attached to host-plant only at one end; antennae
of adult female consisting of seven segments; anal ring
apparently with six anal ring setae Takahashia Ckll.
bb. Wax excreted by adult female horny or opaque or glassy, thick
or thin mass or sheet of wax closely attached to or free from
body of insect, never doughy or cottony mass; abdomen of adult
female never with segmentation indicated.
c. Adult female with legs present and normal in form and size,
d. Adult female with antennae present and normal in form.
e. Test of adult female consisting of opaque waxy wax, more
or less thick, never thin and brittle.
f. Test of adult female not with marginal fringe or with
radiating processes of wax.
g. Abdomen of adult female usually with a more or less
distinct opercularia; antennae of adult female con-
sisting of six to eight segments; margin of body of
adult female with marginal setae, never with spear-
head-shaped spines Ceroplastes Gray.
gg. Abdomen of adult female never with opercularia;
antennae of adult female obscurely segmented; mar-
gin of body with few marginal setae and numerous
spearhead-shaped spines Cardiococcus Ckll.
SUBFAMILY LECANIINAE 171
ff. Test of adult female always with a marginal fringe or
with radiating processes of hard wax.
g. Test of adult female with seven prominent radiating
processes of wax, insect star-like in appearance;
opercularia present but small; antennae of adult fe-
male consisting of six segments Vinsonia Sign.
gg. Test of adult female not with seven radiating pro-
cesses but with marginal fringe of prominent tooth-
like projections of wax; opercularia never present;
antennae of adult female consisting of seven seg-
ments. Ctenochiton Mask.
ee. Test of adult female glassy, thick or thin and brittle,
never opaque and thick.
f. Test of adult female perfectly flat or only slightly con-
vex; antennae of adult female consisting of eight seg-
ments.
g. Test with mesal part of dorsal portion subdivided
into small oval plates; body of adult female about
as broad as long. Lagosinia Ckll.
gg. Test apparently subdivided into four plates, one on
the dorsum, one on each side, and one at the caudal
end; body of adult female nearly twice as long as
broad and anal ring with ten anal ring setae
Alichtensia Ckll.
ff. Test of adult female not flat, usually irregular in form
and strongly convex, approaching spherical.
g. Test of adult female conical with its surface fluted;
antennae of adult female consisting of five segments;
marginal setae numerous, short, sharp, and conical.
Edwallia Hemp,
gg. Test of adult female not conical and if approaching
conical form, not longitudinally fluted,
h. Test of adult female with distinct air-cells.
i. Insect living free, never forming galls on
plants; test of adult female always divided into
plates like the shell of tortoise and striated
with rows of air-cells, not free from body of
insect; antennae of adult female consisting of
six segments. Inglisia Mask.
ii. Insects not living free, forming galls upon
roots of plants; test of adult female not divided
into plates and striated, air-cells running to-
gether, forming long air-spaces, not free from
body of insect; antennae of adult female con-
sisting of seven or eight segments
Cryptinglisia Ckll.
hh. Test of adult female never with distinct air-cells,
i. Test of adult female divided by transverse de-
172 THE COCCIDAE
■ pression, portion on each side of depression ele-
vated and subcorneal with grooves radiating
from their apices, probably not free from body
of insect; antennae of adult female consisting
of eight segments. Parafairmairia Ckll.
ii. Test of adult female convex, subglobular, free
from body of insect, surface roughened by
irregular granules or symmetrical polygonal
plates of wax; antennae of adult female consist-
r". i^S of eight segments GeropVastodes Ckll.
dd. Antennae wanting or indicated only by two impressions;
insect enclosed in test that is deeply incised at stigmatic and
anal clefts; opercula twice as long as wide; insects living in
nests of ants Myxolecanium Beccari.
cc. Adult female with legs rudimentary or wanting.
j d. Adult female with antennae present, consisting of one to
seven segments.
e. Adult female covered by hard brown test, mesal portion
formed during nymphal life; antennae consisting of seven
segments; cuticle of dorsal aspect not with large alveolae.
Lecanochiton Mask.
ee. Adult female covered by thin glassy test, all formed by
adult; antennae short with indistinguishable segments;
cuticle of dorsal aspect with large alveolae
Neolecanium Ckll. & Parr.
dd. Adult female with antennae wanting.
e. Test of adult female flat and circular, divided into two
parts on dorso-meson, with rows of air-cells
Platinglisia Ckll.
ee. Test of adult female always more or less convex and never
provided with air-cells.
f. Test of adult female convex, thin glassy scale, divided
into two parts by longitudinal furrow, forming sub-
lateral low cones with concentric but not longitudinal
striae; anal ring with seven anal ring setae
Pseudokermes Ckll.
ff. Test of adult female convex, semitransparent thin
glassy scale, divided into two parts by longitudinal
furrow, with low eminence on each side of furrow, pro-
vided with radiating and concentric striae; anal ring
with six anal ring setae Schizochlamidea Ckll.
aa. Body of adult female always naked, never attached to or encased in
distinct mass of wax of varying form and consistency or with body
covered by thin sheet of opaque or glassy wax, at mosit only with
powdery bloom or thin covering or patches of pulverulent wax on
dorsal or ventral aspects; abdomen of adult female never with
segmentation indicated.
SUBFAMILY LECANHNAE 173
. Adult female with legs and antennae always present and antennae
segmented.
c. Abdomen of adult female conspicuously transversely furrowed,
appearing as if distinctly segmented, as seen in unmounted
specimens from dorsal aspect; antennae of adult female short
and thick, with six segments; legs rudimentary, segments short
and swollen. Alecanopsis Ckll.
cc. Abdomen of adult female never with prominent transverse fur-
rows so as to appear as if segmented.
d. Adult female with margin of the body fringed with marginal
setae, never with scales.
e. Body of adult female always alveolate, never with tesserae.
f. Abdomen of adult female always with distinct anal
cleft with opercula near margin of the body.
g. Cuticle of dorsal aspect with minute microscopic pits
or alveolae, distance between alveolae always several
times diameter of alveolae.
h. Adult female with alveolae never symmetrically
arranged, bounding tesserae-like areas.
i. Body of adult female flat or slightly convex;
antennae consisting of seven or eight segments;
median spina about twice as long as lateral
spinae; opercula usually longer than broad;
anal ring with eight anal ring setae
Lecanium Burm.
ii. Body of adult female distinctly convex, oper-
cula usually broader than long,
j. Adult female with antennae consisting of
seven segments; median spina usually less
than twice as long as lateral spinae; anal
ring with eight anal ring setae
Eulecanium Ckll.
jj. Adult female with antennae consisting of
seven or six segments; median spina usually
three times as long as lateral spinae
Mesolecanium Ckll.
hh. Adult female with alveolae symmetrically ar-
ranged and bounding tesserae-like areas; antennae
consisting of seven or eight segments; anal ring
with six anal ring setae Stictolecanium Ckll.
gg. Cuticle of dorsal aspect with large often nearly
adjacent alveolae, the distance between alveolae not
several times diameter of alveolae,
h. Body of adult female slightly convex or hemis-
pherical, lateral portions never vertical; antennae
of adult female consisting of eight segments; anal
ring with eight anal ring setae Saissetia Depl.
174 THE COCCIDAE
hh. Body of adult female with lateral portions per-
pendicular, edge of dorsal portion overlapping at
top like roof of house; antennae of second
nymphal stage of female with seven segments
Platysaissetia Ckll.
ff. Abdomen of adult female with anal cleft and opercula
located near middle of dorsal aspect.
g. Body of adult female spherical with opercula located
near middle of length of body; antennae minute,
consisting of eight segments in adult female and of
six in first nymphal stage, anal ring of first nymphal
stage with seven anal ring setae; dorsum of adult
female and of nymphs not with two cephalic and two
caudal groups of cerores Cryptes Mask.
gg. Body of adult female not spherical, flat and cor-
rugated, with opercula located near middle of length
of body; antennae of adult female consisting of nine
segments and of first nymphal stage of six; dorsum
of female in all stages with two prominent cephalic
and two caudal groups of cerores
Hemilecanium Newst,
ee. Body of adult female flat and convex, never spherical,
never with alveolae, but always with tesserae; antennae
of adult female with eight segments; anal ring with six
anal ring setae Eucalymnatus Ckll.
dd. Adult female with margin of body fringed with flattened
setae or scales; antennae consisting of eight segments; anal
ring with six anal ring setae Paralecanium Ckll.
bb. Adult female with legs and antennae, when present, all or in
part rudimentary, one or both wanting.
c. Adult female with body not greatly elongated, lateral margins
strongly convex, never subparallel; anal cleft and opercula dis-
tinct,
d. Body of adult female convex, sometimes approaching hemis-
pherical, never spherical.
e. Body of adult female convex, sometimes approaching
hemispherical, fringed with long slender, spear-like mar-
ginal setae; legs and antennae wanting.. _^ncerws Westw.
ee. Body of adult female always convex, not approaching hem-
ispherical, not fringed with slender spear-like marginal
setae.
f. Body of adult female with margin of dorsal aspect
divided into twenty plates of which one is cephalic, one
is about the anus, four are spiracular, and others are
intermediate in position; legs are wanting, antennae
tuberculate; anal ring with ten anal ring seate
Houardia Marchal.
ff. Body of adult female with dorsal aspect never divided
into plates; legs and antennae rudimentary, latter con-
SUBFAMILY LECANHNAE 175
sisting of six segments; cuticle of dorsal aspect with
minute distant alveola; anal ring with ten anal ring
setae. Toumeyclla Ckll.
dd. Body of adult female spherical, never convex or subdepressed.
e. Adult female with opercula not prominent and not dorsal
in position.
f. Cuticle of the dorsal aspect with minute distant
alveolae; legs and antennae rudimentary or wanting;
opercula obscure. Akermes Ckll.
ff. Cuticle of dorsal aspect with large polygonal sub-
adjacent alveolae; legs and antennae rudimentary
Megasaissetia Ckll.
ee. Adult female with opercula prominent, swollen, dorsal in
position; antennae and legs wanting; body with lateral
portions infolded forming marsupium on each side
Physokermes Targ.
cc. Adult female with body greatly elongated with lateral mar-
gins subparallel; anal cleft and opercula wanting; legs want-
ing; antennae represented by unsegmented tubercle on each
side; mesothoracic spiracles sometimes located considerable
distance cephalad of attachment of rostrum Aclerda Sign.
Exaeretopus Newst. — The type species, formiceticola Newst., was
collected in the nests of ants in England, the other species, caricis Ehrh.,
occurs on Carex and Trisetum in California.
Luzulaspis Ckll. — A single species, luzulae Dufour, is found in
Europe on Luzula. A second species, spinulosa Leon., has been recorded
from Argentina on Atriplex.
Spermococcus Giard. — This poorly defined genus includes a single
species, fallax Giard, from France on the roots of grass.
Lichtensia Sign. — There are twelve species included in this genus.
The following are found in America: — argentata Hemp, from Brazil;
colimensis Ckll. from Mexico on ?Celtis ; crescentiae Ckll. from Mexico on
"Guanabano"; lutea Ckll. from Mexico on Croton, wild fig; lycii from
New Mexico on "Lycium bush"; mimosae Towns. & Ckll. from Mexico
on Mimosa; parvula Ckll. from Mexico on Mimosa; simillima Ckll. from
Paraguay; viburni Sign, from France, England, Wales, Massachusetts on
Viburnum, Spiraea, Prinos, Hedera; zapotlana Ckll. from Mexico on copal.
Stotzia Marchal. — This genus includes a single species, striata
Marchal, from Algeria on Ephedra.
Euphillipia Berl. & Silv. — A single species, olivina Berl. & Silv.,
occurs on olive in Italy.
Eriochitin Mask. — A genus containing three species from New
Zealand and India, hispidus Mask, from New Zealand on Olearia;
spinosus Mask, from New Zealand on Atherosperma, Melicope, Elaeo-
carpus, Muhlenbeckia, and theae Green from India on tea-plant.
Mallococcus Mask. — Two species are described, sinensis Mask,
from China on Callicarpa and lanigerus Hemp, from Brazil.
Austrolichtensia Ckll. — A single species, hakearum Fuller, oc-
curs in Australia on Hakea.
176 THE COCCIDAE
Eriopeltis Sign. — Four species are included, bracliypodii Giard
on Brachypodium, festucae Fonsc. on Festuca, and lichtensteinii Sign,
on Festuca and other grasses from Europe. Festucae Fonsc. has been
Introduced into Nova Scotia, Eastern Canada, Maine, Illinois, Indiana,
and Dakota. The fourth species, coloradensis Ckll. occurs in Colorado
on grass.
Fillipia Targ. — A single species, oleae Costa, occurs on olives in
Italy.
Lecanopsis Targ. — Four species are found in Europe upon the
roots of grass, Asperula, and from the nests of Lasius. A single species,
lineolatae King & Ckll. occurs in Massachusetts in the nests of ants.
Ceronema Mask. — A genus of five species, three from Australia,
one from India and Japan, and one from Ceylon.
Pseudophillipia Ckll. — There is a single species, quaintancii Ckll.,
from Florida on pine.
Pulvinella Hemp. — A single species, pulchella Hemp., is de-
scribed from Brazil on Baccharis.
Pseudopulvinaria Atk. — There is a single species from India,
sikkimensis Atk. on Cinchona.
Protopulvinaria Ckll. — This genus includes two species, convexa
Hemp, from Brazil on Smilax, is the type, and longivalvata Green from
Ceylon.
Philephedra Ckll. — A single species, ephedrae Ckll. from New
Mexico on Ephedra, is included. According to Ferris this species ap-
pears like a much elongated Pulvinaria in which spinae and stigmatic
clefts are wanting. The marginal setae are large and spine-like.
Tectopulvlnaria Hemp. — A monotypic genus from Brazil with
albata Hemp, as type occurs on Veronia and Trichogonia.
Pulvinaria Targ. — This genus contains over sixty species of
which the following are found in America: — acericola Walsh & Riley
from eastern and western United States on maple; amygdali Ckll. from
New Mexico and Georgia on peach, apple, plum; argentina Leon, from
Argentina on Lycium, Fabiana; bigeloviae Ckll. from California, Colo-
rado on Bigelovia; broadwayi Ckll. from Grenada; camelicola Sign, from
Europe, Australia, Japan, southern United States on Camellia, Euonymus,
Oncidium, Calanthe; cockerelli King from Massachusette on Spiraea,
Prinos; convexa Hemp, from Brazil on Smilax; cooleyi King from Mon-
tana on Picea; coulteri Ckll. from Colorado on Rosa; cupaniae Ckll. from
Jamaica on Cupania, Bignonia, guava; dendrophthorae Ckll. from Ja-
maica on Dendrophthora; depressa Hemp, from Brazil on Miconia;
ehrhorni King from California on alder, willow; eugeniae Hemp, from
Brazil on Eugenia; ficus Hemp, from Brazil on Ficus, Psidium, Mangi-
f era, Ixora ; floccif era Westw. from England, India, Canada, Massachu-
setts, Trinidad on Auguloa, Lyasta, Camellia, Acalypha, Brassia, Phaius;
grandis Hemp, from Brazil on Myrtaceae; hunteri King from Kansas on
maple, honey locust; innumerabilis Rathvon from Canada, United States
on maple, many shade itrees; maclurae Fitch from eastern United States,
New Mexico on osage orange, sumac; mammeae Mask, from Hawaiian
Islands, North America on Mammea, ferns, orange, coffee, plum; occi-
dentalis Ckll. from Washington, British Columbia, Nova Scotia on cur-
rant, gooseberry, hawthorn, plum, pear, mountain ash, willow, poplar.
SUBFAMILY LECANIINAE 177
alder; ornata Hemp, from Brazil on Ariabidaea; plucheae Ehrh. from Cal-
ifornia on Pluchea; pruni Hunter from Kansas on plum, white elm;
psidii Mask, from New Zealand, Japan, California, etc. on tea, plum,
coffee, Citrus, etc.; pulchella Hemp, from Brazil on Baccharis; piri-
formis Ckll. from Trinidad, Jamaica, Grenada, Madeira on guava, cinna-
mon, etc.; rhois Ehrh. from California on Rhus; simulans Ckll. from
Trinidad, Mexico; spinulosa Leon, from Argentina on Atriplex; tiliae
King & Ckll. from Massachusetts on Tilia; tinsleyi King from New Mex-
ico on Celtis; tuberculatus Bouche from South America on Malvaceae;
urbicola Ckll. from Jamaica, Barbadoes, Trinidad on Capsicum; viburni
King from Canada on Viburnum; and vitis Linn, from Europe, United
States on Grape. The two common eastern species of the United States
can be separated as follows: —
SPECIES OF PULVINARIA
a. Cuticle of dorsum with distinct large subadjacent alveolae; ovisac
much longer than body of adult female and formed upon leaves of
host-plant. Eastern United States on Maple acericola W. & R.
aa. Cuticle of dorsum with small distant alveolae; ovisac not much
longer than body of adult female and formed upon stems and
branches of host-plant. — Eastern United States on maple, grape, elm,
box elder, etc. vitis Linn.
Takashasia CklL — The type species, japonica Ckll. is from
Japan on Mulberry A second species, jaliscensis T. & W. Ckll. occurs
in Mexico on Rhus(?). A third species, citricola Knw. has been de-
scribed from Japan on Citrus.
Ceroplastes Gray. — This genus occurs in all parts of the world
and includes over sixty species, of which over fifty are found in Amer-
ica. The following species are found in the United States: — cirripedi-
formis Comst. from Florida, Louisiana, Mexico, West Indies on orange,
quince, myrtle, persimmon, Solanum, Eupatorium, etc.; cistudiformis
Towns. & Ckll. from Mexico, California on Chrysis, Bignonia, Chrysan-
themum, Cordia, etc.; floridensis Comst. from southern United States,
West Indies, Ceylon, Asia, Australia on Oleander, Citrus, quince, red
bay, Anona, Andromeda, etc.; irregularis Ckll. from Mexico, New Mexico,
Texas on Atriplex. Cockerell has proposed the following subgeneric
groups for the genus Ceroplastes: —
1. "Typical Ceroplastes, with convex scales, in which the lateral
plates are large and reach the lower margin; caudal horn (opercularia)
usually moderate and directed posteriorly."
2. Ceroplastina Ckll. — Type, C. lahillei Ckll. "Wax abundant, pushing
the plates to the dorsal surface; caudal horn mammiform, directed up-
wards. Wax of different individuals often confluent."
3. Ceroplastidia Ckll. Type C. bruneri Ckll. "In this group the
wax of the several individuals is thick and always confluent, and the fe-
male has become high and narrow."
Cardiococcus Ckll. — Three species are included, two from Aus-
tralia and the third, umbonatus Ckll., is from Mexico on wild guava.
Vinsonia Si|^. — A single species, stellifera Westw., occurs com-
monly on many plants throughout tropical America and Ceylon.
178 THE COCCIDAE
Ctenochiton Mask. — There are fifteen species listed in Fernald's
Catalogue. The following occur in America: — aztectus Towns. & Ckll.
from Mexico on "Cafetilla Cimarron" and flavus Maskell from Brazil oa
Brachyglottis, Panax, Leptospermum, Elaeocarpus.
Lagiosinia Ckll. — A single species, strachani Ckll. from west
Africa on Anona, is known.
Alichtensia Ckll. — There is a single species, attenuata Hemp.,
occurs in Brazil on Baccharis.
Edwallia Hemp. — A monotypic genus described from Brazil on
Eugenia includes rugosa Hemp.
Inglisia Mask. — This genus includes eleven species. The follow-
ing are from America: — malvacearum Ckll. from Mexico on Malva, Hi-
biscus, cotton and vitrea Ckll. from Trindad on Acacea.
Cryptinglisia Ckll. — A single species, lounsburyi Ckll., occurs in
Cape Colony on the roots of Vitis.
Ceroplastodes Ckll. — There are five species listed in Fernald's
Catalogue. The following are from America: — acaciae Ckll. from Ari-
zona, New Mexico on Acacia; daleae Ckll. from New Mexico on Dalea;
and dugesii Sign, from Mexico on Mimosa.
Parafairmairia Ckll. — This genus includes a single species,
bipartita Sign, from France on Agropyrum, Mesembrianthemum.
Myxilecaiiium Beccari. — This genus was based upon the presence
of what was supposed to be a peculiar structure associated with the
rostrum, a crumena, which is found in most if not all coccids. A single
species is included, kibarae Beccari, from New Guinea on Kibara.
Lechanochiton Mask. — Two species from New Zealand are in-
cluded in this genus.
Neolecanium Ckll. & Parr. — The following American species have
been referred to this genus: — chilaspidis Ckll. from Mexico on Chilopsis;
O cornuparvum Thro from New York on Magnolia; herrerae Ckll. from
\ Mexico on Agave; imbricatum Ckll. from Mexico, New Mexico on Mi-
mosa; leucaenae Ckll. from Mexico on Leucaena, Mimosa; manzanillense
Ckll. from Mexico; perconvexum Ckll. from Brazil on Nectandra;
plebeium Ckll. from Mexico on Ficus; sallei Sign, from Mexico; silveirai
Hemp, from Brazil on roots of grape; tuberculatum Towns. & Ckll. from
Mexico on "cafetillo"; urichi Ckll. from Trinidad, Grenada, Brazil on
Smilax and from nests of ants.
Platinglisia Ckll. — A single species, noacki Ckll., is recorded
from Brazil on Myrtaceae and Laurus.
Pseudokeiines Ckll. — Two species are referred to this genus;
armatus Ckll. from Mexico on "Palo de gusano" and nitens Hemp, from
Brazil on Myrtus and Psidium.
Schizochlamidia Ckll. — A single species from Mexico on Mimosa,
mexicana Ckll. & Parr., is included.
Alecanopsis Ckll. — Maskell has described a single species, filicum
Mask., from New South Wales on Doodia.
Lecanium Burm. — The species included here are those referred
to the genus Coccus in Fernald's Catalogue. The following species
\\^ occur in America: — citricola Campbell from California on Citrus;
V, flaveolus Ckll. from New Mexico, Colorado on Pilea; hesperidum Linn.
from all parts of the world on a great variety of plants; incisus King
SUBFAMILY LECANIINAE 179
from South America on nutmeg; longulum Dougl. from New Zealand,
China, India, Mauritius, eastern United States, Mexico, West Indies,
Hawaiian Islands on Psidium, Bambusa, Acacia, Citrus, Ficus. etc.^
mangiferae Green from Ceylon, West Indies on Mango, cinnamon, nut^
meg, bread-fruit, etc.; melaleucae Mask, from Australia, China, Massa-
chusetts on Melaleuca; minimum Newst. from England, Mexico on Areca,
Abutilon, fan-palm; nanus Ckll. from Trinidad on "Balata"; pseudo-
hesperidum Ckll. from Canada on Cattleya; rubellus Ckll. from Jamaica;
schini Ckll. from Mexico on "Nancem"; terminaliae Ckll. from Jamaica,
Mexico on Terminalia; ventralis Ehrh. from California, Japan; viridis
Green from Ceylon, Brazil, Mauritius on Cinchona, Citrus, Gardenia, tea,
coffee, etc.
Eulecanium Ckll. — This genus contains over seventy species of
which nearly fifty are found in America and of these forty-five are re-
corded from the United States. It is likely that a considerable number
of these are synonyms. Six of the more common species can be sepa-
rated by means of the following table which is based for the most part
upon a table by Dietz and Morrison: —
SPECIES OF EULECANIUM
a. Adult female with median spina more than twice as long as either
lateral spina; dorso-meson with row of dorsal tubercles extending
cephalad from opercula; fringe setae four in number; operculum
with outer and basal margins subequal and slightly longer than
greatest width of operculum. — East of Rocky Mountains on peach,
plum, apple, maple, etc. nigrofasciata Perg.
aa. Adult female with median spina always less than twice as long as
either lateral spina; dorso-meson not with row of dorsal tubercles.
b. Each operculum distinctly more than twice as long as wide.
c. Alveolae promiscuously arranged, not in irregular transverse
groups; species large, ten to thirteen millimeters long. — East-
ern United States, Canada on apple, plum, linden, poplar, etc.
caryae Fitch.
cc. Alveolae arranged in irregular transverse groups; species
small, four to six millimeters long. — Europe, Indiana, Oregon
on apple, Crataegus. bituberculatum Targ.
bb. Each operculum never more than twice as long as broad.
c. Lateral spinae never twice as long as marginal setae of aver-
age length,
d. Alveolae arranged in more or less irregular bands.
e. Alveolae at least near middle of the dorsum large and
arranged in single rows radiating from meson; cuticle
nearly smooth, not wrinkled or roughened. — Eastern
United States, Canada on arbor-vitae fletcheri Ckll.
ee. Alveolae all subequal in size and small, arranged in ir-
regular rows radiating from meson; cuticle irregularly
roughened. — Europe, United States on Cornus, Ribes,
Corylus, Pyrus, Tilia. corni Bouche.
180 THE COCCIDAE
dd. Alveolae promiscuously arranged, not in transverse rows;
four fringe setae, mesal setae longer than lateral and ar-
ranged in two rows converging cephalad. — New York on
blackberry. obtusum Thro.
cc. Lateral spinae always twice or more as long as marginal setae
of average length; alveolae promiscuously arranged; four
fringe setae; arranged in single transverse row. — Eastern
United States and Canada on Quercus quercijex Fitch.
The species of the genus Eulecanium according to Sulc should be
distributed among four genera. These genera are based upon the
characters of the male, as follows: —
a. Species reproducing avongenetically, males with two pairs of waxy
abdominal filaments.
b. Males with six pairs of ocellanae. Eulecanium Ckll.
bb. Males with three pairs of ocellanae.
c. Males with wings and halteres present, with pocket for hook
of halteres. Palaelecanium Sulc.
cc. Males with wings and without halteres and pockets for hooks
of halteres. Sphaerolecanium Sulc.
aa. Species reproducing parthenogenetically, males not known
Parthenolecanium Sulc.
Sulc refers caprae L,, douglasi Sulc, ciliatum Newst,, and alni Mod.
to Eulecanium; bituberculatum Targ. to Palaeolecanium ; prunastri Fonsc.
to Sphaerolecanium; and coryli Linn, and persicae Geoff, to Partheno-
lecanium.
Globulicoccus Lindinger. — This name was proposed by Lindinger
as of subgeneric rank to include Eulecanium fuscum Gmel. from Europe
on oak, also capreae Linn, from Europe, Nova Scotia on Salix, poplar,
linden, Pyrus, Prunus, etc.
Mesolecaniuin Ckll. — An American genus of thirteen species.
One of these, phoradendri Ckll., occurs in Arizona on Phoradendron,
the other species are limited to Mexico and Brazil.
Stictolecanium Ckll. — This genus includes a single species, orna-
tum Hemp, from Brazil on Eugenia.
Saissetia Depl. — This genus contains over twenty species, several
of them of wide distribution. The three more important species occur-
ring in the United States can be separated by the following table based
upon the analysis of Thro: —
SPECIES OF SAISSETIA
a. Fringe setae eight in number.
b. Opercula with outer margins distinctly longer than basal mar-
gins, outer angle broadly rounded, fringe setae convergent toward
meson. — Europe, South Africa, Mauritius, Ceylon, China, Japan,
Australia, New Zealand, Hawaiian Islands, Brazil, Mexico, West
Indies, United States on orange, apple, pear, plum, olive, apricot,
etc. oleae Bern.
' SUBFAMILY LECANIINAE 181
bb. Opercula with' outer margins subequal in length with basil mar-
gins, outer angle not rounded, but angulate, fringe setae in trans-
verse row. — Europe, United States, Mexico, West Indies, Brazil,
Australia, etc. on ferns, Oleander, Gamellia, etc
hemisphaericum Targ.
aa. Fringe setae four in number; opercula with outer and basal mar-
gins subequal, outer angle angulate, fringe setae in transverse row. —
Europe, United States, Australia, Jamaica, etc. on ferns
filicum Bdv.
Platysaissetia Ckll. — A single species, castilloae Ckll., from
Mexico on Castilloa is included.
Cryptes Mask. — There is a single Australian species, baccatus
Mask., occuring on several species of Acacia.
Hemilecanium Newst. — Two species, imbricans Green from West
Africa on Theobroma and from India on Ficus and recurvatum Newst.
from Upper Congo, Africa from hollow stems of Plectronia, are included.
Eucalymnatus Ckll. — A genus containing the following four
species: — brunfelsiae Hemp, from Brazil on Brunfelsia, Laurus; gracilis
Hemp, from Brazil on Sapindaceae; perforatus Newst. from England,
Australia, Hawaiian Islands, Florida, Iowa, Colorado, California; and
tesselatus Sign, from England, France, Australia, Mauritius, Hawaiian
Islands, Ceylon, Jamaica on Laurus, Sapindus, lignum vitae, palms,
ferns, etc.
Paralecaiiium Ckll. — This genus contains seven species, one,
marianum Ckll. from Brazil, and the others from Australia, Ceylon, and
China.
Ericerus Westw. — This genus includes the single species, pe-la
Westw., which is a native of China and Japan where it feeds upon Lig-
astrum, Rhus, Hibiscus, and Fraxinus. The adult females are collected
by the Chinese and raised in order to secure the males in large numbers,
which produce a considerable quantity of wax. This is collected in
June and mixed v/ith tallow and used in the preparation of candles
which are burned in the sanctuaries. The Chinese not only cultivate
the plants upon which the females are grown but transport them a long
distance through dangerous mountain passes from Western China to
the regions where the wax is produced. The pure wax is clear white
in color like the best quality of paraffin and very hard, but not brittle.
The wax harvested each year by the Chinese, according to Sassaki, is
worth over two hundred thousand dollars. Paraffin candles are used at
the present time instead of the candles made from the coccid wax, ex-
cept on very state occasions.
Houardia Marchal. — A single species, troglodytes Marchal, from
Senegal, Africa, is found in the galleries in the branches of Balanites
where they are attended by ants.
Toumeyella Ckll. — This genus contains the following species:
magnoliarum Ckll. from California, Ohio on Magnola, Daphne, Virginia
creeper; mirabilis Ckll. from Arizona, Mexico on Mesquite; parvicornis
Ckll. from Florida on pine; pini King from Canada, New York on pine;
quadrifasciata Ckll. from New Mexico on Robinia; sonorensis Ckll. &
182 THE COCCIDAE
Parr, from Mexico on Beloperone; and turgida Ckll. from Florida on
Magnolia.
Akermes Ckll. — A genus containing ten species from Mexico,
South America, Australia, and India.
Megasaissetia Ckll. — Two species, inflata Ckll. & Parr, from
Mexico on "Laurel-tree" and brasiliensis Hemp, from Brazil, are known.
Physokermes Targ. — A genus containing the following five
species: — coloradensis Ckll. from Colorado on spruce, Pinus; concolor
Coleman from California on Abies; insignicola Craw from California on
Pinus; piceae Schr, from Europe, New York on Abies, Pinus; taxifoliae
Coleman from California on Pseudotsuga.
Aclerda Sign. — There are eleven species included in this genus.
The following are recorded from America: — ariditalis Ferris from New
Mexico on a perennial grass; californica Ehrh. from California, Kansas,
New Mexico on Andropogon; obscura Parr, from Kansas on Andropogon,
Sporobolus; and tokionis Ckll. from California, Japan on bamboo. Ac-
cording to Ferris the marginal spines in californica Ehrh. are acorn-
shaped, rounded at the tip in ariditalis Ferris, while the marginal spines
appear to be entirely wanting in obscura Parr.
CHAPTER XVI
SUBFAMILY ASTEROLECANIINAE
The Ceravitreous Coccids
The body of the adult female is round, oval, or elongate oval in out-
line. The dorsal aspect is convex or strongly convex and the ventral
aspect is flat. The lateral margins are sometimes convex, frequently
with a marginal rim and the two ends are convexly rounded or the
caudal end is truncate or produced. The segmentation is very indis-
tinct or not indicated. The caudal end of the body is not inflated and
the cephalic and lateral margins are not provided with a keel. The
antennae are normal in form, consist of four to six segments or each
is reduced to a single rudimentary tubercle, and are articulated to the
ventral aspect of the head. The eyes are wanting. The mouth-parts,
rostrum and rostralis, are present, the rostrum consists of a single seg-
ment. The legs are rudimentary, mere tubercles or wanting. The
spiracles of the mesothorax and metathorax are normal in size and
ventral in position. The mesothoracic spiracles are never located near
the anus. The abdominal spiracles are always wanting. The stigmatic
clefts may be present or wanting, when present, they are usually pro-
vided with one or two spinae with canellae and spiracerores. The abdo-
men is about as long as the head and thorax together or longer. It is
usually more or less convex on the dorsal aspect and flat on the ventral.
The lateral margins are either sharply defined or obscure or wanting.
The caudal end is rounded or truncate or more or less produced or
emarginate, but never with a mesal cleft and opercula. All the seg-
ments are fully exposed and similar in appearance with or without coriae
between them, none are retracted to form a marsupium. The abdomen
is never provided v/ith an anal cleft and opercula. The anal ring is
usually present and located at the cephalic end of a short retracted
anal tube and provided with six, eight, tv/elve, four, two, or without
anal ring setae. The anal lobes and the anal setae are usually present,
very different In size and form in the different genera. The caudal
abdominal segment can not be identified. The body is provided with
cerores, is never provided with pilacerores, and usually with octacerores,
and sometimes with ceratube. The body is never enclosed in a resinous
cell with three adjacent openings or covered by a scale with or without
exuviae but is usually covered by a test of wax, very diverse in form,
varying from a thin glassy sheet of wax with a glassy fringe to a rather
dense thick mass of wax or with the body naked. Insects sometimes
form depressions or galls on plants. The rectum is never provided with
a long glassy tube of wax or with a chitinized rectal tube bearing rings
of anacerores. The body even if naked is not gall-like in form.
183
184 THE COCCIDAE
The female nymphs of all stages are provided with a rostrum and
a rostralis. The three pairs of legs are present in the first nymphal
stage and in the second nymphal stage of some species and, when pres-
ent, are similar in form. The profemora are never greatly enlarged and
the prothoracic legs are never fitted for digging. The abdomen is
usually provided with octacerores, frequently present when wanting in
the adult, also provided with an anal ring and anal ring setae, the
number of anal ring setate variable, and with anal lobes and anal setae.
An anal cleft, opercula, and pilacerores are never present; ceratubae are
sometimes present. The caudal end of the rectum is not chitinized,
forming a rectal tube provided with one or more rings of anacerores
excreting a long glassy tube of wax. The caudel abdominal segment is
usually distinguishable and bears the anal setae.
The males have four ocellanae, two dorsal and two ventral. The
abdomen is not provided with long lateral filaments or tufts of wax
filaments or provided with long thread-like caudal filaments. The stylus
is long and slender.
This group was first recognized as a tribe by Targioni Tozzetti
when he proposed his generic division Lecanodiaspis and the tribal
name Lecanodiaspidites to include it. This subdivision was re-
jected by Signoret and Comstock. It was later adopted as a
subdivision of his Lecaniinae by Maskell and as a subfamily by
Cockerell in 1896 under the name of Asterolecaniinae. While it
was rejected by Green in 1896 in the first volume of his ''Coccidae
of Ceylon," he recognized it in the fourth volume published in
1909 after he had studied the Ceylonese species. There are less
than thirty-five species of Asterolecaniinae listed in Fernald's
Catalogue from America. The subfamily is not limited in its dis-
tribution, since species are found in practically all parts of the
world.
While this subfamily is apparently a well founded homogenous
group, all its genera do not offer an abundance of characters for
combining them into distinctive units. The most characteristic
feature is the presence of octacerores or the so called 8-shaped
dermal glands. These are formed by the close apposition or ar-
rangement of the cerores in pairs. The adjacent portions of the
margins of the pairs of cerores are often round, but frequently
more or less flattened, emphasizing the resemblance to a figure 8.
In the genus Asterolecanium there is one or two rows of octacerores
arranged around the margin of the body. It is from these octa-
cerores that is excreted the characteristic straight or curved glassy
threads of wax. This forms a margin which is a thin homogenous
sheet produced by a partial or entire fusion of the adjacent threads.
The covering of wax may be opaque and have a distinct waxy
SUBFAMILY ASTEROLECANIINAE 185
consistency, or the test may be dense and of considerable extent
bearing horn-like projections. The test is apparently, no matter
how extensive and complex, freely separable from the body ofthe
adult female. The species that form galls, as those belonging to
the genera Amorphococcus and Frenchia, and those that are at-
tended by ants have the body naked. The octacerores, while
usually confined to the dorsal aspect, are in a few species situated
upon the ventral aspect. They may be wanting in the adult
female, but are generally a characteristic feature of the first
nymphal stage where they are usually arranged in longitudinal
rows. There is considerable variation in the number and arrange-
ment of the octacerores, not only in the adult female but in the
nymphal stage. The test, in which there is an opening for the
escape of the young, is designated the ovisac by Newstead.
There are small tubes extending through the cuticle into the
body cavity which have been designated by Green as the tubular
glands or tubular dermal glands. He believes that the homogenous
part of the test is excreted from these. The cuticular tubes are
evidently similar to the tubes found in certain naked lecaniids
and although characteristic in form for this group are undoubt-
edly homologous with the ceratubae of the diaspidids.
While the species of certain genera form galls that are charac-
teristic in form, other species, as certain of those of the genus
Asterolecanium, form prominent crater-like depressions in the
bark in which the insect rests.
The antennae in the adult female are variable in development,
they may be fully formed and consist of eight or nine segments or
of only a few segments, four or less, or be completely wanting.
They are attached, when present, to the ventral aspect of the head
some distance from the lateral margin. The antennae in the first
nymphal stage consist of six segments.
The legs are usually wanting, or rudimentary in the adult
female, frequently only represented by mere hooks, but are well
developed in Olifiella. In the first nymphal stage and also in the
second in at least certain genera the legs are present and normal
in size, form, and arrangement of parts.
The extent of the thorax, since the legs are wanting in the
adult female in most species, can be determined only by the pres-
ence of the thoracic spiracles. They are normal in size. The
mesothoracic spiracles are located on a transverse line drawn
through the rostrum and the metathoracic spiracles in line with and
186 THE COCCIDAE
a short distance cavidad of the mesothoracic spiracles. There are
canellae with spiracerores which extend from the spiracles to the
margin of the body where there are one or two prominent spinae
in Lecanodiaspis and Anomalococcus. The figures of some authors
of other genera show spiracerores but the absence of spinae.
The dorsal surface of the abdomen in certain genera bears
some cuticular pitted or sieve-like plates, which are known as the
cribiform plates. These plates as dorsal structures are not found
outside the subfamily Asterolecaniinae. Ferris has noted the
presence of cribiform plates in Kuwanina Cocker ell, but these
plates are confined to the ventral aspect of the abdomen. The
cribiform plates are the madreporiform bodies of Comstock. The
function of these plates is unknown, their openings may be cerores.
There are six cribiform plates in Lecaniodiaspis, three on each side
distant from the meson ; two groups in Ceroroccus, one on each side
near the anal ring, each group consisting of four plates; a single
large group extending transversely in an irregular row near the
middle of the abdomen and consisting of one hundred to one hun-
dred and fifty cribiform plates. The surface of the plate is cup-
shaped in Cerococcus and flat in the other genera.
The caudal end of the abdomen may be only slightly indented
or emarginate or it may be deeply indented, in such cases it is said
to be deeply cleft. Such an emargination is different from the
anal cleft of the lecaniids, but is more like the emargination of
the eriococcids. The emargination appears to be formed by the
projecting anal lobes. That these structures are anal lobes is
shown from the almost universal presence of anal setae, the caudal
setae of Green, which in some species are very large and prominent.
The caudal end of the alimentary canal, the anus and anal
ring, are invaginated and form the bottom of a short tube or pocket,
the anal tube, which resembles somewhat the anal tube of the
lecaniids. The form and extent of this tube varies with the genus.
The caudal end of the anal tube is supported on the dorsal side by
a transverse strongly chitinized plate, the dorsal lip. This varies
from a simple transverse plate in Asterolecanium to a shield-shaped
plate in Cerococcus and an irregular crescentic or subcrescentic
plate with the caudal margin emarginate in Lecaniodiaspis. In
this latter genus there is a prominent dumb-bell-shaped thickening
on the ventral aspect caudad of the opening of the anal tube.
The number of anal ring setae is frequently not constant even
for the same genus. The normal number in Asterolecanium is six
SUBFAMILY ASTEROLECANIINAE 187
but may be reduced to two or even all may be wanting. There are
eight to twelve anal ring setae in Lecaniodiaspis, ten to sixteen in
Anomalococcus, eight in Cerococcus, wanting in Pollinia, and-^ight
in Amorphocoecus.
The eggs are deposited by the female in the test under the
body. The first stage nymphs are elongate, oval in outline with
the cephalic end bluntly rounded and the caudal end pointed. The
legs are present and normal in form. The antennae consist typ-
ically of six segments. The spinae, if present in the adult, are also
present in the nymphs. The dorsal lip and dermal plate, if charac-
teristic of the adult may be represented in the nymphs. The
octacerores are frequently more numerous and arranged in two to
six longitudinal rows. The anal ring is present and provided
typically with six anal ring setae. The anal lobes and anal setae
are usually present, but vary greatly in size and prominence in
the different genera. The female nymphs molt twice. The male
nymphs of the second stage may be provided with octacerores.
GENERA OF ASTEROLECANIINAE
a. Abdomen of adult female with dorsal cribiform plates.
b. Antennae of adult female well developed, consisting of four to
nine segments; spinae present at least in nymphs; anal lobes of
adult female small.
c. Spinae large and distinct in adult female; body of insect en-
closed in complete test; anal ring of adult female with eight
to twelve anal ring setae; abdomen with two rows of cribiform
plates. Lecaniodiaspis Targ.
cc. Spinae wanting in adult female, distinct in nymphal stages;
body of insect not enclosed in test, naked; anal ring of adult
female with ten to sixteen anal ring setae; abdomen with large
transverse group of cribiform plates near middle of the dorsum.
Anomalococcus Green,
bb. Antennae of adult female rudimentary, always consisting of less
than four segments; spinae never present in any stage; anal lobes
of adult female large and distinct with shield-shaped dorsal lip
between them; body of insect enclosed in dense test of wax with
or without projections; anal ring of adult female with eight anal
ring setae; abdomen with two groups of cribiform plates, each
cephalo-laterad of the dorsal lip. Cerococcus Comst.
aa. Abdomen of adult female never with dorsal cribiform plates,
b. Adult female living free, never enclosed in woody gall.
c. Body of adult female and first stage nymph always with octa-
cerores.
d. Body of adult female always with peripheral band of dorsal
octacerores, dorsum sometimes with other octacerores than
188 THE coccroAE
peripheral band; anal ring normally with six anal ring
setae, but may have two, four, or none; anal lobes and anal
setae small, sometimes wanting; first nymphal stage with
peripheral band of dorsal octacerores, antennae with six seg-
ments, and anal lobes and anal setae distinct
Asterolecanium Targ.
dd. Body of adult female never with peripheral band of dorsal
octacerores.
e. Dorsum of adult female with mesal band of octacerores;
anal ring obscure and apparently without anal ring setae;
first stage nymph with antennae of four segments, with-
out anal lobes and anal setae, without anal ring setae,
and with peripheral band of dorsal octacerores
Callococcus Ferris
ee. Dorsum of adult female with numerous octacerores, but
not arranged in peripheral or mesal row; anal ring small
and obscure; first stage nymph with antennae of five seg-
ments, anal lobes and anal setae small, anal ring small
with four minute anal ring setae, and with four dorsal
longitudinal rows of octacerores Mycetococcus Ferris.
cc. Body of adult female never with dorsal octacerores.
d. Adult female without prominent anal lobes or anal setae,
if present, small; anal ring with eight anal ring setae or
without; first stage nymph with four longitudinal rows of
octacerores, antennae with six segments, and anal ring with
six anal ring setae Pollinia Targ.
dd. Adult female with prominent anal lobes, with or without
anal setae; first stage nymph unknown or octacerores un-
identified.
' e. Adult female with prominent anal lobes bearing large
anal setae, longer than anal lobes; anal ring with eight
anal ring setae and without shield-shaped dorsal lip;
first stage nymphs with four longitudinal rows of cuticu-
lar circles, octacerores not identified, antennae with eight
or nine segments, anal ring with anal ring setae
Amelococcus Marchal.
ee. Adult female with large anal lobes bearing several small
setae, none as long as anal lobes, without distinct anal
setae; anal ring with six anal ring setae and with prom-
inent shield-shaped dorsal lip between anal lobes; anten-
nae atrophied, apparently with seven segments; nymphal
stages unknown. Olliffia Fuller.
bb. Adult female never living free, always enclosed in a woody gall.
c. Adult female with antennae of six segments and with distinct
legs; dorsal surface of adult female with numerous octa-
cerores. OlliffieUa Ckll.
cc. Adult female with antennae and legs wanting, antennae at
most only tuberculate; dorsal surface of adult female without
octacerores.
SUBFAMILY ASTEROLECANIINAE 189
d. Nymphs of first stage with four rows of octacerores,
antennae with six segments; adult female with anal ring
with eight anal ring setae; thorax with four groups of
spiracerores. Amorphococcus Green.
dd. Nymphs of first stage with six rows of octacerores, antennae
with six or seven segments; adult female apparently with-
out anal ring and anal ring setae, thorax apparently without
groups of spiracerores. Frenchia Maskell.
Lecaniodiaspis Targ. — There are less than twenty-five described
species in this genus. The following have been reported from America: —
celtidis Ckll. from Texas, Ohio on Celtis; dendrobii Dougl. from British
Guiana on Dendrobium; manihotis Towns, from Mexico on Manihot;
prosopidis Mask, from Arizona on Prosopis; pruinosa Hunter from Kan-
sas on elm; radiata Ckll. from Mexico; rufescens Ckll. from Arizona,
California on Adenostoma, Pouquera; rugosa Hemp, from Brazil; tessa-
lata Ckll. from Georgia, Florida on Disospyros, hickory; and yuccae
Towns, from New Mexico on Yucca, Dasylirion, Parthenium.
Anomalacoccus Green. — This genus includes a single species, cre-
mastogastri Green, from Ceylon from the nests of ants.
Cerococcus Comst. — About ten species belong to this genus.
There is only a single American species, quercus Comst., from California
and Arizona on Quercus.
Asterolecanium Targ. — There are twenty-seven species listed in
Fernald's Catalogue and a considerable number of species have been
described since its publication. The fellowing species occur in Amer-
ica:— aureum Bdv. from Europe, West Indies on Calathaea, Oncidium;
bambusae Bdv. from Brazil, Grenada, West Indies, Mexico, Algeria,
Ceylon, Mauritius on bamboo; epidendri Bouche from Europe, Jamaica,
Trinidad on Epidendrium, Oncidium, Angraecum, Broughtonia; fimbri-
atum Fonsc. from Europe, Australia, British Guiana on Leptospermum,
Coronilla; miliaris Bdv. from Brazil, Jamaica, Trinidad, Algeria,
Mauritious on bamboo; palmae Ckll. from Jamaica on cocoanut-palm ;
pustulans Ckll. from Florida, Mexico, West Indies, British Guiana on
fig, oleander, mango, peach, Hibiscus, Anona, etc.; quercicola Bouche
from Europe, eastern United States on oak; townsendi Ckll. from Mex-
ico on "Gusima"; urichi Ckll. from Trinidad, Grenada on palms,
Bactris; variolosum Ratz. from Europe, eastern United States, Canada
on oak; and viridulum Ckll. from Argentine Republic.
Callococcus Ferris. — A single species, pulchellus Maskell, from
Australia.
Mycetococcus Ferris. — Two American species are included,
ehrhorni Ckll. from California on oak and corticis Towns. & Ckll. from
Mexico on oak. These species were previously referred to Cerococcus.
Pollinia Targ. — A small genus of less than five species, the type
species, pollini Costa, has been introduced into California on olive.
Amelococcus Marchal. — A monotypical genus occurring in Mada-
gascar on Euphorbia. The single species, allaudi Marchal, was con-
sidered as an eriococcid. The form of the body, the anal lobes, the
position of the anal ring, the four longitudinal rows of circular tuber-
190 THE COCCIDAE
cles, and the enveloping of the female in a sac open only at the posterior
end are responsible for its location in this subfamily.
Olliffia Fuller. — There is a single species, eucalypti Fuller, from
West Australia on Eucalyptus. The genus is generally placed in the
Eriococcinae. The form of the anal lobes and the shield-shaped dorsal
lip, similar to that found in Cerococcus, suggested its present location
and the possibility that it may not be an adult female that is figured.
The nymphal stages are unknown.
OUiffiella Ckll. — A single species, cristicola Ckll., from New
Mexico, where it produces galls on the leaves of oak is included. The
presence of octacerores suggested its present location. This genus was
originally associated with ithe Australian gall making coccids of the
genus Cyiindrococcus and Sphaerococcus and later with the Eriococcinae.
The tarsi consist of a single segment according to Ferris and not two
as described by Cockerell. The antennae of the adult female consists
of five to six segments and of the first nymphal stage of six segments.
The anal ring of the adult female is without anal ring setae and the
first nymphal stage is provided with six anal ring setae. Ferris records
the presence of ventral octacerores and considers this genus as related
to Kermes.
Amorphococcus Green. — This is a Ceylonese genus, including a
single species, mesuae Green, on Mesua. Ferris has referred leptospermi
Maskell, described as a species of Sphaerococcus from New South Wales
on Leptospermum to this genus.
Frenchia Maskell. — Two species, casuarinae Maskell and semi-
occulata Maskell from Australia on Casuarina, belong to this genus. Its
present position was suggested as probable by Green.
/
vaO
CHAPTER XVII
SUBFAMILY KERMESIINAE
The Pseudogall Coccids
The body of the adult female is rotund, globular, not insect-like. It
resembles a gall or berry and is either uniformly smooth or subdivided
by broad shallow furrows. The cephalic, lateral, and caudal margins
can not be identified. The coriae are uniformly chitinized and are sim-
ilarly colored with the segments so that the segmentation is obliterated.
The caudal end of the body is not inflated and the cephalic and lateral
margins are not provided with a keel. The antennae are distinct, con-
sist of six segments or are greatly reduced and rudimentary or are
wanting, when present, are articulated to the ventral aspect of the head.
The mouth-parts, rostrum and rostralis, are present but inconspicuous,
the rostrum consists of two segments. The legs are present. They are
subequal in length, rudimentary, or wanting. The profemora are never
enlarged and the prothoracic legs are never fitted for digging. The
trochanto-femoral and tibio-tarsal sutures of all normally formed legs
are always distinct. The spiracles of the mesothorax and metathorax
are small and inconspicuous and are ventral in position. The mesotho-
racic spiracles are never located near the anus. The abdominal spir-
acles are always wanting. The stigmatic clefts, spinae, and canellae are
wanting. The extent of the head, thorax, and abdomen can not be de-
termined. The ventral aspect of the body is depressed or concave to fit
the surface of the host-plant, the other aspects are uniformly convex
without division between them. The caudal end is uniformly convex.
The segmentation of the abdomen can not be determined, but no part is
invaginated to form a marsupium. The abdomen is never provided with
an anal cleft and opercula. The anal ring and anal ring setae are want-
ing. The anal lobes are rarely present. The caudal abdominal segment
can not be identified. The body is provided with cerores none of which
are octacerores or pilacerores. Ceratubae are sometimes present. The
body is never enclosed in a resinous cell with three adjacent openings
or covered by a scale with or without exuviae. The surface of the body,
dorsal and particularly the ventral aspect, sometimes bears a slight
covering of mealy or pulverulent wax. The body is usually naked and
gall-like, rarely setaceous or slightly covered with seta-like threads of
wax.
The female nymphs of all stages are provided with a rostrum and
rostralis. The three pairs of legs are present in the first nympal stage
and are similar in form. The profemora are never greatly enlarged
and the prothroracic legs are never fitted for digging. The body is
never provided with an anal cleft and opercula, octacerores, or pila-
191
192 THE COCCIDAE
cerores. The anal ring is distinct in the first nymphal stage and bears
six anal ring setae. The anal lobes are distinct and each bears a
single large anal seta. The caudal end of the rectum is not strongly
chitinized, forming a rectal tube with one or more rings of anacerores
excreting a long glassy tube of wax. The caudal abdominal segment
is not short, narrow, and projecting.
The male has ten ocellanae, five on each side of the head. The ab-
domen is not provided with long lateral filaments or with tufts of long
wax filaments, but bears two long caudal filaments. The stylus is short.
This subfamily contains a single genus, Kermes, and about
forty species all of which with only two exceptions feed upon
various species of oak. The single species found in Australia feeds
upon Acacia and one of the Japanese species feeds upon Quercus
and Rhamnus. The great majority of the species are found in
Europe and America, with the larger number in the latter country.
One of the species, Kermes ilicis, is found abundantly upon
a small species of live oak, Quercus coccifera, in southern Europe.
It has been used as a dye from the earliest times for dyeing cloth
blood red or crimson. The Phoenecians before the time of Moses
used it as a dye and it was also used in ancient times by the
Greeks, Romans, Arabians, and Persians. The latter people knew
it under the name of Kermes. The crimson of the Greeks and
Romans was produced with this color and the imperishable reds of
the Brussels and other Flemish tapestries were produced with this
same dye. It was universally used before the discovery of the
new world and the introduction of cochineal from America by the
Spaniards.
The adult females, which have the body naked or rarely
covered with a slight bloom of mealy wax or sparsely pubescent,
are usually attached to the smaller branches of young trees. The
insects vary greatly in color, brownish or blackish or dirty yellow-
ish, with or without darker markings, which are generally arranged
as light or dark transverse bands. The portion of the body applied
to the plant is flat or concave or irregular so that the insect fits
securely against the bark of the branch. The most of the species
are globular or subglobular in form and resemble very closely the
minute smooth bullet galls found on the stems of the same trees.
The female of Kermes resembles berries or galls so closely in general
appearance that they are frequently regarded or mistaken for galls
and not as the bodies of insects. This resemblance is enhanced
through the fixation of the insects in the angles or near buds,
where the bullet galls are usually located. The species of this
SUBFAMILY KERMESIINAE 193
subfamily have been defined in all the diagnoses of the subfamilies
as body naked and gall-like in form. It is not only unwise^ but
improper to designate the body of any of the naked coccids as the
scale as is frequently done by certain writers. The body in some
species is slightly longitudinally or transversely constricted at
middle or furrowed, so that it is more or less reniform. A few
species resemble certain of the naked lecaniids, others are elongate
and more swollen. The species also vary in that some occur singly
or at most in groups of three or four upon a branch while others
occur in such numbers as to completely cover a considerable por-
tion of the stem.
The appendages are present on the ventral aspect of the adult
female. The rostrum consists of two segments. The antennae and
legs of the adult female may be well developed. The former con-
sisting of six or less segments, or they may be rudimentary or
wanting. The thoracic spiracles are present, normal in position,
and surrounded by numerous spiracerores. The anal lobes and the
anal setae are wanting as well as the anal ring and the anal ring
setae. The ventral aspect of the body may bear numerous cerores
from which is excreted the white wax located between the body of
the female and the host-plant, but this wax is generally not ex-
posed until the insect is removed. The exposed surface of the
body may bear cerores from which is excreted the powdery wax
which is spread over its surface. Some species bear numerous
short ceratubae. The genus Kermes was originally associated with
the lecaniids. In order to justify this location, it was necessary
to show the presence of an anal cleft and opercula. Some writers
have indicated the presence of such structures, but these structures
are wanting in most species and it is very doubtful if they or any
other structures similar to them occur in any species. Maskell in
defining his group Hemmicoccidinae, which is the equivalent of
the Kermesiinae, makes the following statements: — ** Adult female
exhibiting the anal cleft and the lobes of the Lecanidae ; naked or
covered. Larvae presenting at the extremity of the abdomen the
anal tubercles of the Coccidae. From the foregoing characters the
group is very evidently intermediate between the Lecanids and
Coccids."
While the adult females of certain species of Kermes do show
a superficial resemblance to the naked lecaniids, the first stage
nymphs show clearly their true relationship. They are distinctly
eriococcid, not only in general appearance, but in structure. The
194 THE COCCIDAE
body is elongate oval in outline. The antennae are elongate and
consist of six segments which bear numerous setae. The mouth-
parts are well developed and located between the prothoracic legs.
The legs, which are similar in size, are long and slender and project
for some distance beyond the lateral margins of the body. They
are provided with long claws and with two tarsal and two ungual
digitules. The caudal end of the body is provided with two anal
lobes, each of which bears a large anal seta and usually several
much shorter setae. The anal ring is distinct and located cephalad
of the anal lobes. It bears anal ring setae, typically six, and is
sometimes divided into two parts. The entire lateral margin is
fringed with marginal setae. In a few species the marginal setae
are conical in form. There are four longitudinal rows of similar
setae on the dorsum making six longitudinal rows of dorsal setae,
while Signoret figures conical marginal setae in some of the
European species, arranged so as to form two disconnected rows on
each side, and six dorsal longitudinal rows, making eight in all.
This gives an arrangement similar to that found in certain
young eriococcids. There are cerores arranged in rows between
the conical setae. Practically nothing is known regarding the
second stage female nymphs. Maskell states that they are semi-
globular and yellow or brown. The number of molts in the female
has not been determined.
Observations by one of my students, J. Howard Gage, would
suggest that the second nymphal stage may be wanting. The
young nymphs emerge from the eggs in the fall and creep into
minute cracks and crevices near the ends of the smaller twigs,
covering themselves with curly threads of wax. They remain as
minute pinkish specks in such situations without change until
spring. As the weather gets warm they leave their place of con-
cealment and covering of wax. After the buds have opened and
the new twigs have been formed and are about an inch or two in
length, the nymphs migrate out onto the tender branches and onto
the petiole and surface of the leaf. Those on the branches and
petioles attach themselves, but most of those on the leaves migrate
back onto the branches and petioles. Early in June the body of
the insect is swollen, globular, and is beginning to assume the
characteristic form of that of the adult female. They are still
whitish or greenish in color and do not yet have the brownish
color of the adults. The growth after their emergence from hiber-
nation is a very gradual one and if molts occurred, although par-
SUBFAMILY KERMESUNAE 195
ticular care was taken to observe them, none was noted. If the
body of the dried female is opened in the fall, it will be found
completely filled with white empty egg shells. The young nypmhs
resemble miniature mealy bugs.
The adult males according to Signoret resemble the males of
the Eriococcinae. They complete their final transformations in
small white felt-like cocoons of wax attached to the ventral surface
of the leaves. Comstock figures the cocoons of the males of
galliformis Riley on the underside of the leaves. Nothing of their
early development appears to be known. The head bears ten
ocellanae, five on each side of the head. The dorsal and ventral
pairs are much larger than the others. The antennae are very
long and consist of ten segments. The wings are long and the
halteres are distinct with distinct hooks. The legs are long and
slender, the femur and tibia are subequal and each is much longer
than the tarsus which bears a simple claw and two tarsal and two
ungual digitules. The abdomen is elongate. The stylus is about
as broad as long and the caudal end bears two long thread-like pro-
jections, longer than the entire body and evidently formed of wax.
Kermes Boit. — A considerable number of Americair species of the
genus Kermes have been described. These in most cases are based upon
the form and color of the gall-like or berry-like body of the adult fe-
male. The following table for the separation of these species is based
upon the tables and descriptions of Cockerell, King, and Ehrhorn. The
table must be used with caution, unfortunately it includes practically all
the information given in the various descriptions. In every case where
tljie body of the insect is referred to, the body of the adult female is
meant.
SPECIES OF KERMES
a. Body either sparsely pubescent or covered with wax,
b. Body sparsely pubescent and naked; 3.6 mm. broad by 3 mm.
high; light brown with obscure and suffused dark brown bands,
surface shining, with concolorous specks, without dark spots or
pits. — Massachusetts, Ohio, Indiana, Kansas on Quercus macro-
carpa. pudescens Bogue.
bb. Body without setae but covered with wax.
c. Body covered, except on middle of back, with snow-white
powdery wax; 4.5 mm. broad and 3 mm. high; resembles small
species of Lecanium in shape; dark sepia brown in color and
irregularly marbled with black and pale ochreous, beset with
numerous minute dark spots. — Massachusetts on Quercus alba.
nivalis King & Ckll.
cc. Body covered with wax, but never snowy white,
d. Body over three millimeters in diameter.
196 THE COCCIDAE
e. Body 5 mm. in diameter, color mahogany brown, shining,
with three distinct dark brown lines across dorsum, black
spots apparently wanting. — California on Quercus chryso-
lepsis. shastensis Ehrh.
ee. Body 4 to 4.5 mm. in diameter; color brown, shining,
dotted with black spots. — Arizona on Quercus
ceriferus Ehrh.
dd. Body three millimeters in diameter; color dark gray, dull
not shining, with fine black lines upon dark yellowish brown
surface and with depressed darker spots. — Massachusetts on
Quercus ilicifolia. perryi King.
aa. Body always naked, never sparsely setaceous or covered with wax.
b. Insects of large size, 7 to 10 mm. in diameter.
c. Body v/ith transverse furrows and with rounded protuberances,
with three protuberances, one mesal in each transverse row;
8 mm. long and 7.5 mm. broad; color white with dark brown
mottling, usually with white dorsal band. — Colorado, New
Mexico on Quercus undulata. gilletti Ckll.
cc. Body not with transverse furrows and without protuberances.
d. Body globular, 10 mm. diameter; surface dull, peppered or
mottled irregularly with black, grey and ^^hite, white areas
with numerous minute brown spots. — Mexico on Quercus
engelmanni. grandis Ckll.
dd. Body subglobular, 7 mm. in diameter; marked with five
transverse black bands composed of large round dots and
fine lines, pebbled and marbled with black dots and specks
between lines and dots, entire surface covered with minute
black specks. — California on Quercus. miraMlis King.
bb. Insects of smaller size, 3 to 6 mm. in diameter.
c. Body convex without median longitudinal constriction.
d. Body strongly convex, 5 mm. in diameter and 5 mm. high,
distinctly pyriform; surface shining, with three to five dark
lines, color light brown with three to four dark brown
bands of variable length and suffused dark brown, blotchy
spots and round dots. — Massachusetts, Ohio, Indiana, Georgia
on Quercus. andrei King.
dd. Body globular or nearly so, never pyriform or conical,
e. Body variously marked, but not with light or dark bands,
f. Body strongly elevated, 5 to 6 mm. high.
g. Body dark, dirty gray, turning to nearly clear white
in old weathered specimens, with black spots and
minute black specks; 6 mm. high and 6 mm. long. —
East of Rocky Mountains on Quercus
galliformis Riley.
gg. Body dark, irregularly marbled with white and
black and reddish or black and reddish, minute
SUBFAMILY KERMESIINAE 197
points on white intensely black. — Oklahoma on
Quercus alba. hoguei Ckll.
ff. Body not so strongly elevated, 3.5 mm. high; light
ochreous, marked with slightly darker redder tint,
marking wanting on mid-dorsal line, leaving pale band
with transverse rows of black spots, surface closely
beset with minute dark dots. — Massachusetts, Delaware
on Quercus rubra. kingii Ckll
ee. Body always marked with light or dark bands,
f. Body marked with transverse lighter stripes.
g. Body light brown, with several white stripes run-
ning parallel with segments, more or less pitted,
pits dark brown or black; 4.5 mm. broad and 4 mm.
long and 4 mm. high. — California on Quercus
oblongifolius. austini Ehrh.
gg. Body light brown, generally with four transverse
parallel white stripes, stripes with numerous brown
dots; 5 to 6 mm. in diameter. — California on Quercus
chrysolepsis. rattani Ehrh.
ff. Body marked with dark transverse stripes,
g. Body with four dark transverse stripes.
h. Body grayish white, distinctly marbled with
light yellow or reddish brown, with four promi-
nent linear transverse dark brown bands, some-
what wavy, due to quite large pits, surface
speckled with minute black dots; 3 to 5 mm.
broad. — Arizona on Quercus arizonensis King.
hh. Body yellowish brown, with four transverse very
dark brown bands on meson, marbled with darker
brown, some of marbling circular, surface marked
with some dark brown spots; 5 mm. in diameter.
— Connecticut on Quercus waldeni King.
gg. Body with five transverse narrow blackish bands,
broken at intervals by somewhat larger round black
dots, surface marbled light gray-brown between
bands and cot ered with minute black specks. — Cali-
fornia on Que 'cus. occidentalis King.
CO. Body with more or lesa distinct median longitudinal constric-
tion or furrow,
d. Body strongly gibbous or with humps.
e. Body light brown without conspicuous black specks, dor-
sum usually marked with black lines and spots; 5 mm.
long and 4 mm. broad and 4 mm. high. — California on
Quercus lobata. cockerelU Ehrh.
ee. Body light brown with four transverse bands of light
cream color, constriction not pallid, surface peppered with
minute black specks; 6 mm. in diameter and 5 mm. high.
— California on Quercus agrifolia essigi King.
198 THE COCCIDAE
dd. Body not gibbous or humped.
e. Surface of body not speckled with black, color lively
ochreous, with bands and spots of dark brown and black,
hind part with numerous pits. — Kansas on Quercus
macrocarpa. concinulus Ckll.
ee. Surface of body speckled.
f. Body with definite specks and other coloring but not ar-
ranged in transverse rows.
g. Body not globular, somewhat transverse, median
furrow forming rather broad black or dark brown
band, generally crossed by narrow blackish or
brownish transverse lines of various length, between
lines several black specks as large as pin heads, en-
tire surface covered with minute black specks, color
white marbled with gray, turning to nearly white,
light lemon yellow in museum specimens. — Massa-
chusetts on Quercus rubra sassceri King.
gg. Body globular, not very pale ochreous, speckled all
over with minute black specks, with more or less
pallid transverse bands, median constriction obscure;
4.5 mm. long and 5.5 mm. broad and 4 mm. high. —
California on Quercus nigropunctatus Bhrii.
ff. Body with specks and other coloring arranged in trans-
verse rows.
g. Body small, 4 mm. long and 4 mm. broad and 3.5
mm. high; french yellow in color with black spots
and obscure black specks; median constriction
marked by dark line, with distinct rows of black
spots. — New York, Indiana, Massachusetts, Canada,
Ohio on Quercus alba and Quercus rubra
pettiti Ehrh.
gg. Body large, at least 5 mm. long and 6 mm. broad.
h. Body varying in color from bright argillaceous to
dull gray, more or less conspicuously speckled
with black with bands of dark spots; median
constriction crossed more or less by dark lines,
a rounded dark spot on each side of front and
elongated blotch a short distance above and be-
low anal opening; 5.5 mm. long and 6 mm. broad
and 4.5 mm. high. — East Rocky Mountains on
Quercus nigra. trinotatus Bogue.
hh. Body varying from light cream color to nearly
white, often with bluish cast, with four broad
transverse bands of light yellowish brown, sur-
face thickly covered with minute black specks;
median constriction very disitinct on caudal por-
tion; 5 mm. long and 6 mm. wide and 5 mm. high.
— California on Quercus chrysolepsis
hranagani King.
CHAPTER XVIII
SUBFAMILY APIOMORPHINAE
The Pegtop Coccids
The body of the adult female is turbinate or shapfed like a peg-top,
rarely like a spherical top, usually three or four times as long as broad,
only rarely broader than long. The lateral margins are usually convex
and are not margined. The segmentaition of the body is usually dis-
tinct. The caudal end of the body is not inflated and the cephalic and
lateral margins are not provided with a keel. The antennae are rudi-
mentary, consist of three to five inconspicuous segments or are wanting.
They are articulated to the ventral aspect of the head, distant from each
lateral margin and near to each other. The eyes are inconspicuous.
There is apparently an ocellana located on each lateral margin. The
mouth-parts, rostrum and rostralis, are present. The rostrum consists
of two segments. The legs, when present, are greatly reduced in size
and rudimentary, the prothoracic legs are the smallest and the meta-
thoracic the largest. The coxae may be greatly swollen, the femora en-
larged, and the trochanters and claws may be wanting so that the leg
consists of only three segments. The profemora are never greatly en-
larged and the prothoracic legs are never fitted for digging. The tro-
chanto-femoral and tibio-tarsal sutures are present or obsolete. The
thoracic and abdominal spiracles are not definitely described and are not
figured. The mesothoracic spiracles are never located near the anus.
The stigmatic clefts, spinae, and canellae are wanting. The abdomen
is variable in form and length, longer than the head and thorax com-
bined in some species, shorter in others, or reduced to a mere projection
in a few. It is usually convex on the dorsal and ventral aspects, never
depressed. The lateral margins are convex and are never margined.
All the segments are fully exposed and similar in appearance with dis-
tinct coriae between them, none are retracted to form a marsupium.
The abdomen is never provided with an anal cleft and opercula or an
anal ring and anal ring setae. The anal lobes are long and prominent,
frequently strongly chitinized, but distinct anal setae are wanting. The
caudal abdominal segment is long and distinct and not short and narrow
and extending beyond the other segments. The body is provided with
cerores, none of which are octacerores or pilacerores, and is never pro-
vided with ceratubae. The body is never enclosed in a resinous cell with
three adjacent openings or covered by a scale with or without exuviae.
Insects always live in galls and body is covered with mealy or powdered
wax. The rectum is never provided with a long glassy tube of wax or
with a chitinized rectal tube bearing rings of anacerores. The body is
never naked and gall-like in form.
199
200 THE COCCIDAE
The female nymphs of all stages are provided with a rostrum and
rostralis. The three pairs of legs are present in the first nymphal stage
and in at least certain species in the second. The legs of the first
nymphal stage are similar in form, those of the second are not, the pro-
thoracic legs are smallest and the metathoracic are the largest, the pro-
thoracic and mesothoracic are much farther apart than the mesothoracic
and metathoracic. The profemora are never greatly enlarged and the
prothoracic legs are never fitted for digging. The hody is never provided
with an anal cleft and opercula, octacerores, pilacerores, or ceratubae.
The anal ring is distinct and hears six anal ring setae. The anal lobes
are inconspicuous in the first nymphal stage, but the anal setae are large
and prominent. The anal lobes are prominent projections in the second
nymphal stage and the anal setae are small. The body is depressed or
subdepressed, ellipitical or oval in outline, and completely margined with
flabellate scales. The caudal end of the rectum is not strongly chitinized,
forming a rectal tube with one or more rings of anacerores excreting a
long glassy tube of wax.
The male apparently has four ocellanae, two dorsal and two ventral.
The abdomen is not provided with long lateral filaments or with tufts
of long wax filaments, but is provided with two white filaments at the
caudal end, which are probably formed of wax. The stylus is stout.
The insects of this subfamily all produce galls. The species
are confined to Australia and New Zealand. The great majority
make their galls on different species of Eucalyptus, less than one
per cent produce galls on plants of other species. Both sexes
form galls and, what is very unusual, the galls formed by each
sex is different in size and shape. As a rule the males make their
galls on the leaves and the females on the stems and branches,
although there are some exceptions to this. The galls are appar-
ently in every species abnormal growths produced from the cam-
bium layer of the plant and not through modification of the buds.
The insects evidently attach themselves to the surface of the
leaves or stems where they normally produce their galls. This
effects in some way the cells of the plant and an abnormal growth
begins which surrounds and eventually covers them, forming the
mature gall. Whether the galls jare due to irritation produced by
the mouth-parts or from excretions from the salivary glands, Mal-
phigian tubules, rectal glands, or some other internal structure is
not known.
The galls produced by the females of the different species
show a great variety of forms, varying in size from one-half to
seven or eight inches in length and in some species may be as
much as three inches in diameter although most are much smaller.
Some resemble conical fruits, others nuts and globular fruits.
While all the galls of a species are fairly constant, there is variation
SUBFAMILY APIOMORPHINAE 201
in size and form and in the case of certain species, as with all
gall forming groups, there is some variation between the individ-
uals of the same species. The difference in the form of the various
galls serves as one of the easiest methods of identifying the differ-
ent species.
The galls of the males are short cylindrical tubes. Some are
simple tubes with a notched rim at the free end or with the rim
dilated forming a saucer-like tip. While the male galls are gen-
erally attached to the leaves, in a few species they are attached
to the gall of the female.
The nymphs of the first stage are quite similar in appearance.
They are bright or pale red or yellow in color. The body is flat
or depressed, and elliptical or broad oval or almost round in outline.
The legs are long and well developed in Apiomorpha, normal in
form, bear several setae and a pair of digitules. In Opisthoscelis
the prothoracic legs are wanting and the mesothoracic legs are
rudimentary and the metathoracis legs are long and distinct. The
antennae are slender and consist of three to seven segments. The
single ocellana on each side of the head is large and prominent,
and brownish red in color. The periphery of the body is distinctly
margined or fringed with minute alate setae or scales, similar in
size to the marginal scales of certain lecaniids, but differing from
them in form in that their distal ends are truncate. Fuller de-
scribes the scales as stout alate spines, the wings of which are thin
and transparent and soon broken. This would suggest that the
wings might be wax excreted from marginal cerores supported by
a seta as a central matrix. The scales are apparently limited to
the head in Opisthoscelis. Froggatt in describing the nymphs of
Ascelis and Opisthoscelis states that their tarsi are each provided
with two claws, which is a very unusual number for the members
of this family, and is evidently an error. The first stage nymphs
are provided with an anal ring with anal ring setae, but the num-
ber of anal ring setae is not stated, it is probably six. The male
nymphs differ from the female nymphs according to Froggatt only
in being more elongate.
The female apiomorphids molt twice. The nymphs of the
second stage are very different in appearance from those of the
first. The body is broadest through the mesothorax, the two
lateral margins converge cephalad and caudad from this region.
The segmentation is distinct, but the margins have lost their
marginal rim and their fringe of flattened scales. The antennae
202 THE COCCIDAE
are greatly reduced in size and the ocellanae are apparently want-
ing. The legs are also reduced, the prothoracic pair is directed
cephalad and the others are directed caudad. The distance be-
tween the first and second pairs is much greater than that between
the second and third. The abdomen tapers and the caudal segment
is long and deeply cleft, the portion on each side of the cleft forms
a stout, divergent, sharply pointed projection bearing several
setae. Each of these projections is a modified anal lobe. The
cephalic part of the ventral aspect bears a distinct anal ring with
six anal ring setae. The surface of the body is covered with mealy
wax.
The body of the adult female has been generally compared by
the writers on the Apiomorphinae, so far as their shape is con-
cerned, to a peg-top. It is yellow or brownish in color with a
distinct segmentation, the distinctness of which is emphasized by
the constrictions between the segments. The head and prothorax,
as is usual with coccids, are poorly separated and form a large
cephalic region bearing the mouth-parts, the antennae, and the
prothoracic legs when these are present. There is a deep trans-
verse furrow on the ventral aspect separating the mesothorax and
metathorax, from which there extends on the meson a broad furrow
connecting with the curving furrow between the prothorax and
mesothorax. This latter has been called the facial furrow by
Fuller. The furrows produce a chin-like elevation upon which the
mouth-parts are situated. There is considerable variation in the
width of the mesothorax, which is usually the widest segment of
the body. There is a corresponding variation in the shape of the
facial furrow and its surrounding parts. The antennae and legs,
even when present, are greatly reduced and the antennae some-
times consist of only three segments. Fuller states that the tho-
racic spiracles are distinct in the adult females of Ascelis and
Cystococcus. The abdomen is conical and the three or four caudal
segments are strongly chitinized. It is usually as long as the head
and thorax together but in Cystococcus the abdomen is a mere
tubercle. The anal ring and anal ring setae are apparently want-
ing in the adult female. The caudal margin of each of the
abdominal segments is usually fringed with a transverse row of
conical spine-like projections, this appears to be always true in
Apiomorpha. These projections are probably used by the insect
in moving about in its cell in the gall. The caudal end of the
abdomen bears two long slender anal lobes in Apiomorpha and in
SUBFAMILY APIOMORPHINAE 203
some species of Opisthoscelis, but are wanting in Ascelis. It is
the presence of these lobes that gives the caudal end of the body
of the insect a cleft appearance. It is claimed by Schrader that
the anal lobes are used to keep the orifice in the gall open. This
orifice is in the free end of the gall. While the anal lobes bear
numerous setae, none of them can be identified as the anal setae.
The body bears long slender setae, conical setae, and cerores.
From these latter there is excreted powdery or mealy wax which
completely coats the body of the insect.
The galls formed by the males have already been described.
The males as they are figured by Froggatt resemble miniature
two-winged ant-lions. The wings as represented are much broader
than in the case of other male coccids. The abdomen is long,
slender, and cylindrical and bears two long thread-like filaments at
the caudal end which are twice as long as the entire body and are
probably formed of wax. The antennae are long, setaceous, and
consist of about ten segments. Froggatt states that, ''when full
grown the males emerge from the neighboring galls, and by means
of their slender, pointed abdomen impregnate the imprisoned
female through the apical orifice, through which the latter can
exsert her anal appendages. After impregnation the males die,
and the females become a mass of eggs, from which the young
larvae soon emerge, crawling through the opening in the gall and
leaving the empty shell of the mother behind in the gall."
The writers upon the apiomorphids have \]not giiven any
synopsis of the species, but have contented themselves with de-
scribing the galls, the adults, and certain of the nymphal stages.
Froggatt makes the following comment as to the females: ''To a
casual observer the female coccids would appear very much alike
but though there is a very strong resemblance in most cases, there
are besides the difference in form, several very good specific peculi-
arities; firstly, in the form, shape, and situation of the anal append-
ages; secondly, in the hairy coating on the abdominal segments;
and thirdly, in the number, shape, and regularity of the distribu-
tion of the tubercles and fine-toothed spines covering the upper
side of the abdominal segments. ' '
The four genera of Apiomorphinae can be distinguished by
means of the following table : —
204 THE COCCIDAE
GENERA OF APIOMORPHINAE
a. Adult female with at least one pair of legs.
b. Adult female with three pairs of legs and rudimentary antennae
of three to five segments; abdomen elongate, longer than head
and thorax together, disitinctly segmented; body tapering or pear-
shaped. ApioTnorpha Rubs.
bb. Adult female with prothoracic legs wanting, mesothoracic vesti*
gial, and metathoracic legs very long and slender; antennae pres*
ent and vestigial, consisiting of three to four segments or want-
ing. Opisthoscelis Schrader.
aa. Adult female without legs or antennae; abdomen very short, much
shorter than either head or thorax, segmentation obsolete, head and
thorax globular, abdomen appearing as tubercle upon one side of
globule.
b. Body not longitudinally striate, globular but not with cephalic end
truncate; spiracles present. Ascelis Schrader.
bb. Body longitudinally striate, globular with cephalic end truncate;
thoracic spiracles large Cystococcus Fuller.
Apiomorpha Rubs. — This name was proposed by Riibsamen for
the genus Brachyscelis Schrader, which was preoccupied. By far the
larger number of species of the subfamily belong to this genus. There
are thirty species listed by Mrs. Fernald.
Opisthoscelis Schrader. — A genus of twelve species, all of which
produce galls upon Eucalyptus.
Ascelis Schrader. — The adult females of this genus are greatly
modified. There are four species described.
Cystococcus Fuller. — This genus was erected by Fuller to include
his species echiniformis which forms a spherical gall from three-fourths
to one inch in diameter. There is some question as to whether this
genus can be distinguished from Ascelis. The walls of the gall are thin
and uniform in thickness. The body of the female nearly fills the cav-
ity in the gall. These galls according to Fuller are not only edible but
are considered as a great delicacy by all the natives and all whites who
know of them. The galls are broken between the teeth and the insect
sucked out. They are considered as cool and refreshing on a hot day.
Unless better characters are discovered than have been noted, the genus
Cystococcus will fall as a synonym of Ascelis which has the thorax glob-
ular and the abdomen a mere protuberance on one side.
CHAPTER XIX
SUBFAMILY CYLINDROCOCCINAE
The Lubberly Coccids
The body of the adult female is variable in form, circular and thick-
ened, spherical or globular, or cylindrical and elongate. The lateral
margins are broadly convex or subvertical, rarely if ever sharply mar-
gined. The cephalic and caudal ends are broadly rounded, sometimes
continuous with the lateral margins, so that the outline of the body is
almost a complete circle, or truncate. The segmentation may be dis-
tinct, obscure, or wanting, rarely sharply defined. The caudal end of
the body is not inflated and the cephalic and lateral margins are not
provided with a keel. The antennae are variable in form, when pres-
ent, consist of less than seven segments, frequently rudimentary, mere
tubercles, or wanting. They are articulated to the cephalic part of the
ventral part of the head. The eyes are apparently wanting. The mouth-
parts, rostrum and rostralis, are always present, the rostrum consists
of two segments. The legs are present or wanting, when present, they
may be subequal in size and similar or may have one pair much longer
than the other two pairs or they may all be tuberculate. The profemora
are never enlarged and the prothoracic legs are never fitted for digging.
The trochanto-femoral and the tibio-tarsal sutures of all normally formed
legs are always distinct. The spiracles of the mesothorax and metatho-
rax are large and prominent, and ventral in position. The mesothoracic
s.piracles are never located near the anus. The abdominal spiracles are
always wanting. The stigmatic clefts, spinae, and canellae are wanting.
The abdomen is variable in length, about as long as the head and thorax
together or longer or not as long as the metathorax alone, never de-
pressed, but may be thick and fiattened. The lateral margins are uni-
formly convex without a margin and the caudal end may be rounded or
truncate. All the segments are fully exposed and subsimilar in appear-
ance, none are retracted to form a marsupium. The abdomen is never
provided with an anal cleft and opercula, an anal ring and anal ring
setae, or anal lobes and anal setae. The caudal abdominal segment, if
short and narrow, is never protruding. The body is provided with
cerores none of which are octacerores or pilacerores and is never pro-
vided with ceratubae. The body is never enclosed in a resinous cell with
three adjacent openings or covered by a scale with or without exuviae.
The insects either live in galls or live free. The body may be wholly
or in part covered by a compact mass of wax with mealy or powdery
wax. The rectum is never provided with a long glassy tube of wax or
with a chitinized rectal tube with rings of anacerores. The body is
never naked and gall-like in form.
205
206 THE COCCIDAE
The female nymphs of all stages are provided with a rostrum and a
rostralis. The three pairs of legs are present in the first nymphal stage
and in the second nymphal stage so far as known and are similar or
subsimilar in form. The profemora are never greatly enlarged and the
prothoracic legs are never fitted for digging. The body is never provided
with an anal cleft and opercula, an anal ring and anal ring setae, octa-
cerores, pilacerores, or ceratuhae. The anal lobes are generally small,
but even if not prominent, frequently with prominent anal setae in the
first nymphal stage. The caudal end of the rectum is not strongly chiti-
nized, forming a rectal tube with one or more rings of anacerores ex-
creting a long glassy tube of wax. The caudal abdominal segment is not
short, narrow, and projecting.
The male has ocellanae. The abdomen is not provided with lateral
filaments or with tufts of long slender glassy threads of wax, but with
two long white caudal filaments. The wings are present. The stylus is
about as long as broad.
The subdivision Idiococcidae was proposed in 1893 by Maskell
to include two genera, Cylindrococcus and Sphaerococcus, which he
had described. The group was defined to include the following:
*' Adult females active or stationary; gall-making, or naked, or
producing cotton or wax. Anal tubercles entirely absent. Anal
ring hairless. Antennae with usually less than seven joints. Body
not prolonged posteriorly." When this group was erected, the
first genus contained two species and the second genus five. In
Fernald's Catalogue the first of these genera, Cylindrococcus, con-
tains four species and the other, Sphaerococcus, nineteen. Maskell
in 1895 in a list of the Australian coccids gives the Idiococcidae as
a section of the subfamily Coccinae, the Eriococcinae of this work,
and associates it with the genera included in the Eriococcinae as
given on preceeding pages. Cockerell in his "Check-list" pub-
lished in 1896 adopts Maskell 's name and recognized it as a group,
but combined it with the Eriococcinae in 1899. Green in the first
volume of his Coccidae of Ceylon also adopts Maskell 's name and
characterization with this addition: "Larvae with anal tubercles,
adult without." The term Idiococcidae is not tenable or usable
as a group name, as a subfamily or family name, because it is not
based upon a generic name. For this reason the subfamily name
Cylindrococcinae has been used for this group.
At the time when Maskell characterized his Idiococcidae, the
tendency was to rely upon the characters of the adult female alone
and not to use, at least to any extent, the characters of the nymphs.
Several of the genera now placed in the Eriococcinae, would ac-
cording to Maskell 's definition, fall into the Cylindrococcinae,
since their adult females do not have an anal ring or anal ring
i
SUBFAMILY CYLINDROCOCCINAE 207
setae. These structures are present, however, in the first nymphal
stage and frequently in the other nymphal stages of the leniale.
Maskell figures for the second nymphal stage of the female of the
type species, casuarinae Maskell, of the genus Sphaerococcus, an
anal ring with two anal ring setae. This genus for this reason
should be removed to the subfamily Eriococcinae. It does not
follow that the remaining eighteen species of this genus are
eriococcids. Two of these have already been removed as the types
of new genera, one being referred to the Efiococcinae and the
other to the Asterolecaniinae. Several of the other species repre-
sent another generic type, particularly the species forming galls
from the buds of plants.
The females of the generalized Anal Eing Conservers have a
well developed anal ring with anal ring setae, not only in the
adult, but in all nymphal stages of the female. Associated with
the anal ring is the presence of anal lobes and anal setae. The
anal lobes, when the abdomen is swollen with eggs, become incon-
spicuous. One of the lines of specialization in the eriococcids is
the gradual suppression of the anal lobes in the adult female fol-
lowed by an obsolescence of the anal setae. These structures are
both very prominent in the second nymphal stage of the generalized
genera, but with specialization are crowded out of this stage into
the first nymphal stage. The anal ring and the anal ring setae
pass through a similar series of stages of obsolescense. First there
is a reduction in the size of the anal ring setae in the adult female
followed by a reduction in the number until they are all wanting
and the adult female lacks not only the anal ring but the anal ring
setae. The second nymphal stage passes through a similar series
of changes and reductions. This results in the inclusion in the
subfamily Eriococcinae of all those genera whose species are pro-
vided with at least some anal ring setae in the first nymphal stage.
There is considerable question as to whether the Cylindrococ-
cinae are a natural group or not. Their structure would suggest
that they were highly specialized eriococcids that had gone one step
farther. That is they lack an anal ring and anal ring setae not
only in the adult female and its second nymphal stage, but also
in the first nymphal stage. They usually lack anal lobes and anal
setae in all stages or, if present, are greatly reduced in size. If
this interpretation is correct, the Cylindrococcinae is not a natural
subdivision but simply the highly specialized end of the Eriococ-
cinae that has lost through specialization by reduction the charac-
208 THE COCCIDAE
ters used in separating this group from the Cylindrococcinae. The
following quotation would suggest that Maskell held a similar
view. ''Moreover, I propose to characterize the Idiococcidae by
such wide and comprehensive features as will permit the future
inclusion therein of other genera which may hereafter be discov-
ered ; in fact, I mean, the subdivision to serve as a receptacle for,
perhaps many insects which cannot possibly enter into the others,
and so we may avoid, so far as possible, multiplication of names."
The species of one genus of this subfamily form galls upon
plants. A considerable number of these are from the Australian
region. While the galls of the Apiomorphinae are formed upon the
tissue of the stems and leaves of the plants, those of the gall mak-
ing species of the Cylindrococcinae are in most cases modifica-
tions of the buds. The galls resulting from such parts of the
plant, are, as in most of the galls formed in similar situations by
other insects, miniature cones. The species of a few genera live
in blister-like swellings upon the bark or leaves. These swellings
are abnormal growths upon the plant and should probably be con-
sidered as galls. The galls made by the species of Cylindrococcus
are usually comparatively small, never so voluminous as in the
female galls of many apiomorphids.
The adult females of a number of the species are enclosed in
a test of wax which completely encloses them and frequently has
an opening at one end or on one side. The wax of the test, when
a definite one is formed, is not voluminous, but is hard and not
easily soluble in caustic potash. The test in two genera is shaped
like a Greecian lamp. In the case of Ourococcus the insect lives
in a crevice in the bark, excretes a black covering of wax and a
long glassy tail of wax. The three species of Capulina are sug-
gestive of the conditions found in this subfamily. The body of the
adult female of sallei, the type, is covered with a white cottony
sac of wax with a single long caudal tube of wax, jaboticabae makes
neither gall nor definite sac and desposits her eggs in a fluffy mass
of white cottony wax, while crateriformis makes a small crater-
shaped gall or depression.
There is great variation in the external shape of the adult
female. The body may be elongate with parallel sides, cylindrical
or subcylindrical in form, or may be orbicular in outline, like a
thickened pancake with the peripheral margin convex. The mouth-
parts, including the rostralis and rostrum, are always present.
The head and prothorax form a single region in the gall forming
SUBFAMILY CYLINDROCOCCINAE 209
species and this region is distinctly separated from the meso-
thorax. The segmentation of the thorax and abdomen is usually-
distinct in these species, but in the wax forming species it is
generally completely effaced, frequently indicated only by the
marginal setae. In many coccids there is a definite number of
setae or cerores or groups of cerores along each lateral half of the
body. This is illustrated by the cerari found in many eriococcids.
The number of setae or cerores varies somewhat with the species,
usually the number is about eighteen, of which the first four of
each side belong to the head, two to each thoracic segment, and
one to each abdominal segment. The number may be slightly
increased or decreased, or the number may be doubled, twenty-
eight on each side. While the antennae and legs and all indication
of the intersegmental coriae may be wanting, the extent of the
various segments of the body can be closely approximated by a
study of the number and arrangement of the marginal setae or
other marginal structures that may be present.
The antennae of the adult female, even when present, is greatly
reduced and consists of only a few segments, always less than
seven, according to Maskell. The legs like the antennae are
greatly modified. They are frequently wanting and when all are
present, one pair is much larger than the others. The thoracic
spiracles although not described or figured for all the species are
undoubtedly present and located on the ventral aspect. The
abdomen is elongate in the gall forming species but is very short
in some of the species that are covered with wax. Abdominal
spiracles are not recorded as present in any of the species and the
anal ring and anal ring setae are also wanting.
The nymphs of the first stage, where known, are elongate,
ovate in outline and resemble very much the first stage nymphs
of the eriococcids. The segmentation is indicated, the ocellanae
are present, the antennae are long and slender and consist of six
segments, and the legs are long and typical in form. The caudal
end of the body may bear anal lobes, which are distinct or may be
wanting. The anal setae are also frequently present. The dorsal
aspect of the body may bear six longitudinal rows of setae, known
as dorsal setae, or transverse rows of ten setae each or the setae
may be very small and not arranged in rows or they may be
wanting. The anal ring so far as indicated is always wanting. It
needs more careful study than has been given it thus far in this
group.
210 THE COCCIDAE
The males in the wax forming species excrete an elongate
white sack of loose cottony filaments of wax or a small cylindrical
horny ferruginous sac of wax in which transformation takes place.
The males of three genera have been described, Capulina, Car-
pochloroides, and Halimicoccus. The descriptions are incomplete,
but there is nothing to suggest that they differ from the males of
the associated subfamilies. The genera referred to the subfamily
Cylindrococcinae can be separated by means of the following
table : —
GENERA OF CYLINDROCOCCINAE
a. Thorax of adult female with at least one pair of complete legs
present.
b. Thorax of adult female with three pairs of legs, prothoracic and
mesothoracic pairs small and rudimentary, metathoracic pair large
and prominent; nymphs of first stage without rows of dorsal
conical setae. Sphaerococcopsis Ckll.
bb. Thorax of adult female with only single pair of legs present;
nymphs of first stage with rows of dorsal conical setae.
c. Thorax of adult female with metathoracic legs present and
prothoracic and mesothoracic legs wanting; nymphs of first
stage with two complete rows of dorsal setae; females either
not forming galls or living in crater-like depressions
Capulina Sign.
cc. Thorax of adult female with prothoracic legs present and
mesothoracic and metathoracic legs wanting; nymphs of first
stage with four rows of dorsal setae; females forming galls
on plants. Cylindrococcus Mask.
aa. Thorax of the adult female with all three pairs of legs wanting.
b. Abdomen of adult female emarginate ati caudal end with a pair of
anal lobe-like projections, anus located at bottom of emargination
from which projects long glassy tube of wax. Ourococcus Fuller.
bb. Abdomen of adult female not emarginate at caudal end and not
with anal lobe-like projections.
c. Nymphs of first stage with dorsal longitudinal rows of conical
setae.
d. Antennae of adult female tuberculate, bearing terminal
bunch of stiff setae; anal tubercles wanting but indicated by
groups of conical setae; body with numerous setae near
anus. Apiococcus Hempel.
dd. Antennae of adult female wanting; anal tubercles wanting
and not indicated by groups of setae; body not with numer-
ous setae near anus Carpochloroides Ckll.
cc. Nymphs of first stage not with dorsal longitudinal rows of
conical setae.
d. Body enclosed in horny sack of wax shaped like lampshell;
cuticle of adult female not tuberculate and without conspicu-
ous cerores; antennae and legs wanting Halimicoccus Ckll,
SUBFAMILY CYLINDROCOCCINAE 211
dd. Body not enclosed in horny sack of wax; cuticle of adult
female with numerous hyaline tubercles without conspicuous
cerores and without setae; antennae tuberculate and legs
wanting. Phoenicococcus Ckll.
Sphaerococopsis Ckll. — This genus includes a single species,
inflatipes Maskell from Australia on Eucalyptus. It is somewhat simi-
lar in general appearance to Pseudoripersia turgipes Maskell, but the
adult female lacks an anal ring and anal ring setae. The legs are also
different in size and form.
Capulinia Sign. — The three species referred to this genus are:
cra)teraformis Hempel from Brazil on Eugenia; jaboticabae von Ihering
from Brazil on Eugenia and Myrciaria; and sallei Signoret, the type, from
Mexico on "Capulino" and "Escobillo".
Cylindrococcus Mask. — An Australian genus of four species oc-
curring upon Casuarinia: amplior Mask., casuarinae Mask., gracilis
Fuller, and spiniferus Mask.
Ourococcus Fuller. — A genus from West Australia found upon
Eucalyptus and Casuarinia. The species are: casuariniae Fuller, cobbii
Fuller, and eucalypti Fuller. The immature stages are unknown. The
anal ring and anal ring setae are not described. The form of the caudal
end of the abdomen and the fact that in each species a caudal glassy
tube of wax is formed suggests that the anal ring and anal ring setae
are present and that this genus should be placed in the subfamily
Eriococcinae, falling in the table of genera near Antonia, or if without
an anal ring the rectum must be provided with rings of anacerores.
Apiococcus Hempel. — A Brazilian genus found upon Myrtaceae.
It contains the four following species: asperatus Hemp., globosus Hemp.,
gregarius Hemp., and singularis Hemp.
Carpochloroides Ckll. — This genus contains a single species,
viridis Ckll., which is found upon Eugenia and Myrtaceae in Brazil.
Halimococcus Ckll. — A single species, lampas Ckll., occurring
upon the leaves of palm in Natal. The abdomen is provided with four
long setae at the caudal end instead of two as in Phoenicococcus.
Phoenicococcus Ckll. — There is a single species, marlatti Ckll.,
which occurs upon date palm in Algeria. It has been identified on date
palm received at quarantine from Algeria.
CHAPTER XX
SUBFAMILY CONCHASPINAE
The Exuvialess Scales
The body of the adult female is depressed, hroadest across the pro-
thorax. The lateral margins gradually converge caudad, they are thin
and convex but not margined. The segments are well defined. The
caudal end of the body is not inflated and the cephalic and lateral mar-
gins are not provided with a keel. The antennae are prominent, consist
of three segments, and are articulated to the ventral aspect of the head
near each lateral margin. The eyes are present. There are two ocellanae
located on the dorsal aspect, nearer to the meson than to the lateral mar-
gin of the head. The mouth-parts, the rostrum and rostralis, are pres-
ent, the rostrum consists of two segments. The legs are present. They
are subequal in length and normal in form. The profemora are never
enlarged and the prothoracic legs are never fitted for digging. The
trochanto-femoral sutures are present and the tibio-tarsal sutures are
wanting. The spiracles of the mesothorax and metathorax are normal in
size and ventral in position. The mesothoracic spiracles are never lo-
cated near the anus. The abdominal spiracles are limited to a single
pair located on the ventral aspect of the first segment, their spiracular
tracheae are never provided with rings of cerores. The stigmatic clefts,
spinae, and canellae are wanting. The abdomen is about as long as the
head and thorax togeither, and is convex on the dorsal and ventral as-
pects, always more or less depressed. The lateral aspects are convex
but distinct, not distinctly margined, the caudal end is convex. All the
segments are fully exposed and subsimilar in appearance with distinct
coriae between them, none are retracted to form a marsupium. The
abdomen is never provided with an anal cleft and opercula, or anal
lobes and anal setae. An anal ring and anal ring setae may be present.
The caudal abdominal segment is never short and narrow and extending
beyond the other segments. The body is provided with cerores none
of which are octocerores or pilacerores and is never provided with
ceratubae. The body is never enclosed in a resinous cell with three
adjacent openings, but is always covered by a scale which does not
contain exuviae. The rectum is never provided with a long glassy tube
of wax or with a chitinized rectal tube with rings of anacerores. The
body is never naked and gall-like in form.
The female nymphs of all stages are provided with a rostrum
and rostralis. The three pairs of legs are present in all nymphal stages
and are similar in form. The profemora are never greatly enlarged and
the prothoracic legs are never fitted for digging. The body is never
provided with an anal cleft and opercula, anal lobes and anal setae, octa-
212
SUBFAMILY CONCHASPINAE 213
cerores, pilacerores, or ceratubae. An anal ring and anal ring setae
may be present. The caudal end of the rectum is not strongly chiti-
nized forming a rectal tube provided with one or more rings of ana-
cerores excreting a long glassy tube of wax. The caudal abdominal seg-
ment is not short, narrow, and projecting.
The male has four ocellanae, two dorsal and two ventral. The ab-
domen is not provided with long lateral filaments or tufts of long wax
filaments or with two long thread-like filaments. The stylus is long,
one-half or more the length of the body.
This subfamily contains three genera, Conchaspis of Cockerell,
Fagisuga of Lindinger, and Scutare of Brittin. The typical genus
was based upon specimens collected in the West Indies. A descrip-
tion of them was published in February, 1893, under the name of
Conchaspis angraeci. In July of the same year Newstead described
this same species as Pseudinglisia rodrigueziae from plants im-
ported from Trinidad. Four other species of the typical genus
are now known. The genera of Lindinger and Brittin each con-
tain only a single species.
The scale of the adult female of Conchaspis is circular in out-
line, elevated at the center and bluntly conical. The surface of
the scale is unmodified in socialis Green, while there are six to
eight ridges radiating from the apex of the cone but not reaching
the periphery of the scale in angraeci Cockerell. The ventral scale
is complete, free from the dorsal scale, and adheres to the host-
plant. The dorsal or outer scale is opaque to grayish brown or
black. The female scales resembles the convex light colored scales
of certain diaspids, but they can always be separated from these
scales, because in the Conchaspinae the molted skins or exuviae
are never fixed in the dorsal or ventral scale. It is evident that
the molted skins, which are probably not more than two, although
the number is unknown in either sex, after they are shed, are
pushed out under the margin of the scale as in the males of certain
lecaniids.
The scale of the male is known only in socialis Green. It is
described by Green as ''white, oblong, flattened, closely felted,
completely enveloping the pupa. The hinder extremity has a
valvular opening. Length 1.50 mm. and breadth at widest part
is about 0.75 mm. The male puparia do not occur separately, but
are always collected together, in groups of ten or more, beneath the
parent scale (that of the female) which they completely fill." It
is questionable whether this is the true scale of the male and
whether it is comparable to the scale of the female, but rather a
cocoon. May it not be that the male does not form a scale.
214 THE COCCIDAE
The body of the adult female as viewed from above resembles
somewhat in general appearance an eriococcid nymph of the first
stage, but the segmentation is indicated more distinctly and the
body is more depressed. The eyes are present, located on the
dorsal aspect, near together and near the cephalic portion of the
body. The antennae are small, consist of three or six segments,
and are articulated to the ventral aspect of the head, distant from
each other and from the rostrum, which consists of two segments.
The legs are long and project beyond the lateral margin of the
body, probably due in great part to their articulation to the
sternum very distant from each other. They usually lack the
suture between the tibia and tarsus. Each bears a few long
setae, but lacks digitules. The claws are bluntly toothed on the
ventral side at the proximal end. The mesothoracic spiracles are
located near to and laterad of the prothoracic legs while the
metathoracic spiracles are located a short distance caudad of the
mesothoracic legs. A third pair of spiracles, those of the first
abdominal segment, are located just caudad of the metathoracic
legs. The caudal abdominal segments have chitinized areas which
are considered by Newstead as similar to the pygidium of the
diaspidids. These plates bear large circular cerores which are dis-
tributed over the segments or are arranged in groups as in socialis
near each lateral margin of the ventral aspect of segments one to
three. There is in this species on the lateral margin of segments
one to four small groups of short broad tubular ducts which open
on the dorsal surface. The lateral margin of each segment of the
thorax and abdomen bears a group of three or four long slender
setae. The abdomen does not contain ceratubae or oraceratubae,
structures which are so characteristic of all diaspidids. The size of
the body of the adult female is remarkable, as Green states, in
socialis it is 1.25 mm. long and 0.80 mm. wide, while the scale has
a diameter of 5 mm. The female deposits her eggs which are long
and purplish in color under her scale where they are mixed with
the cocoons of the male.
The adult male is known only in socialis Green. The body is
yellowish in color. The head is rounded, bears four simple eyes
or ocellanae, two dorsal and two ventral, and two antennae, each
consisting of seven segments. The distal segment bears four
knobbed setae and the other segments bear numerous small conical
setae. The wings are large and held in place by prominent halteres.
The caudal end of the abdomen bears a slender stylus that is more
SUBFAMILY CONCHASPINAE 215
than one-half the length of the body. Green was unable to follow
the life history of the male, but found some nymphs in their cocoons
that were oval in outline, still retained their nymphal legs, and
bore antennae consisting of four segments.
Our knowledge of the development of the female is very lim-
ited. Newstead states that the nymphs of the first stage resemble
the adult female except that they are smaller. They are short
and ovate in outline and have antennae of six segments, of which
the two distal segments are the longest. The legs lack the tibio-
tarsal suture as in the adult female but differ in that there are
two tibial and two ungual digitules present. The abdominal seg-
ments are also provided with plates. The three genera of this
subfamily can be separated by means of the following table : —
GENERA OF CONCHASPINAE
a. Body of adult females not with broad multidenticulate projections;
dorsum of caudal end of abdomen not with thickened plate-like pro-
jection on each side of meson; setae of lateral margin of each seg-
ment long, in groups or two or more.
b. Abdomen with caudal end projecting, not with anal lobes bear-
ing setae; tergum of thorax with ventral aspect never with con-
tinuous marginal fringe of setae Conchaspis Ckll.
bb. Abdomen with caudal end deeply hollowed out, hollow bounded
on each side by a prominent anal lobe bearing one long slender
seta and three short stout setae; tergum of thorax with continu-
ous fringe of short setae Scutare Brittin.
aa. Body of adult female with three broad, multidenticulate, pectinae-
like projections; dorsum of caudal end of abdomen with thickened
plate-like projections on each side of meson; each plate with row
of three short setae; lateral margins of segments apparently with-
out setae; caudal end of abdomen truncate, bearing near margin six
ventral and four dorsal setae Fagisuga Lindinger.
Conchaspis Ckll. — This genus with its four described species is
placed beitween the Phenacoleachiinae and Dactylopiinae by Mrs.
Fernald in her Catalogue. Green and Newstead place it before or after
the Diaspidinae, a more logical location it would seem. The species of
this genus can be separated by means of the following table: —
SPECIES OF CONCHASPIS
a. Antennae of adult female with five or six segments, scale of adult
female conical but not fluted.
b. Antennae of adult female with six segments, segments subequal
in length; abdominal segments with four groups of cerores,
larger number on cephalic segments, with scattered cerores be-
tween groups. — Mexico on Plumieria newsteadii Ckll.
bb. Antennae of adult female with five segments, two distal seg-
216 THE COCCIDAB
menits obscure, segments variable in length, third twice as long
as fifth, distal segment with two thick compressed setae on distal
end, antennal formula 35142; abdomen broader than head and
thorax, cephalic partt of ventral aspect with two much com-
pressed thick setae; caudal abdominal segments with glandular
areas on dorsal and ventral aspects. — Brazil on a woody shrub— _
fluminiensis Hemp,
aa. Antennae of adult female with less than five segments.
b. Antennae of adult female with four segments; cerores found on
all abdominal segments and generally distributed; scale of adult
female fluted.
c. Body of adult female purplish red; scale conical with apex at
center. — England, West Indies, United States, Mexico on
Angraecum, Rodriguezia angraeci Ckll.
cc. Body of adult female colorless, except brown plates of abdo-
men; scale conical with apex on one side. — Mexico on Hibiscus.
angraeci MMsci Ckll.
bb. Antennae of adult female with three segments; cerores arranged
in group on each side of abdominal segments one ito three; scale
of adult female not fluted. — Ceylon. socialis Green.
Scutare Brittin. — Scale of female is flat, subcircular, glassy,
fringed, and ventral scale is complete. The adult female is widest
through the thorax. The antennae consist of six segments and the ros-
trum of two segments. The thoracic spiracles are represented by two
pairs. The legs are present and normal in form. The abdomen is dis-
tinctly segmented with distinct chitinized areas. The anal lobes are
distinct, subtriangular. On each side of each abdominal tergum within
the margin there are four shorifc cylindrical setae, these are continued
around the margin of the thorax. The tergum of the abdomen between
the anal lobes and fifth segment is provided with a row of similar setae
on each side of the meson. The anal ring located between the anal
lobes, bears six long fine setae. AduHt insect with covering scale does not
resemble other species of subfamily. The presence of an anal ring with
anal ring setae and a scale fringed with glassy threads of wax needs to
be confirmed. A single species, fimbriata Brittin, is the type.
Fagisuga Liindinger. — A single species, triloba Lindg., is the type.
It occurs in Chile on Nothofagus. The scale is about 1 mm. in diameter,
without exuviae, conical, not fluted, dirty white in color. The body is
0.69 mm. long and distinctly segmented. The antennae consist of four
segments. Its formula is 4312. The body on each side adjacent to the
lateral margins of the denticulate projections is provided with several
tooth-like projections.
CHAPTER XXI
SUBFA>nLY DIASPIDINAE
The Armored Scales
The body of the adult female is variable in form, frequently slightly
longer than broad, often distinctly broader than long, sometimes four
or more times as long as broad, when comparatively short, broadest
through the spiracular region, always more or less depressed. The lat-
eral margins, at least of the caudal part, always thin and distinct and
converging caudad. The caudal end of the body is not inflated and the
cephalic and lateral margins are not provided with a keel. The antennae
are usually wanting, sometimes each is represented by an unsegmented
tubercle located on the ventral aspect, bearing one or more se^tiae. The
eyes are wanting. The mouth-parts, rostrum and rostralis, are always
present, the rostrum consists of a single globular segment. The legs
are always wanting. The spiracles of the mesothorax and m^fcathorax
are always present, normal in form and size, ventral in position. The
mesothoracic spiracles are never located near the anus. The abdomi-
nal spiracles are always wanting. The stigmatic clefts, spinae, and
canellae are wanting. The abdomen is rarely if ever longer than the
head and thorax together, when the body is elongate, the elongation is
confined for the most part to the thorax. The dorsal and ventral sur-
faces near together, particularly on the caudal half, where the body is
distinctly depressed. The lateral margins sharply defined and convex.
The caudal end is bluntly rounded, bluntly pointed, or rarely truncate.
All the segments are fully exposed or at least no part is retracted to
form a marsupium. The abdomen is never provided with an anal cleft
and opercula, an anal ring and anal ring setae, or anal lobes and anal
setae. The abdomen is divided into two regions, a cephalic segmented
preabdomen and an unsegmented caudal pygidium upon which the caudal
segment can not be identified. The body is provided with only a few
cerores, limited, when present, to the spiracerores and the genacerores.
The body is never provided with pilacerores or octacerores. The cera-
tubae are always present. The body is never enclosed in a resinous
cell with three adjacent openings. It is always covered by a scale with
one or two exuviae. The rectum is never provided with a chitinized
rectal tube bearing rings of anacerores excreting a long glassy tube of
wax. The body is never naked and gall-like in form.
The female nymphs of all stages are provided with a rostrum and
a rostralis. The three pairs of legs are present in the first nymphal
stage and wanting in all others, those of the first nymphal stage are
normal in form and structure. The profemora are never greatly en-
larged and the prothoracic legs are never fitted for digging. The body
is never provided with an anal cleft and opercula, an anal ring and
217
218 THE COCCIDAE
anal ring setae, pilacerores, or octacerores. The anal lobes in the first
nj^mphal stage are indicated with varying degrees of distinctness and the
anal setae are rarely wanting. The ceratuhae, although usually few in
number, can generally be recognized. The caudal end of ithe rectum is
not strongly chitinized with one or more rings of anacerores excreting a
long glassy tube of wax. The caudal abdominal segment can not be
identified.
The male has six ocellanae, two dorsal, two sublateral, and two
ventral. The abdomen is not provided with long lateral filaments of
wax or two thread-like caudal filaments of wax. The wings are some-
times wanting. The stylus is long and slender.
The eggs, in the case of those species laying eggs, are deposited
under the scale behind the body ofi^e female. They are elongate
in form with the two ends rounded and are whitish, yellowish, or
reddish in color. The surface of the eggs is frequently more or less
dusted with fine granules of wax, excreted by the genacerores.
The young nymphs when they hatch remain for a time under
the scale of the parent. All the eggs on the same branch or tree
apparently hatch at about the same time and the young first stage
nymphs swarm out from under the various scales, so that the
smaller branches of the host-plant, where the infestation is heavy,
are so densely covered with the young mite-like coccids that the
branches appear grayish in color. The young insects after wan-
dering about on the branch or branches for a few hours insert
their rostralis and attach themselves to the host-plant and begin
to draw nourishment.
The first stage nymphs, in which the males are indistinguish-
able from the females, are normally provided with antennae of six
segments. The distal segment is long and constricted and appears
as if composed of several fused segments. There is a single
ocellana on each side near the articulation of an antenna. The
rostrum, consisting of a single globular segment, and the rostralis
are easily identified. The loop of the rostralis extends into a
prominent crumena which projects into the cavity of the abdomen.
The legs consist of the normal number of segments and are
articulated near together distant from the lateral margin of the
body. They are long enough, however, so that they project beyond
the margin of the body. The tarsi usually bear two tarsal and two
ungual digitules. The body, which is more or less depressed and
oval in outline, has a fairly distinct segmentation of the thorax
and abdomen. This latter, which apparently consists of six seg-
ments, is in reality composed of two parts, the cephalic five seg-
ments constitute the preabdomen and the apparent sixth segment,
SUBFAMILY DIASPIDINAE 219
which is formed by the fusion of four others, is known as the
pygidium. The abdomen, therefore, consists of nine segments.
The five preabdominal segments are distinct in Lepidosaphes, but
in Chionaspis and some other genera, the fifth preabdominal
segment is combined with the pygidium, so that in such genera
there are apparently four segments in the preabdomen and five in
the pygidium. The margin of the pygidium and of the abdominal
segments usually bears projections, but they are quite different in
form from those found in the adult and described later. The
caudal portion on each side of the meson bears a large prominent
seta, an anal seta, which is one-half or more the length of the
body. The first stage nymphs of certain genera show structures
that may be anal lobes, but they are never distinct. The anal
ring and anal ring setae are never present. The anus, a circular
opening located on the dorsal aspect of the pygidium, is always
distinct.
The first nymphal stage is a comparatively short one. There
is considerable variation among the different genera in the way
in which the cuticle is molted at this time. In the genera Diaspis,
Lepidosaphes, and their allies the old cuticle is ruptured trans-
versely between the points of attachment of the antennae and the
rostrum. The cuticle is then ruptured along each lateral margin
to the pygidium and the ventral portion of the cast skin or exuvia
is rolled caudad and comes to rest upon the pygidium. This
explains why the mouth-parts are located over the pygidium in
certain mounted individuals, a condition that nonpluses many
beginners. This method of dividing the exuvia separates the
antennae which remain attached to the cephalic portion of the
dorsal exuvia and project horn-like beyond its cephalic margin,
even in old dried exuviae, while the caudal portion of the ventral
exuvia is folded upon the pygidium. In the genus Aspidiotus and
its allies the split extends entirely around the margin of the body
and the entire dorsal aspect is separated from the ventral, forming
two distinct plates, a dorsal exuvia and a ventral exuvia. The
dorsal plate in this case always lacks the dried porrect antennae.
The dorsal exuvia forms a part of the dorsal scale and the ventral
exuvia forms a part of the ventral scale, but is frequently lost.
The female nymphs at the first molt lose their antennae and
legs, become greatly enlarged in size, and are more like the adult
in general form. This is also true of the structures on the margin
of the pygidium. The second stage females may live for a con-
220 THE COCCIDAE
siderable time, several months, the length of this stage varying
greatly with the species. When the insects of this stage molt,
there is no variation in the way in which the cuticle ruptures, but
in all species there is a transverse rupture on the ventral aspect
cephalad of the rostrum which connects on each side with the
lateral margin, the further rupture of the cuticle extends caudad
along each side to the pygidium. The ventral portion of the
exuvia is rolled back upon the caudal portion of the ventral aspect
of the body.
The females at the completion of the second nymphal stage
and after the second molt assume the adult condition. The most
striking variation found in the adult female from that of the
second nymphal stage is the great increase in size and the greater
complexity in the structure of the caudal portion of the abdomen.
The male of the second nymphal stage is similar to the same
stage of the female. It lacks both legs and antennae, but possesses
a rostrum and rostralis. At the second molt all similarity between
the male and female is lost and the males of the third nymphal
stage are elongated. The rostrum and rostralis are wanting and
the developing legs and antennae have become of such size that they
show externally, each enclosed in a cuticular sac. The male
nymphs during this stage because of their similarity to the pupae
of insects with a complete metamorphosis are often incorrectly
designated as pupae and those of the second stage as propupae.
The third nymphal stage requires only a few days for its comple-
tion and after the final or third molt the insect transforms to an
adult.
The adult males, after completing their nymphal development,
push the exuvia of the last molt out under the margin of the scale
and the fully formed insect normally remains for several days
under the scale. This is evidently for the further completion of
the development of the adult organs. The wings are fully ex-
panded during this time.
The completely formed male is very different in appearance
from the adult female. The constrictions separating the segments
of the body are distinct. The eyes are simple and six in number,
four dorsal ocellanae, two of which are large and subequal in size
to the two ventral ocellanae. The antennae are typical in form
and consist normally of ten segments. The thorax is characteristic
in form and bears the nine-segmented abdomen. The stylus is
long and pointed, often one-half or more the length of the abdomen.
SUBFAMILY DIASPIDINAE 221
It is a valvular structure through which the flexible penis can be
extruded. The wings, which are normally present and prominent,
are sometimes wanting or greatly reduced in size in certain indi-
viduals.
The adult females vary greatly in form, size, and structure
in the different genera. The body may be elongate oblong, two or
three times as long as wide, linear, five or more times as long as
wide, oval or circular, varying from slightly longer than broad to
broader than long.
The constriction between the head and prothorax in the female
is rarely if ever marked while those bounding the other thoracic
segments may be distinct, obscure, or wanting. The antennae are
at most represented only by unsegmented tubercles bearing a
single large seta and in some species a few small setae. These
tubercles are articulated to the cephalic part of the ventral aspect
near to each other and to the meson. The ocellanae are always
wanting in the second nymphal and adult female. The mouth-
parts are well developed. The rostrum consists of a single segment
which is globular in form. The rostralis is long and forms a
distinct rostral loop within the crumena except in individuals
where the rostralis has been pulled out of the crumena in removing
the specimen from the host-plant. The rostrum, rostralis, and
crumena are always associated with a well developed endoskeleton.
The thoracic spiracles are always distinct and the spiracular
trachea connecting with each is modified into a distinct hour-glass
or trumpet-shaped cavaera. The mesothoracic spiracles are located
on each side of the attachment of the rostrum, while the meta-
thoracic spiracles are frequently located farther apart and about
as far caudad as the two mesothoracic spiracles are distant from
each other. This distance will vary slightly not only with the
species but with the different individuals of the same species.
When there are indentations in the lateral margin marking the
constrictions between the thoracic segments, the spiracles are gen-
erally located in line with these constrictions. It is doubtful if
these indentations represent the rudiments of the stigmatal clefts
and canellae, but in many species there are often a few spiracerores
associated with the spiracles, more frequently with the mesa-
thoracic than the metathoracic.
The great majority of the structures used in distinguishing
and classifying the various species of Diaspidinae are located upon
the abdomen. Morphologists are fairly well agreed that the typical
222 THE COCCIDAE
number of abdominal segments present in insects in general is
eleven. This number is readily recognized in many generalized
species of insects, but a study of the abdomen of several coccids
leads to the conclusion that the typical number of abdominal seg-
ments for this group is nine. That there has been a suppression
of the number of abdominal segments in other groups of insects is
easily proven. This suppression of segments may take place at
the caudal end, due to the modifications necessary to accommodate
the organs for copulation and for depositing the eggs, and at the
cephalic end, due apparently to the enlargement of the thorax to
accommodate the muscles for the organs of flight and its consequent
encroachment upon the cephalic abdominal segments. The ex-
planation of just how this reduction has taken place should be
sought, not only among the early nymphal stages of the female
coccids but especially among the groups showing less departure
from the progenitors of the coccids, the plant-lice and the psyllids.
The abdomen in all diaspidids is divided into two regions, a ce-
phalic segmented region, the preabdomen, and a caudal unsegment-
ed region, the pygidium. The segmentation of the preabdomen is
always more distinct on the dorsal than on the ventral aspect.
The limits of the segments are generally indicated on each lateral
aspect by distinct constrictions. The preabdomen frequently
consists of four segments in the adult female, but in the first and
second nymphal stages and the adult females of certain genera,
one and sometimes more segments form a part of the preabdomen,
so that it may consist of ^ve or six segments.
The caudal, strongly chitinized, unsegmented pygidium in
the adult female consists typically of four segments. The pygidium
is strongly depressed so that the dorsal and ventral aspects are
close together. The lateral margins of the pygidium are provided
with projections and indentations which are known collectively as
the pygidial fringe. It is also designated simply as the fringe but
this term has already been used for the marginal band of glassy
threads of wax excreted from the octacerores in the Asterolecaniinae
and should be reserved for this use. The structure generally
designated as the pygidium is also known as the last segment or
anal plate. The early students of the Coccidae believed this
strongly chitinized caudal end of the abdomen represented a sing]
segment and hence the name of pygidium, but it is in fact a com-
plex of four or five segments. The number of segments and their
limits can only be approximated because all the coriae or sutures
SUBFAMILY DIASPIDINAE 223
bounding them are obsolete. The females of the conehaspids have
been described as having a pygidium, but between all of their
abdominal segments distinct coriae can be identified. While the
pygidium of the Diaspidinae has probably been derived from a
progenitor with an abdominal structure similar to that of the
Conchaspinae, yet the form of abdomen found in this latter group
can not properly be designated as a pygidium.
The wax in the great majority of the Diaspidinae is poured out
through openings which are located in the external cuticle. They
are the external outlets of cuticular tubes. These tubes vary
greatly not only in length and diameter but in the size and shape of
their external openings. Each of these tubes is known as a cera-
tuba. It is not known just how the ceratubae function, whether
they serve simply as passage ways for the wax or whether they
serve as reservoirs into which the wax, as it is elaborated, is stored
until the insect is ready to use it. The ceratubae are formed by an
invagination of the body-wall. The inner end is truncate and
always bears on its ental surface a minute knob or nipple, the bulla.
Whether the ceratubae and the cuticular wax tubes found in the
Eriococcinae, Kermesiinae, Lecaniinae, and Asterolecaniinae have
a common origin is not known, but the presence of the bulla in the
ceratubae readily distinguish those of the Diaspidinae from the
wax tubes of the other subfamilies. There is a lumen extending
through the bulla which connects with the lumen of the ceratuba
and it is through this lumen the wax is poured into the ceratuba.
The ceratubae and their external openings are also known as fusi,
filiere isolate, filiere separee, filieres isolee, tubuli, tubular spin-
nerets, dorsal tubular spinnerets, orifici delle grosse filiere margin-
ale, fuse marginales, elongated pores, oval pores, marginal pores,
spinnerets, dorsal glands, dorsal gland orifices, wax ducts, elongated
pores, marginal gland openings, semi-lunar marginal pores, mar-
ginal lunar pores, or dorsal pores.
There are several types of ceratubae in the Diaspidinae and
each type seems to be more or less characteristic of certain groups
of genera. The external opening of each ceratuba, located in the
external cuticle, is an oraceratuba. This opening, which is usually
small, is the external opening of the ceratuba and not a ceroris or
wax pore or pygidial wax pore as it has been generally considered
and described. The ceroris of the ceratuba, if present, would be
located at the opposite or inner end adjacent to the bulla. The
great majority of the oraceratubae are located on the dorsal aspect.
224 ^ THE COCCIDAE
They are also known as macropores and micropores. In the genus
Chionaspis and its allies, the number of oraceratubae on each side
of the meson is sometimes designated by the use of a formula in
the same way that the number of plates are sometimes indicated.
There are at least six types of ceratubae.
The altaceratubae are found in the Lepidosaphini, Diaspidini,
Parlatoriini, and Fioriini. They are the large broad cylinders,
the mouths of which are usually oblique and located at or near
the margin of the pygidium.
The brevaceratubae are of the same general form as the alta-
ceratubae, but they never open at the margin of the pygidium.
Their oraceratubae are never oblique and they are always much
shorter and smaller in diameter than the altaceratubae. They are
usually about as broad as long and their oraceratubae are generally
known as dorsal pores or dorsal tubular spinnerets.
There is of common occurrence in the pygidia of the Aspidio-
tini long slender ceratubae in which the two sides are parallel or
nearly so. They are sometimes fifteen or twenty times as long as
broad. These are the linaceratubae, the wax ducts of Comstock
and others, and their oraceratubae are sometimes designated as
macropores.
In the pygidia of the Aspidiotini, there is in addition to the
linaceratubae, other ceratubae which are as long or longer than
the linaceratubae, but in which the greater part of the tube is
reduced to a fine thread. The inner truncated end of these cera-
tubae which bears the bulla is much larger. This enlarged inner
end gives these ceratubae a distinct club-shaped or clavate appear-
ance and they are consequently known as the clavaceratubae.
In addition to the linaceratubae and clavaceratubae, there are
other long slender ceratubae in the pygidia of most Diaspidinae.
These are the lamaceratubae. They differ from all other ceratubae
in that their oraceratubae are always located in the distal end of a
plate or pectina, projections of the pygidial fringe.
The lateral portions of the segments of the preabdomen bear
projections which are known as bract eae. They are not only
similar in general form to plates or pectinae but are undoubtedly
their precursors and bear oraceratubae at their free end. The
ceratubae opening through the bracteae are paraceratubae. They
are, so far as observed, always short slender tubes that are ordi-
narily so slender and delicate as to be identified only with great
difiiculty.
SUBFAMILY DIASPIDINAE 225
To Berlese belongs the credit for determining and describ-
ing the structure of the cells forming the threads of wax in the
Diaspidinae. These cells are modified hypodermal cells but are
very different from the flask-shaped cells connected with the
spiracerores and genacerores which excrete powdery or pulveru-
lent wax. The thread excreting cells are long and club-shaped,
each connected with the ental surface of the ental or free trun-
cated end of a ceratuba. Each ceratuba bears a group of three
cells, known as a diaspicera, one of which is supposed to excrete
the wax and the other two a covering for the wax. The wax cell,
the silk gland or ghiandola sericipara of Berlese, is attached to
the end of the bulla. The other cells, the ganogene cells, the
ghiandole ganogene of Berlese, are also attached, one on each side
of the wax cell, to the bulla. There are openings in the bulla
through which the excretion of these cells is poured. The struc-
ture of the wax cell is anomalous, according to the account and
figures of Berlese they are more like a multicellular gland in
structure than a single cell. The distal portion of this cell or
the secreting portion is flask-shaped and contains a single large
nucleus. The secreting portion is connected by a greatly elon-
gated neck-like or tubular portion through which the excretion is
passed to the bulla and ceratuba. The tubular portion of the
cell adjacent to the bulla is thickened and striated and apparent-
ly functions for compressing the thread of wax in somewhat the
same way that the press of the silk duct in lepidopterous larvae
compresses the two components of each thread. The ganogene
cells are about one-fourth the length of the wax cell. Each con-
sists of a distal, club-shaped, excreting portion which contains a
single nucleus. There is a short tubular portion connecting the
excreting portion with its opening in the bulla. Berlese believed
that the function of the ganogene cells was to excrete a substance
which was used for coating or covering the exterior of the threads
of wax. If this suggestion is correct, the threads must be fully
formed when they make their exit from the bulla into the cera-
tuba and this latter structure can serve only as a reservoir for
storing the threads. The attachment of the cells to the ental
surface of the ceratuba would suggest that the bulla and associ-
ated parts might be a ceroris that had been invaginated. These
structures were figured by Childs, but no discussion of his con-
clusions is given.
226 THE COCCIDAE
The portion of each lateral margin of the pygidium extend-
ing on each side from the coria forming the cephalic limit of the
pygidium to its cephalic lobe, second, third, fourth, or fifth as
the case may be, is known as a lateris. It is also known as the
pygidial margin, lateral margin, thickened lateral margin, margo
lateralis pygidii, or simply as the margin. The lateres should
not be confused with the pygidial fringe, of which it is the ce-
phalic part.
Each lateris frequently contains indentations bounded by pro-
jections which are lobe-like in form. These projections, the
latadentes, bounding the indentations in the lateres have been
designated as the lateral teeth by Maria tt. The term latadentes
is applied to all the tooth-like projections cephalad of the cephal-
ic pair of lobes. In certain genera, as Diaspis, Chionaspis,
Lepiodosaphes and their allies, all of a considerable part of the
latadentes bear the oraceratubae of altaceratubae. Each of these
together with the tooth bearing it has been called a gland bear-
ing prominence by Cooley and others. The margin of each lat-
eris, whether the latadentes are present or not, may be finely or
coarsely dentate or denticulate. The denticulations bear no re-
lation to the latadentes. In those tribes provided with altacera-
tubae, the oraceratubae of those located between the lobes some-
times open in tooth-like projections. These are also known as
latadentes.
The margin of the pygidium generally contains a number of
distinct indentations or notches, the incisurae. They were so named
by Leonardi, but are also known as echancrures, incisions, in-
cised notches, interlobular incisions, or simply as notches. It is be-
lieved that the incisurae mark the points on each lateral margin
where the coriae reached the margin and, therefore, limit the seg-
ments. In those species where the pygidial fringe is well devel-
oped, the full number of incisurae, five, can usually be identified,
but they are greatly reduced or wanting where the pygidial
fringe has been wholly or in great part eliminated. This is par-
ticularly true where the lateres are long and well developed as in
most species of Aspidiotini. The pygidium of those species
where the adult female does not escape from the last nymphal
exuviae, as in the species of Aonidia, the incisurae are obsolete.
They are named, beginning with the one on the meson between
the median pair of lobes as the median or first incisura, the next
one on each side cephalad of the median incisura and the median
SUBFAMILY DIASPIDINAE 227
pair of lobes as the second incisura, and the next on each side
as the third incisura, the fourth, and the fifth. There is some dis-
erepancy as to the way in which the incisurae are numbered.
Most writers designate the median incisura as the median or
mesal and then number the others. A few writers number all the
incisurae and such writers always begin with the median. Most
of those who designate the first as the median do not designate
the next, the one between the median and second lobe of each side
as the second but as the first, the one between a second and third
lobe as the second, and the one between a third and fourth lobe
as the third, and so on. The median incisura is disregarded in
numbering the others. This method has not been followed.
There are distinct thickenings associated with the incisurae
in certain species. These thickenings are located on the ventral
aspect and may be known as the densariae. A furrow extends
cephalad from the bottom of one or more of the incisurae onto the
ventral surface of the pygidium and one or both sides of each of
these furrows may be thickened. The thickenings are usually
club-shaped in outline with the clavate portion at the cephalic end.
The number of incisurae with densariae is usually less than the
total number of incisurae present. There are frequently more
pairs of densariae present than there are pairs of lobes. The
densariae are only rarely associated with the median incisura.
Comstock, who was the first to make use of the densariae in tax-
anomic work, called them incisions with thickened edges. They
were designated as paraphyses by Leonardi and Marlatt, but
these structures are confined to the dorsal aspect. The furrow
between the densariae, when there is one on each side of the in-
cisura, is frequently provided with a row of prominent oracera-
tubae.
The broad or elongate semioval projections of the pygidial
fringe are the lobes. They are also known as truUae, pallae,
lobules, lamelles, or palette, and are also designated sometimes as
the primary lobes to distinguish them from the pseudolobes de-
scribed later. The lobes are typically arranged in pairs and sep-
arated by incisurae, although, through the absence of the mesal
incisura and the fusion of the mesal pair, this is sometimes appar-
ently untrue. The lobe on each side of the median incisura is
a median lobe, the two being distinguished as the median pair of
lobes. They are also known as the mesal or anal lobes. The lobes
cephalad of each median lobe are known collectively as the lateral
228 THE COCCIDAE
lobes. The lobe cephalad of each second incisura is a second lobe,
that cephalad of a third incisura is a third lobe, that cephalad of
each fourth incisura is a fourth lobe, and that cephalad of each
fifth incisura is a fifth lobe. The lobes cephalad of each median
lobe are sometimes numbered, beginning with the second lobe, as
the first lateral lobe, second lateral lobe, third lateral lobe, etc.
The lobes of the two sides are grouped together in pairs and des-
ignated as the second, third, fourth, and fifth pairs of lobes. The
maximum number of pairs of lobes is five, when there is more
than this number, the cephalic so called lobes are lobe-like lata-
dentes. The usual number of lobes is three, two, or a single pair.
The median lobes are generally the broadest and longest and each
successive pair is smaller and smaller until it becomes difficult to
distinguish the greatly reduced lobes from the latadentes. The
number of latadentes is greatly increased in certain species or
until such species are said to have ten or more pairs of lobes
present. A projection is not considered as a lobe in the follow-
ing descriptions unless it projects beyond the general outline of
the margin of the pygidium and has associated with it the other
structures of the pygidial fringe, plates and pectinae, which are
usually associated with lobes. The lobes when greatly reduced
in size frequently become narrower and in certain species are
long, blunt or pointed, spear-shaped projections. The lobes, even
when greatly reduced or plate-like in form, are never provided
with oraceratubae. The median lobes are sometimes adjacent on
the meson so that the median incisura is linear or they may be
fused for a part of their length so that it is represented as a
distinct notch or they may be completely fused forming a single
lobe and effacing the median incisura.
The lobes vary considerable in form. The simplest type is
where the margin of the lobe is without indentations and in such
cases is said to be entire. Where there is one or two indenta-
tions in the distal portion, the lobe is said to be notched. The
notch on the mesal side of the lobe is the median or mesal notch
and the one on the lateral side is the lateral notch. Either the
mesal or the lateral notch may be wanting. When there are a
few notches, more than two in the margin of the lobe, it is said
to be crenulate and when there are many notches to be serrate.
The notches in the crenulate or serrate lobes are always smaller
than in the ordinary notched lobes. A lobe may be divided into
two distinct parts by an incision, when it is said to be incised or
SUBFAMILY DIASPIDINAE 229
emarginate. Sometimes there may be two or more incisions so
that the lobe is divided into three or more parts. Each of these
subdivisions is described by some authors as a distinct lobe, they
are best known as lobelets and numbered in order beginning witli
the mesal one. The mesal lobelet, when there are only two
is generally larger than the lateral, if there are more than two,
the lateral lobelets are usually successively smaller than the
mesal.
The margin of the pygidium in those species that transform
in a puparium or do not escape from the last nymphal exuvia, as
the species of the genus Aonidia, may bear a few or a considerable
number of lobe-like projections. These lobes are frequently
asymmetrically arranged and are not homologous with the lobes
of those species not transforming in a puparium. Such projec-
tions are known as pseudolobes. The development of the species
transforming in puparia has been so greatly accelerated, that
true lobes are usually not found in the pygidium of the adult fe-
male but in the pygidium of the second nymphal stage and in
some species only in the pygidium of the first nymphal stage.
The pseudolobes are not only inconstant in different individuals
of the same species but may be different on the two sides of the
same individual. The exuviae of the second and first nymphal
stage of these species should be mounted for a study of the true
lobes and the other parts of the pygidial fringe.
The thin projections with toothed or dentate ends located in
the incisurae between the lobes and upon the lateres are known
as pectinae. This name was given these structures by Leonardi.
They are also known as squamae, squames, pols, squameaux,
petinii, scaly hairs, notched plates, serrate plates, serrated ducts,
serrulate plates, fimbriate plates, fringed plates, and furcate
plates. The pectinae are considered as extensions of the lateral
margin of the pygidium. They differ from the lobes in that there
is always an oraceratuba located in the distal end of each. The
pectinae are regarded as the primitive form of these projections
and not the plates as is held by those who consider the plates as
developed from setae and the pectinae as developed in turn from
the plates. While there is considerable variation in the number
and arrangement of the pectinae, their usual disposition is two
in the median incisura, two or three in each second incisura,
usually two, and two or three in each third incisura, usually three.
The pectinae also vary in form, but are readily arranged in four
230 THE COCCIDAE
groups; in the first group they are broad, the distal end is trun-
cate or subtruncate and the teeth are limited to the distal end.
These are the distapectinae and are of general occurrence in the
Parlatoriini. In the second group the shaft may be comparatively
broad or narrow, the distal end is pointed, so that each pectina
is subtriangular in outline and the teeth are arranged along both
sides of the shaft. These are the latapectinae. In the third
group the pectinae are similar in size and form to those of the
second group, but the teeth are limited to one side of the shaft.
These are the unapectinae. The fourth group includes those pec-
tinae in which there has been the greatest amount of reduction,
those with a slender shaft with two or three small inconspicuous
teeth at the distal end. These are the furcapectinae and are gen-
erally indicated as furcate plates by systematists. The latapecti-
nae, unapectinae, and furcapectinae are all found in the Aspidi-
otini.
The thin cuticular projections with pointed ends located in
the incisurae between the lobes are the plates. They are also
known as lamellae, gland spines, spine ducts, fusi piliformis, or
fusi spiniformis. The plates are considered as having been de-
veloped from pointed pectinae by the loss of their lateral pro-
jections or teeth. They are similar in number and position to the
pectinae and like them each bears an oraceratuba at its distal
end but it is generally more difficult to identify the oraceratuba
than the cylindrical part of the ceratuba extending through the
cavity of the plate. If the plates and pectinae are continuations
of the margin of the pygidium, each is typically provided with
a lumen which is a continuation of the lumen of the body. The
plates are frequently minute and often difficult to distinguish
•Prom setae, the spines of most authors. It is not strange, there-
fore, that they have been sometimes confused with setae, but they
can always be distinguished from setae by the absence of a calyx.
The number of plates and pectinae located in each of the
incisurae is of great value in the separation of species. Cooley
has suggested that for the sake of brevity the number of plates
be indicated in the form of a formula, but this scheme is just
as serviceable in indicating the number of pectinae. The for-
mula is written, as follows : 1-0, 1-2, 3-3, 1-2, 2-2, 3-7. The first
group of numbers shows the number of plates or pectinae in the
median incisura, the cypher indicating their absence, the second
group those of the second incisura, the third group those of the
SUBFAMILY DIASPIDINAE 231
third incisura, the fourth group those of the fourth incisura, the
fifth group those of the fifth incisura, and the sixth group ^ose
located on each lateris. Since the number of lobes and incisurae
is usually three or less, this formula would also indicate the
number of incisurae, for if there was only a single incisura pres-
ent, the formula would consist of only two groups, if of two in-
cisurae of three groups. Such a formula, 0-0, 1-3, 2-5, would
show the presence of two incisurae, the absence of plates or pec-
tinae in the median incisura, the presence of one to three in the
second, and the presence of two to five on the lateris. The formula
also suggests the number of lobes present.
In many of his descriptions Lindinger has abbreviated by
indicating many facts as to the pygidial fringe by the use of a
formula. This is well illustrated by the use of the one he gives
in ''Die Schildlause" for Chionaspis euonymi, as follows:
OPi; Li; 2P2; Dr-m; 2L2 ; 2P3 ; Dr-m; 2L3 ; 2?^ ; 2Dr-m; 2-3P5.
Translated, this formula means, no plates in median incisura,
median pair of lobes present, two plates and oraceratuba on a
latadentis in each second incisura, second pair of lobes present
and incised, two plates and an oraceratuba on a latadentis in each
third incisura, third pair of lobes present and incised, two plates
and two oraceratubae on latadentes in each fourth incisura, and
two to three plates on each lateris.
The arrangement of the plates on the lateres in the Lepido-
saphini, Parlatoriini, and Diaspidini is such in many species as
to suggest a segmental arrangement. The cephalic group, par-
ticularly in the Diaspidini, generally contains the largest num-
ber. Where these plates are separated so that each must be con-
sidered separately or where the plates of the other groups are sepa-
rated, those of a group are enclosed in a parenthesis. The
groups of plates on each lateris are separated by hyphens. The
following shows such a formula: (1-1) -2-2- (1-1-1-1).
The proximal portion of each of the lobes or the surface of
the marginal portion of the pygidium adjacent to the lobes and
the lateres adjacent to the latadentes usually bear small setae,
the pygidial setae. These setae are generally constant in num-
ber and position for a given species and are usually designated
as spines. Some of the setae are located on the dorsal aspect
and others on the ventral and for this reason they are some times
designated as the dorsal or ventral setae. There are a few
species in which a second row of setae, the submarginal setae,
232 THE COCCIDAE
have been identified on the ventral aspect a short distance cephalad
of the pygidial setae. The pygidial setae vary somewhat in size.
They are sometimes so large and so placed as to be mistaken
for plates, particularly in the median incisura when it lacks plates
or pectinae. But the insertion of each seta in a calyx should
remove any doubt as to its identity and structure. They are
also known as spines, polls, hairs, pili simplices, or peli semplici.
The dorsal and ventral setae appear in generalized species td be
segmen tally arranged.
In the region of the vulva there is located a varying number
of minute clear rings or spots. These rings are of quite general
occurrence in many species, but they are especially distinct in
most species of Lepidosaphini and Diaspidini, and, where ever
they occur, their presence is usually easily determined. There
are four to six located just cephalad of the vulva, which are sit-
uated on the ventral aspect; there is another pair located slightly
farther caudad which are situated on the dorsal aspect; there is
sometimes another pair located cephalo-laterad of each group of
pregenacerores which are situated on the dorsal aspect; there is
one or more pairs located between this last pair and the proximal
ends of the lobes which are situated on the ventral aspect; and
there may be a considerable number near the caudal margin of
the pygidium which may be situated on the dorsal or the ventral
aspect. All of these clear spots have been considered by some
writers and a part of them by other writers as micropores or the
openings of wax-pores, that is minute oraceratubae. The shaft of
a seta, although in most cases extremely minute, has been identi-
fied in every instance where these micropores have been care-
fully examined with an oil emersion objective. The rings are,
therefore, not wax-pores but the calices of setae. I was gratified
after this conclusion was reached to discover that Green figures
these structures as setae.
There are in certain species near the proximal ends of the
lobes distinct club-shaped thickenings, which are known as par-
aphyses. They are also known as thickenings of the body-wall,
club-shaped thickenings of the body-wall, processes, elongate
thickenings of the body-wall, or processi chitenosi. They vary
greatly in size and form even in the same species. Their num-
ber, size, and form afford excellent characters for the differenta-
tion of species. The paraphyses are thickenings of the cuticle
of the dorsal aspect as was demonstrated by Comstock and differ
SUBFAMILY DIASPIDINAE 233
in this respect from the densariae which are confined to the ven-
tral aspect of the pygidium and are never present, so far as ob-
served, in those species provided with paraphyses. These two
very different structures are confused by Leonardi and Marlatt
under the single name of paraphyses. The paraphyses are con-
tinuous, at least for the most part, with the proximal prolonga-
tions of the lobes while the densariae are always located between
the lobes. The number of paraphyses associated with each lobe
varies, one to three are the usual numbers, but there may be
more than three and some of the lobes may lack them, if want-
ing, it is usually from the mesal lobes. There is frequently a
considerable number present in a single pygidium, as many as
twenty-eight or fourteen pairs in some.
The anus in the Diaspidinae is the prominent opening located
on the meson of the dorsal aspect, frequently near the middle of
its length and usually some distance from the caudal end of the
pygidium. It is normally located in the caudal abdominal seg-
ment, but the depression of this part of the body, the close apposi-
tion of the dorsal and ventral aspects, and the consequent reduc-
tion in the lumen between these areas, the anus has migrated
cephalad and is found in the cephalic portion of the pygidium of
certain species. The anus is also known as the anal opening or
anal aperture.
The external opening of the female reproductive organs, the
vulva, is apparently constant in position in the Diaspidinae. It
is situated on the ventral aspect of the pygidium in the region
of what is believed to be the sixth segment. The position of the
vulva is often difficult to determine, but is generally more easily
identified because of the structures associated with it, the gena-
cerores, when these are present, and the fine radiating ridges in
the surface of the cuticle. The vulva is also known as the vagi-
nal opening, genital aperture, genital orifice, aperture sessuale
feminae, or genital otvor.
Figures of pygidia usually show the position of the anus and
vulva as if they were both on the same aspect. The comparative
position of the anus and vulva, their distance from each other
and from the caudal margin of the pygidium, is of considerable
value, but the fact that they are situated on different aspects
must not be overlooked. The variation in position between the
anus and vulva is due wholly or for the most part to the differ-
ence in position of the anus. In the Parlatoriini the anus is
234 THE COCCIDAE
usually located midway between the vulva and the caudal mar-
gin, but in Gymnaspis they are superimposed; in the Lepido-
saphini the anus is situated far cephalad of the vulva, the latter
is frequently located on a line drawn through the caudal margin
of the postgenacerores and the anus some distance cephalad of
the mesogenacerores ; in the Diaspidini the anus may be located
caudad of the vulva, superimposed, or cephalad, but is usually
placed near to or slightly cephalad of the vulva; in the Fioriniini
the anus is usually placed cephalad of the vulva, frequently abou»>
its own diameter, rarely slightly caudad; in the Aspidiotini the
anus is situated caudad of the vulva, the distance varying with
the genus and the species, in some it is located adjacent to the
proximal ends of the median pair of lobes, in others midway be-
tween the vulva and the caudal margin, or nearer to the vulva.
In measuring this distance, where reference is made to the caudal
margin, the distal ends of the median pair of lobes is always used
and not the bases or proximal ends of these lobes. In making
such measurements account must be taken of the fact that the
anus is not always round, the length may be two or three times
the width, in all such cases it is the transverse diameter or width
that is used.
The mesal portion of the caudal margin of the pygidium is
deeply emarginate in certain species. This is variously described
as emarginate, indented, notched, deeply concave, or semicircular-
ly or concavely hollowed out. This emargination involves more
than the median incisura, because it is frequently of such size
that the median lobes are situated for the most part within the
emargination. These lobes are usually strongly oblique, the lat-
eral margin of each is fused to the pygidium, and only a small
portion of the distal end of each median lobe projects beyond the
general contour of the pygidial margin. This emargination is
known as the median notch or pygidial incision.
The dorsal surface of the pygidium cephalad of the anus
sometimes contains a reticulated area. This area may be of some
size and cover a considerable part of the cephalic portion of the
pygidium or it may be very small in extent. The reticulations
are closely crowded together and the appearance is due to the
thickenings of the margins of the small areas. Green has com-
pared the pattern of the reticulations to that of crocodile-leather.
Their pattern and shape are responsible for their name of lattice-
shaped thickenings. The functioning of these thickenings is not
clear and they have evidently arisen independently several times
SUBFAMILY DIASPIDINAE 235
since they occur in widely separated groups.
There is located in the cephalic region of the dorsal aspect
of the pygidium of certain species a curved transverse band of
thickenings which were named by Leonardi the calli and here
designated as the calles. They are the transverse thickenings of
Marlatt and the basal thickenings of Dietz and Morrison. The
calles exist in four conditions. There may be four distinct sepa-
rate thickenings, two on each side of the meson, the one on each
side adjacent to the meson is known as a mesal callis and the
one laterad of each mesal callis as a lateral callis; there may be
a transverse band of three thickenings, the mesal one much long-
er than either of the lateral and evidently formed by a fusion of
the mesal calles; there may be a transverse band of two thicken-
ings, one on each side of the meson, each evidently formed by a
fusion of the mesal and lateral calles of a side; or there may be
a single transverse thickening, evidently formed by a fusion of
the two mesal and two lateral calles.
In certain species there are thickenings which appear to be
proximal prolongations of the lobes. These thickenings extend
through the cuticle and for some distance beyond the proximal
ends of the lobes. They are characteristic in form for certain
species and only of certain individuals of other species. They
seem to vary considerable in extent and appearance depending
upon the amount of treatment given the specimen with caustic
potash. They are designated as anomaladensae to distinguish
them from the other thickenings of the pygidium. They are in-
correctly designated as ventral thickenings by some writers.
On the ventral aspect of the pygidium of many species, there
are more or less distinct thickenings that are similar in general
appearance to the calles, but they are situated on the ventral
aspect and extend longitudinally instead of transversely. They
may be known as the paradensae and in common with some of
the other thickenings of the pygidium have been previously called
ventral thickenings. The paradensae are located laterad of the
vulva, but vary somewhat in position. They are more frequently
located on the mesal than on the lateral side of the genacerores.
In some species they are located mesad of the postgenacerores
and laterad of the pregenacerores. In such cases there are two
portions which may be connected by an oblique thickening or the
connection may be wanting. If necessary to distinguish these
two portions of each side, the cephalic portion may be known as
a preparadensa and the caudal as a postparadensa. The portion
236 THE COCCIDAE
laterad of each pregenaeeroris, the preparadensa, is sometimes
wanting, the postparadensae are always so far as observed more
distinct than the preparadensae. The paradensae are generally
obsolete in species lacking genacerores.
The cerores located on the ventral aspect near the meson and
the middle of the pygidium are the genacerores. They are, so
far as known, peculiar to the adult females of certain species of
Diaspidinae, but they may be homologous with the cerores which
are peculiar to the vulvular region of certain adult females of
other subfamilies. The genacerores are also known as filieres,
spinnerets, circumgential glands, circumgential gland openings
or orifices, circumgential pores, ventral glands, grouped glands,
ventral grouped glands, wax glands, fusi, filiere aggregate, disculi
ciripari, disculi ciripari perivulvarea, plaque de filieres, dischi-
ciripari, or paragenitals.
The genacerores are usually arranged in four or five groups.
When there are five groups present, one group is located on the
meson cephalad of the vulva and two groups on each side laterad
of the vulva. The mesal group, the mesogenacerores, is
variously designated as the anterior, mesal, median, or upper
group. The cephalic group of each side, the pregenacerores, is
variously known as the anterior-lateral group, cephalo-lateral
group, upper lateral group, cephalolateral group, or the anterior
laterals. The caudal group of each side, the postg6nacerores, is
variously known as the posterior lateral group, the caudo-lateral
group, lower lateral group, caudolateral group, or posterior lat-
erals.
In the following descriptions of species, the maximum and
minimum number of cerores in the groups of genacerores are in-
dicated in the following way: 0-9 (20-25) 10-19 or (7-9)2-6.
The first group of numbers in front of the parenthesis shows the
number of mesogenacerores, none to nine, the second group, al-
ways enclosed in a parenthesis for this group, shows the number
of pregenacerores, and the third group the number of postgena-
cerores. "Where only two groups of numbers are shown, the first
^•group is included in parenthesis and represents the number of
pregenacerores and the other the number of postgenacerores, the
3nesogenacerores being wanting.
The number of cerores in the groups of genacerores are indi-
cated by different authors by the use of formulae of various forms.
Those of Leonardi are indicated thus:
SUBFAMILY DIASPIDINAE 237
24-24 Or 6X5 Or
17-21
8-13
15-17 2X2
These translated into the form just given, would be: 14(24-24)
15-17 or 3(5-6)2-2 or (17-21)8-13. The most intricate formulae
are those used by Lindinger. The following are a few samples
translated in each case to the form given in the preceding para-
graph: 5-9: 5-10: 5-7: 5-10: 5-9 equals 5-7(5-10)5-9; 23: 27: 19:
32: 19 equals 19(27-32)19-23; 1-4: 1-4: 0: 1-4: 1-4 equals (1-4)
1-4; 0: 0-2: 0-2: 0 equals 0(0-2)0; 10: 11 10: 9: 8; 4: 10: 8:
7: 9; 10: 18: 11: 15: 10 equals 10(9-11)8-10 and 8(7-10)4-9 and
11(15-18)10-10. The last formula is to show the number in
three individuals. If colons have be^n accidentUy substituted
for the semicolons in the printing, as sometimes happens, there is
difficulty in the interpretation of the formula unless one is famil-
iar with the underlying principal of the construction of the for-
mula. Each formula is apparently to consist of five groups of
numbers separated by colons, a group of numbers for each group
of genacerores, and the second five groups of numbers, whether
separated by a semicolon from the first five groups or not, repre-
sents another individual and is a different formula.
The mesogenacerores usually have fewer cerores than either
of the other groups and are frequently wanting or represented
by one or two cerores. They sometimes combine with the pre-
genacerores and form a single large crescentic cephalic group.
Such a group is known as a solaceroris. The postgenacerores
being distinct, in such cases, the pygidium is said to have three
groups of genacerores. The pregenacerores and the postgena-
cerores sometimes combine forming a single elongate group on
each side of the vulva. The group of each side is known as a
latagenaceroris. The pygidium in such cases is said to have three
groups of genacerores if the mesogenacerores are present and
two if they are wanting. In a few cases the genacerores form
a large U-shaped group, from their form evidently arising from
a fusion of all five groups of genacerores, the omnagenaceroris.
When two or more groups of genacerores combine or fuse they are
sometimes described as genacerores confluent. There are in cer-
tain genera as Poliaspis, more than five groups of genacerores
present. In certain species of this genus, as media Maskell and
cycadis Comstock, the three extra groups of genacerores appear
to be groups of cerores of the abdominal segments cephalad of
the one bearing the mesogenacerores. In other species the extra
groups appear to have been formed by the subdivision of some
238 THE COCCIDAE
of the primary groups. This latter method may have been the
original method of formation and the appearance in the species
named may be due to a reduction in the number of cerores and
their later migration and isolation.
The number of cerores in each group varies greatly. There
may be only a few, as one, two, or three, or there may be forty or
more. This variation in the number of cerores, unfortunately
is not fixed even within specific limits, but the comparative num-
ber in a group is often of value in separating species or groups
of species.
Each genaceroris is a disk-shaped ceroris with several irreg-
ularly arranged comparatively large openings. The fact that
they are present only in the adult is of value in those species
possessing them in deciding whether a given individual is an
adult or a second stage female nymph. The genacerores were
originally considered as forming a considerable part of the scale
and because of this belief they were named spinnerets. They
excrete a fine powdery wax which is mixed with the eggs as they
are extruded. Green has called attention to the fact that most
of the species provided with genacerores lay eggs, while those
species without these structures give birth to living young.
Those species provided with genacerores and giving birth to liv-
ing young possess only a very few cerores in each group. The
genacerores are similar in structure to the spiracerores, but in
certain species have a fewer number of openings than the gena-
cerores. The fact that the genacerores are present only in the
adult female and their location about the vulva would suggest
that they must be homologous with the cerores peculiar to other
adult females, certain eriococcids, and from which the wax for
covering the eggs is excreted.
Each female and male of the Diaspidinae is covered by a thin
sheet or pellicle known as a scale. It is frequently incorrectly
designated as a puparium. This term should be applied only
to the last nymphal exuvia in which the adult female of certain
coccids, as the females of the genus Aonidia, pass their life and
the last larval cuticle or exuvia in which the pupae of certain
Coleoptera and Diptera complete their pupal life. The scale is
composed of three distinct products, the molted skins or cuticle
of the first and second nymphal stages in the female and the first
in the male, a mass of threads of wax which are poured from the
oraceratubae and fuse upon coming into contact with each other,
and according to Berlese to a mass of excretion formed by the
^
SUBFAMILY DIASPIDINAE f^ 239
Malphigian tubules and poured out through the anus. In the
case of the dark or blackish scales the coloring matter isjocated
in the excretion from the Malphigian tubules. The remarkable
white varieties, as in the case of Lepidosaphes ulmi Candida, are
imperfect individuals which were unable to produce the secretion
necessary to color the scale. It is not unusual to find individuals
in large colonies containing many individuals in which the cephal-
ic half of the scale is normal in color, black or brownish, and the
caudal half is white or uncolored.
The English and some other writers usually designate the
exuviae as the pellicles, the first exuvia as the larval pellicle and
the second exuvia as the nymphal pellicle. Most American
writers usually employ a word derived from the Latin word
exuviae, which was used by the Romans to designate the cast off
skin of animals. It was used in the pleural only. Some writers
apparently do not differentiate between the cast skins of the first
and second nymphal stages and always use the word exuviae but
evidently only in the pleural form. Comstock and Cockerell em-
ploy the word exuviae for both female and male scales, evidentlj-
using it in both a pleural and a singular sense. Some have used
a neuter form, exuvium, for the singular and the regular pleural
form, exuvia. These forms have found their way into the Ameri-
can dictionaries. Ferris writes of the first exuvium, the second
exuvium, and the exuviae. So far as observed, Cooley was the
first to use the singular form, exuvia. He is followed in this by
Marlatt. The regular Latin singular of exuviae would be exuvia.
These forms have been adopted.
The scale consists of two parts, a dorsal scale, sometime
simply designated as the scale, and a ventral scale. The ventral
scale is frequently very thin and adheres closely to the surface
of the host-plant, but it may be well developed and fused with
the dorsal scale. As each exuvia is molted, it is secured to the
sheet of wax of which the scale is formed by a coating on its
inner surface. As the second exuvia is cast later, this explains
its position under the first, but there is always a sheet of wax
between them. In the case of those aspidiotids which rupture
their exuvia around the margin of the body, the dorsal portion
of the exuvia is imbedded in the dorsal scale and the ventral
portion in the ventral scale. Because the ventral portion of the
exuvia is thin and delicate, it is frequently overlooked, but in
many cases where the ventral scale is very thin, it is likely that
the ventral portion of the exuvia is lost. In all other Diaspidinac
240 THE COCCIDAE
the first exuvia remains in a single piece and since the antennae
remain attached to the dorsal portion of the exuvia, it is possible,
unless they have been broken off by the elements, to identify the
antennae on the first exuvia of the adult scale. The presence
or absence of the antennae on the dorsal portion of the first exuvia
may prove of value in determining the tribal location of a given
species.
The scales vary greatly in shape. In some species the
exuviae are nearly as large as the entire scale, while in others
they are not only much smaller but form only a small portion of
the scale in comparison with the much greater extent of the wax
or excreted portion. "Where the scales are elongate, the exuviae
are placed at one end and usually project beyond its margin. The
portion of the scale bearing the exuviae is always narrow and
the scale gradually widens toward the caudal end. In the circu-
lar scales the exuviae are usually placed within the periphery and
in the Aspidiotini, which usually have this type of scale, they
are located at or near the center of the scale. The body of the
insect is generally located under or near the exuviae.
The young nymphs of the first stage usually form a minute
ball or button of threads of wax as the first part of their scale.
This becomes melted into a compact mass and can be identified
as the minute dot over the first exuvia of the scales of many
adult females. This dot may be known as the cicatrix. Com-
stock has described and figured the peculiar rolls of cicatrices
formed by certain individuals which do not get beyond this stage
in the formation of their scale.
The scales of all adult females contain typically two exuviae.
A few genera in most of the tribes as Aonidia do not molt their
second exuvia, but shrink away from it and transform to adults
within it, using this second exuvia as a puparium which conse-
quently never becomes attached to the scale. Some of the species
transforming in a puparium, particularly those with a long or
pyriform scale, have a transparent scale so that the puparium
can be identified through the scale and from its position appears
to be the second exuvia. With the exception of these genera, the
presence of a single exuvia indicates that the enclosed insect is
in the second nymphal stage if a female or if a male, that it is in
any stage after the first nymphal stage. The scales of the males
are always much smaller than those of the adult female, are
usually quite similar to those of the second nymphal female, and
are always provided with a single exuvia. In a few genera as
SUBFAMILY DIASPIDINAE 241
Diaspis, Aulascapis, Chionaspis, etc., the male scales are easily
identified because the secretionary portion is frequently distinctly
longitudinally carinate.
The tribes of Diaspidinae, of which there are six, have been
arranged in the order of the following tabulation which indicates
something not only as to the relation of the tribes but of their
lines of specialization: —
SYNOPSIS OF THE DIASPIDINAE
GENERALIZED DIASPIDINAE.— Pygidum with Altaceratubae.
Pygidium with Pectinae.
Preabdomen with Bracteae.
1. The Fringed Scales. — Parlatoriini.
Preabdomen without Bracteae.
2. The W^ite-shield Scales. — Leucaspidini.
Pygidium with Plates.
Brevaceratubae Conservers.
Altaceratubae opening near Margin.
3. The Oyster-shell Scales. — Lepidosaphini.
Altaceratubae opening in Latadentes.
4. The Double-shield Scales. — Diaspidini.
Brevaceratubae Loosers 5. The Uhlerian Scales. — Fioriniini.
SPECIALIZED DIASPIDINAE.— Pygidium without Altaceratubae.
6. The Deltoid Scales.— Aspidiotini.
What is considered as the most generalized condition of the
Diaspidinae is found in certain species of Parlatoriini. The re-
tention of distapectinae in the Leucaspini, although they have
departed considerably from the condition in the parlatoriids,
allies them with the latter. The presence of plates allies the
Lepidosaphini, Diaspididni, and Fioriniini. In the first two the re-
tention of brevaceratubae distinguishes them from the third,
while the presence of plates in the mesal incisura of most Lepido-
saphini shows a more generalized condition than their absence
in most Diaspidini, but the location of the anus much nearer the
caudal margin of the pygidium than in the latter would show a
more generalized condition for the Diaspidini. The Aspidiotini
are without altaceratubae, have pectinae in the generalized genera,
plates in the specialized, and the absence of both and the develop-
ment of pseudolobes in the most specialized.
The Parlatoriini, Lepidosaphini, Diaspidini, and Aspidiotini
contain among their most highly specialized genera those that
transform in a puparium. In the Leucaspini and Fioriniini all or
most of the species transform in this way. There is, except in
in the last two named tribes, the same line of specialization
242 THE COCCIDAE
shown in all in the modification of the pygidium and the pygidial
fringe. The generalized pygidium contains five distinct groups
of genacerores, at least three pairs of lobes, and two or three
broad distapectinae in each incisura and several on each lateris.
The line of modification is a transformation of the pectinae into
plates, a reduction in the number of plates, a reduction in the
size of the lobes, and finally in the number of lobes. The modi-
fication proceeds always along each side of the pygidium from
the cephalic end to the meson or caudal end. With the reduction
in the pygidial fringe there proceeds also a reduction in the num-
ber of cerores in the groups of genacerores and finally their com-
plete loss. One of the highest types of specialization is repre-
sented by a few species where lobes, plates, pectinae, and gena-
cerores are wanting and the projections of the pygidium are
limited to latadentes. Whether there is an acceleration of the
development and in the last named species the pygidial fringe is
to be found only in the pygidium of the second nymphal stage
is not known. But representing the highest type of specializa-
tion in each of the four tribes first named are genera in which
the adult female shrinks away from the last or second nymphal
skin but never escapes from it. The young are born alive and
escape through openings in the cuticle, that is the second exuvia
or puparium as it is designated in such species. The pygidium
of the adult female may be entirely without projections or bear
a large number, none of which are homologous with any part of
the pygidial fringe. These projections, the pseudolobes, are fre-
quently asymmetrical. There has been an acceleration in the de-
velopment of the pygidial fringe and it is frequently found in
the pygidium of the second nymphal stage, but in certain species
the pygidial fringe is wanting even in the second nymphal stage
and is to be found only in the pygidium of the first nymphal
stage. The species transforming in a puparium represent the
highest type of modification and specialization.
The expression in the following table, of the adult female or
of the second nymphal female, does not imply that both of these
stages in all species must be examined before the position of any
given species can be determined. It should not be forgotten that
the characters of the pygidium of the second nymphal female can
be secured at all times by mounting the second exuvia of the
scale, It does mean, that, if the structure named is wanting in the
adult female, the pygidium of the second nymphal female should
be examined to determine whether it is present or absent in this
SUBFAMILY DIASPIDINAE 243
stage. This statement is added so that the species transforming
in puparia, if it is known that the species transform in this way,
need have only the second nymphal stage examined. Specimens
belonging to the various tribes of Diaspidinae can be distinguished
by means of the following table: —
TRIBES OF DIASPIDINAE
a. Pygidium of adult female or of second nymphal female always with
altaceratubae and frequently with brevaceratubae ; scale of adult
female usually elongate, several times longer than broad with
exuviae at one end, if shorter or round, exuviae placed at or just
within periphery, exuviae never central or superimposed, placed al-
most end to end, overlapping slightly; exuvia of first nymphal fe-
male with remains of antennae forming porrect appendages; pygi-
dium, if lacking altaceratubae, with elongate scale with single termi-
nal exuvia with porrect antennae.
b. Pygidium of adult female or of second nymphal female usually
with pectinae, rarely with plates; mesogenacerores frequently
wanting and, when present, rarely with more than eight cerores;
scale of adult female sometimes almost completely covered by
second exuvia.
c. Preabdomen of adult female or of second nymphal female with
lateral margins of its segments, at least caudal ones, bearing
bracteae; adult female sometimes enclosed in puparium, if
not enclosed, pygidium of adult female with distapectinae and
altaceratubae, if transforming in puparium, second nymphal
female with similar structures; second exuvia of scale of adult
female large, often almost covering scale Parlatoriini.
CO. Preabdomen of adult female or of second nymphal female
never with lateral margins of any of its segments bearing
bracteae; adult female always enclosed in puparium and fre-
quently without altaceratubae and with furcate pseudolobes,
second nymphal female with altaceratubae and distapectinae;
second exuvia of scale of adult female moderate in size, not
attached to scale, and always less than one-half length of
scale. Leucaspidini.
M. Pygidium of adult female or of second nymphal female usually
with plates, at most only with f urcapectinae ; mesogenacerores
usually present and usually with more than eight cerores; scale
of adult female with second exuvia always small, rarely if ever,
covering one-half of scale.
c. Pygidium of adult female or of second nymphal female usually
with distinct brevaceratubae, irregularly arranged or arranged
in transverse rows; adult female rarely enclosed in puparium;
pygidium never with linaceratubae or clavaceratubae.
d. Pygidium of adult female or of second nymphal female with
plates between median lobes, rarely wanting; body of adult
female always twice or more as long as broad; altaceratubae
usually arranged in pairs, rarely opening in latadentes, their
244 THE COCCIDAE
oraceratubae elongate oval in outline, frequently twice as
long as broad; scale of adult female elongate, coriaceous,
or horny, usually brownish or blackish in color, shaped and
ridged like an oyster-shell, exuviae always at one end, sides
sometimes parallel or subparallel ; scale of adult male never
carinate and usually similar in color to that of female
Lepidosaphini.
dd. Pygidium of adult female or of second nymphal female
rarely with plates between median pair of lobes; body of
adult female always distinctly longer than broad, rarely
twice or more as long as broad, if so, always without plates
between median pair of lobes; altaceratubae usually arranged
singly, frequently opening in latadentes, their oraceratubae
circular or subcircular in outline; scale of adult female
pyriform, elongate or round, sometimes shaped and ridged
like oyster-shell and, if so shaped, always without plates be-
tween median pair of lobes, rarely if ever with sides parallel
or subparallel, generally white in color or light colored, and
closely felted; scale of adult male often longitudinally cari-
nate and chalky in color and appearance Diaspidini.
cc. Pygidium of adult female or of second nymphal female always
without brevaceratubae ; scale of adult female always elongate
and with single exuvia; adult female always enclosed in
puparium, excretionary part of scale thin and puparium,
which is generally nearly as large as scale, apparently form-
ing second exuvia; pygidium sometimes apparently lacking
altaceratubae and provided with linaceratubae and clavacera-
tubae. FioRiNiiNi.
aa. Pygidium of adult female or of second nymphal female never with
altaceratubae, rarely with brevaceratubae, and if provided with
brevaceratubae, not arranged in transverse rows, usually with lina-
ceratubae or clavaceratubae ; scale of adiult female circular or
nearly so, rarely elongate, if so, exuviae superimposed; exuviae
always central or nearly so, first exuvia superimposed over center
of second, never projecting beyond periphery or attached at peri-
phery; exuvia of first nymphal female never with remains of
antennae forming porrect appendages; pygidium rarely with dista-
pectinae or furcapectinae, frequently with latapectinae, unapectinae,
or plates, if apparently with altaceratubae scale always round with
central exuvia and first exuvia without remains of antennae; scale
of adult female with second exuvia small and never covering one-
half of scale; adult female sometimes enclosed in puparium
ASPIDIOTINI.
>j ' CHAPTER XXII
TRIBE PARLATORIINI
The Fringe Scales
The body of the adult female is usually longer than broad with the
cephalic and caudal ends bluntly rounded and with the sides parallel
or subparallel or as broad or broader than long with the body triangu-
lar in outline and the caudal portion with converging lateral margins
and bluntly pointed caudal end. The lateral margins of the segments
of the preabdomen are provided with bracteae either in the adult female
or the second nymphal female. Lobes are present in the adult female
or second nymphal female. There are usually three pairs, rarely four
pairs, sometimes with the fourth and fifth pairs rudimentary, and
sometimes with only two pairs, if wanting, present in the second
nymphal stage. Distapectinae are present in either adult female or
second nymphal female, rarely reduced until they approximate furca-
pectinae in form, somatinues wanting in adjult jEemale and second
nymphal female. Plates are rarely present. Incisurae are broad and
shallow, usually completely filled by pectinae, median incisura rarely with-
out pectinae. Densariae or paraphyses are never present. Altacera-
tubae are present and are usually very large. Brevaceratubae may be
numerous, few in number, or wanting, but are never arranged in trans-
verse rows. Genacerores are present or wanting, when present, repre-
siented by four or five groups, the mesogenacerores are generally want-
ing and, if present, rarely with more than eight cerores. Scale of adult
female variable in form, round or almost round with exuviae small
and peripheral, subquadrangular with the exuviae large and covering
miost of surface, or two or three times as long as wide with exuviae
at one end. Exuviae usually two in number, female sometimes trans-
forms in a puparium, in such cases with only a single exuvia. Scale
of male is usually elongate, twice as long as wide, with the exuvia at
one end, rarely subcircular with the exuvia subcentral.
The genera of Parlatoriini fall into two groups, those whicla
are normal in their development and those in which the adult
female is enclosed in a puparium. This latter group includes a
single genus, Gymnaspis. The pygidium in the species of this
genus is greatly modified in the adult female. In some there is
no indication of lobes, pectinae, or incisurae, but the typical
pygidial structures are found in the second stage of the female
nymph.
The scale of the adult female varies considerable in form.
245
246 THE COCCIDAE
It may be elongate, over three times as long as broad; ovate, not
quite twice as long as wide; or circular or subcircular. The
exuvia overlap and are placed at one end or project beyond the
periphery of the circular or subcircular scales. In the ovate
scales the second exuvia is very large in comparison to the ex-
tent of the w^ax and forms the greater part of the scale. In the
elongate and circular scales the area of wax is two or more
times the area covered by the exuviae. The wax of the scale is
thin and delicate and usually not strongly colored. The exuviae
in a few species are black.
The scale of the male is elongate, two or more times as long
as broad with the sides nearly straight and subparallel. The
single exuvia is attached at one end and the area of wax is much
greater than the area covered by the exuvia. The scale is similar
in color to that of the female and its exuvia is sometimes black.
The name of fringed scales is derived from the characteristic
appearance of the pygidial fringe of many species, which consists
of a continuous margin of lobes and large distapectinae. Equally
conspicuous are the large altaceratubae with their oraceratubae
located near the proximal ends of the pectinae. The cephalic
limits of the pygidium are usually not distinct and there is a
gradual mergence of the segments of the preabdomen and pygi-
dium. This is still more striking in the nymphal stages of the
female. The dorsal aspect bears numerous brevaceratubae, but
they are promiscuously distributed and offer no indication as to
the segmentation of the pygidium. There are typically three
pairs of lobes present, but this number may be reduced in some
species while as many as five pairs can be identified in others.
This arises fom the fact that in some species the cephalic pairs
of lobes are not sharply different in form from the pectinae, in
others they are distinctly bluntly or pointedly rounded at the
distal end and are lobe-like and not pectinae-like in shape. The
pectinae are generally broad and quite constant in number, two
in the median incisura, two in each second incisura, three in
each third incisura, three in each fourth incisura, and three or
four in each of the other incisurae. There are slender lamacera-
tubae extending through the pectinae. The genacerores are
wanting in all of the adult females enclosed in a puparium, while
there are four or five groups in the species of other genera. The
mesogenacerores are frequently wanting and when present con-
sist of only a few cerores, usually less than eight. There is con-
siderable variation in the number of cerores in the groups of
TRIBE PARLATORIINI 247
pregenacerores and post genacer ores, ranging from four to twenty
or more. In Euparlatoria viridis the pregenacerores and postgena-
cerores are continuous, form latagenacerores, with only a slight
constriction marking the division between the two groups and
the pygidium appears, from a cursory examination, to have only
two groups of genacerores, each consisting of 23 to 26 cerores.
Species of Parlatoriini are found in all parts of the world,
but they are more abundant in tropical and subtropical regions.
In temperate regions they are able to survive only on plants
grown under glass. Several of the species are pests upon the
fruits, trees, and shrubs of cultivated tropical fruits of different
kinds.
The only complete account of the species of Parlatoriini is
that of Leonardi: "Generi e specie di Diaspiti, saggio di siste-
matica delle Parlatoriae, " published in 1903. This paper is now
out of date since many species have been described since its pub-
lication. In this work two genera are recognized, Parlatoria and
Gymnaspis. The former genus, Parlatoria, which has ziziphi
Lucas as type, was divided into two subgenera, Euparlatoria for
which calianthina Berl. & Leon, may be taken as the type, and
Websteriella which has blanchardi Targ. as type. Since blan-
chardi and ziziphi are congeneric, Websteriella must give way to
the older name of Parlatoria. Apteronidia Berl., not Leonardi as
given in the Catalogue of Coccidae, was first suggested in 1896
and also has blanchardi Targ. as type. The name Euparlatoria
published in 1900 is antedated by Syngenaspis Sulc published in
1895 with parlatoriae Sulc as type. This species of Sulc and cali-
anthina Berl. & Leon, are congeneric.
The species of Gymnaspis are liable to be confused with those
of the genus Aonidia, a genus of Aspidiotini. The second exuvia
or puparium in Gymngtspis is almost as large as the entire scale
and the first exuvia is peripheral instead of central in position.
In Aonidia the second exuvia or puparium is larger than is usual
in the scales of the Aspidiotini, but it is much smaller than those
of the Parlatoriini and in addition the first exuvia is central in
position. The practical difficulty in differentiating the two genera
is that the first exuvia is lost. It is apparently not firmly at-
tached to the scale. The presence or absence of altaceratubae in
the pygidium of the second nymphal female or both the second
and the first nymphal females needs to be recorded and this char-
acter used for identifying the genera of Aspidiotini that trans-
form in a puparium. The species of this tribe, so far as observed,
248 THE COCCIDAE
always lack altaceratubae. It is likely that several of the speciesf
described as belonging to the genus Aonida may be found to be«
long to Gymnaspis when the pygidia of their nymphal stages are
described or figured.
GENERA OF PARLATORIINI
a. Pygidium of adult female with groups of genacerores.
b. Pygidium of adult female always with lohes and pectinae or
plates; adult female never enclosed in puparium.
c. Pygidium of adult female with four or five groups of gena-
cerores.
d. Pygidium of adult female always with pectinae and more
than two pairs of lohes.
e. Scale of adult female with second exuvia of medium size,
never covering one-half of scale Syngenaspis Sulc.
ee. Scale of adult female with second exuvia large, covering
two-thirds or more of scale Parlatoria Targ.
dd. Pygidium of adult female without pectinae and with plates;
scale of adult female with second exuvia large, covering
greater part of scale Cryptoparlatoria Lindgr.
cc. Pygidium of adult female with two groups of genacerores.
— Type, Gymnaspis aberemoae Lindgr Lindingeria MacG.
hb. Pygidium of adult female always with pseudolohes and without
lobes and pectinae or plates; adult female enclosed in puparium.
Porogymnaspis Green,
aa. Pygidium of adult female without groups of genacerores.
b. Pygidium of adult female and of second nymphal female with
lobes and pectinae; scale of adult female with two exuviae, adult
female never enclosed in puparium. — Type, Parlatoria pseudaspidi-
otus Lindgr. Genaparlatoria MacG.
bb. Pygidium of adult female with simple or toothed pseudolohes and
without pectinae; second nymphal female with lobes and pecti-
nae; scale of adult female with single exuvia, adult female al-
ways enclosed in puparium. Gymnaspis Newst.
SPECIES OF SYNGENASPIS
a. Pygidium with five groups of genacerores.
b. Pygidium with four pairs of lobes, all with dintinct lateral notch,
fourth pair of lobes small; pectinae broad, two in each incisura,
and filling space; genacerores 0-7(10-25)11-23; altaceratubae 19,
one mesal and nine on each side; scale of adult female circular
or subcircular, white or whitish with yellow to fuscous exuviae. —
Parlatoria (Euparlatoria) calianthina Berl. & Leon. — Italy, Al-
geria on Rosa, Fraxinus, Olea, peach, apple, almond olea Colv.
bb. Pygidium with three pairs of lobes.
c. Pygidium with each group of pregenacerores consisting of
about twenty cerores.
TRIBE PARLATORnNI 249
d. Pygidium withL the median pair of lobes with prominent
lateral and mesal notches,
e. Scale of adult female oval in outline, pale ochrecnis, sec-
ond exuvia black or blackish; pygidium with median pair
of lobes with lateral and mesal notches; genacerores 1-1
(20-20)7-7. — Japan, New York on tea, maple theae Ckll.
ee. Scale of adult female circular in outline, white or gray-
ish, second exuvia dark greenish to black; genacerores
1-4(9-16)16-17. — Japan on Ilex theae viridis Ckll.
dd. Pygidium with median pair of lobes with inconspicuous
lateral notch; second pair of lobes similar in shape with
more distinct lateral notch; third pair of lobes broad,
truncate; lobes close together, pectinae long and narrow,
arranged 2, 2, 3, 6; genacerores 8-10(20-22)20-22; scale of
adult female pyriform, pale ferruginous, exuviae pale red-
dish.— ^Australia on Banksia. hanksiae Mask.
cc. Pygidium with each group of pregenacerores consisting of
about eight to ten genacerores.
d. Pygidium of adult female with lobes broadly truncate at
distal end with notch on lateral margin; lobes far apart and
incisurae filled with broad distapectinae ; altaceratubae 7,
one mesal and three on each side; mesospiracerores none;
anus located midway between vulva and caudal margin;
genacerores 3-4(7-13)7-10. — Bohemia on Abies
parlatoriae Sulc.
dd. Pygidium of adult female with lobes subtriangular, mesal
margins straight, median pair of lobes largest, with one
mesal and two lateral notches; second pair of lobes with
distal end rounded, with two lateral notches; third pair of
lobes similar to second but smaller; fourth pair of lobes
sometimes represented by crenate projection; incisurae
strongly chitinized; pectinae subequal in length to lobes,
narrow, of usual number, unapectinae; genacerores 1-2(10-14)
9-14. Pygidium as figured is distinctly of aspidiotid type. —
Society Islands on orange. cinerea Doane & Hadden.
Pygidium with four groups of genacerores.
b. Pygidium with four pairs of lobes.
c. Scale of adult female elongate, pyriform, distinctly longer
than broad, whitish to yellow brown, exuviae dark; lobes with
median, second, and third pairs with lateral and mesal
notches; fourth pair of lobes narrow, elongate, bluntly point-
ed, unnotched; anus located a little caudad of vulva; gena-
cerores (9-14)8-12. — Australia on Myrtus, Viburnum
myrtus Mask,
cc. Scale of adult female variable, usually circular, sometimes
elongate or pyriform.
d. Pygidium of adult female with genacerores (6-10)6-9.
e. Scale of adult female dusky whitish tinged with brown-
ish, exuviae yellowish; median, second, and third pairs
250 THE COCCIDAE
of lobes with mesal and lateral notches; fourth pair
of lobes variable in size and form, always shorter than
third, subequal in width, usually pointed or tridentate;
distapectinae broad and filling incisurae. — District of
Columbia, Florida, Louisiana, Ohio, Kansas, California,
Mexico, Hawaiian Islands, Japan, Europe, Algeria, Cey-
lon on Japonica, lemon, orange, pergandei Comst,
ee. Scale of adult female with secretionary portion pale trans-
parent ochreous and exuviae bright ca&taneous or brown-
ish orange with broad black mesal band; pygidium as in
typical specimens. — Ceylon on Phyllanthus
pergandei phyllanthi Green,
dd. Pygidium of adult female with genacerores (6-6)5-5; scale
pale to dark slate color, exuviae yellowish; median, second,
and third pairs of lobes narrow, bluntly rounded, with mesal
and lateral notches; fourth pair of lobes minute, similar in
general form to third, distinctly pointed, notched; dista-
pectinae subequal in length with lobes, filling incisurae. —
Philippine Islands on Cocus greeni Banks.
bb. Pygidium with three pairs of lobes.
c. Pygidium with four distinct groups of genacerores.
d. Pygidium with third pair of lobes not notched.
e. Pygidium with the median and second pair of lobes of
adult female faintly notched, third pair with distal end
rounded, sides subparallel; pectinae of unapectina type;
scale of adult female elongate, very wide, slate-gray,
exuviae dark brown. — ^West Australia on Petrophila,
Hakea. petrophilae Fuller.
ee. Pygidium with all three pairs of lobes of adult female
obliquely truncate, mesal margin longer than lateral,
entire; fourth pair of lobes represented by broadly
rounded area; pectinae narrow, hardly as long as lobes,
finely toothed; scale of adult female pale yellow or brown-
ish, exuviae yellow; mesospiracerores 2, metaspiracerores
wanting; genacerores (10-14)9-14; anus located nearer
transverse line drawn midway between vulva and caudal
margin than vulva; altaceratuba 19, one mesal and nine
on each side. — Ceylon on Flacourtia, Scolopia
cingala Green,
dd. Pygidium with all lobes notched.
e. Pygidium with each group of pregenacerores consisting
of about twehty cerores; scale circular or ovate, brownish,
exuviae brownish or greenish; genacerores (21-23)12-18.
— Japan on Euonymus. thea euonymi Ckll.
ee. Pygidium with each group of pregenacerores consisting
of fifteen cerores or less.
f. Pygidium of adult female with median pair of lobes
elongate, pointed, mesal and lateral margins crenulate;
TRIBE PARLATORIINAE 251
second pair of lobes broader and longer than first pair,
distal end bluntly rounded, mesal and lateral margins
crenulate; third pair of lobes smaller with mesal and
lateral notches; pectinae slightly longer than lobes,
those of median incisura approaching latapectinae in
form, others distapectinae ; genacerores (4-4)7-7. —
Borneo on Gnetum. hastata Lindgr.
ff. Pygidium of adult female with median pair of lobes as
broad as or broader than second pair, their distal ends
broadly rounded, never pointed.
g. Scale of adult female circular or oval, never twice as
long as wide.
h. Scale of adult female pinkish-grey and finely
punctate, exuviae dark brown to blackish brown;
genacerores (10-11)10-11. — Australia on Dryandra.
dryandrae Fuller,
hh. Scale of adult female orange-yellow, sometimes
tinged with green, exuviae yellowish, darker than
excretionary portion; lobes comparatively nar-
row, subconical, bluntly pointed, with more or
less distinct mesal and lateral notches; pectinae
broad and deeply toothed; altaceratubae 19, one
mesal and nine on each side; anus located midway
between vulva and caudal margin; spiracerores
wanting; genacerores (5-7)4-7. — North America,
South America, Europe, northern Africa, Aus-
tralia, Japan, China, Ceylon, Hawaiian Islands on
Selenipedium, Vanda, Macrosamia, Pinus, Myrtes,
Citrus, Camellia, Cymbidium, Pyrus, apple, date-
palm. proteus Curtis.
gg. Scale of adult female elongate with subparallel sides,
more than twice as long as wide, brownish fulvous,
exuviae yellow; lobes comparatively narrow, dis-
tant, distal end bluntly pointed, with mesal and lat-
eral notches; fourth pair of lobes sometimes repre-
sented by rudiment; pectinae of median, second, and
third incisurae narrow, others wide; spiracerores
wanting; genacerores (6-6)5-5; brevaceratubae nu-
merous; altaceratubae distinct, one mesal and nine
on each side; anus located midway between vulva
and caudal margin. — Ceylon on Psychotria
mytilaspiformis Green,
cc. Pygidium apparently with only two groups of genacerores,
pregenacerores and postgenacerores forming continuous group
on each side, limits indicated by constrictions near middle;
scale of adult female elongate, ovate, rounded behind, whitish
to brown, first exuvia brown, second exuvia veridian green;
lobes with distinct mesal and lateral ndtches, constricted at
proximal end; pectinae narrow; genacerores 23-26. — West Aus-
tralia on Pittosporum. viridis Fuller.
252 THE COCCIDAE
SPECIES OF PARLATORIA
a. Pygidium of adult female with four pairs of lobes, first, second, and
third pairs with mesal and lateral notches.
b. Scale of adult female and both exuviae black; pygidium of adult
female with lobes distant, median, second, and third pairs blunt-
ly rounded, with mesal and lateral notches; four^th pair of
lobes small, bluntly rounded, entire; distapectinae broad, not
truncate, approaching latapectinae in form; anus and vulva lo-
cated near line drawn between pregenacerores and postgenacer-
ores; altaceratubae aboult 17, one mesal and eight on each side;
genacerores (6-7)7-10. — United States, Europe, Australia, Algeria,
China on Zizyphus, date-palm, orange, lemon zizyphi Lucas.
bb. Scale of adult female and both exuviae light colored, exuviae
yellowish, never black; pygidium of adult female with lobes dis-
tant, the median, second, and third pairs typical in form; fourth
pair of lobes small, short, bluntly rounded, entire; distapectinae
broad, truncate, not approaching latapectinae in form; genacerores
(8-11)9-13. — Cryptoparlatoria parla;toreoides Lindg^r. — India on
Xanthophyllum. parlatoreoides Lindgr.
aa. Pygidium of adult female with three pairs of lobes; exuviae of scale
of adult female pale, never black,
b. Pygidium of adult female with median and second pair of lobes
distinctly notched.
c. Pygidium of adult female with sides of lobes parallel or con-
vergent distad, never convergent proximad or constricted;
lobes broader than long, margins serrate; peotinae long and
narrow, slightly longer than lobes, distal ends obscurely
toothed, arranged 2, 2, 2, 2 or 3, those of each lateris adjacent
to lobe; altaceratubae obscure, three on each side; anus cen-
tral in position; spiracerores wanting; genacerores (6-6)4-4;
scale pale yellow, second exuvia almost covering scale, fringed
with narrow area of whitish wax. — Cryptoparlatoria atalantiae
Green. — Ceylon on Atalantia. atalantiae Green.
cc. Pygidium of adult female with sides of lobes not parallel, con-
vergent ait least proximad and sometimes distad, always con-
stricted.
d. Pygidium of adult female with lobes constricted on proxi-
mal half, pointed at distal end with one mesal and two lat-
eral notches, distant; fourth pair of lobes sometimes fully
developed; pectinae long and narrow, subequal in length to
lobes, not filling incisurae, finely toothed, altaceratubae 7,
one mesal and three on each side; brevaceratubae few in
number or wanting; spiracerores wanting; genacerores (4-5)
5-6. — Cryptoparlatoria aonidiformis Green. — Ceylon on Notho-
pegia. aonidiformis Green.
dd. Pygidium of adult female with lobes distinctly constricted
on proximal portion and with distal portion rounded with
mesal and lateral notches; pectinae broad, distinctly longer
than lobes, deeply toothed; altaceratubae 13, one mesal and
six on each side; brevaceratubae numerous; genacerores
TRIBE PARLATORHNI 253
(13-16)11-13; scale of adult female dark greenish-gray, some-
times almost black. — Australia on Pittosporum .
pittospori Mask,
bl). Pygidium of adult female with median and second pair of lobes
constricted at proximal and broadly truncately rounded without
notches at distal end, entire; third pair of lobes with single lat-
eral notch; pectinae broad, not truncate, irregularly toothed;
genacerores (6-10)3-8. — Australia, northern Africa, Arizona on
daJte-palm. hlanctiardi Targ.
SPECIES OF CRYPTOPARLATOREA
a. Pygidium of adult female with three pairs of lobes.
b. Pygidium of adult female with two plates between median pair of
lobes and two plates in each second and third incisura; plates
thorn-like and three or four times as long as lobes; with three
pairs of short, conical, pointed, colorless lobes; each incisura with
latadentis bearing oraceratuba; genacerores (9-11)7-11; meso-
spiracerores 3-4; pygidium of second nymphal female with three
pairs of lobes and with narrow distapectinae. — Japan on Juni-
perus, Chamaecyparis. leucaspis Lindgr.
bb. Pygidium of adult female with single plate between median pair
of lobes and two in each second and third incisura and three
subadjacent plates on each lalteris near each third lobe; plates
longer than lobes, sides parallel, distal fourth nipple-like con-
stricted portion; with three pairs of lobes, one-half or more
length of plates, pointed, each with lateral notch; latadentes
wanting between lobes; genacerores (6-6)4-4; mesospiracerores 1;
pygidium of second nymphal female with three pairs of broad
lobes with lateral and mesal notches; each incisura with two
broad distapectinae and several on each lateris. — Celebes, Philip-
pine Islands on Artocarpus, Mallotus uherifera Lindgr.
aa. Pygidium of adult female with two pairs of lobes.
b. Pygidium of adult female with furcapectinae in median incisura;
with two pairs of lobes, median pair largest, with faint mesal
and lateral notches; second pair of lobes with lateral notch only;
furcapectinae subequal in length to lobes, arranged, 2, 2, 9,
those of each lateris arranged 3-2-2-2; lateres deeply crenulate;
genacerores (10-12)10-12; scale elongate oval, formed of large
dark brown second exuvia and white excretionary portion form-
ing rim in front and lance-shaped projection behind. — Parlatoria
destructor Newst. MSS. — Australia on apple destructor Frogg.
bb. Pygidium without furcapectinae or plates in median incisura.
c. Pygidium of adult female with median lobes subadjacent,
deeply noltched on lateral margin and rounded unnotched on
mesal; second pair of lobes long and narrow, bluntly pointed,
deeply notched on lateral margin; third pair of lobes faintly
indicated; plates arranged, 0, 2, 2, 5, those of each incisura
small and serrate; lateres long and without latadentes; alta-
ceratubae 13, one mesal and six on each side; anus localted on
cephalic side of line drawn midway between vulva and caudal
254 THE COCCIDAE
margin and nearer line than vulva; genacerores 1-4(8-10)8-12.
— Parlatoria pyri Marl. — China on apple, pear pyj-i Marl.
cc. Pygidium of adult female with median pair of lobes subad-
jacent, crenulate with fine notches on oblique lateral margin
and obscure notch on mesal margin; second pair of lobes
small, lateral margin with two to three faint notches; third
pair of lobes not indicated; plates arranged, 0, 2, 2, 3, those
of incisurae small; lateres long without latadentes; anus lo-
cated slightly cephalad of vulva; altaceraltubae about 12, six
on each side, mesal wanting; genacerores (5-6)6-6. — Parlatoria
chinensis Marl. — China, Japan on crab-apple. Hibiscus,
Xanthoxylon, Thuja. chinensis Marl.
SPECIES OF LINDINGERIA
Pygidium of adult female with three pairs of lobes; median pair of
lobes large, as broad as long, broadly rounded, entire; second and
third pairs of lobes similar, slightly smaller, entire; plates truncate,
subequal in length to lobes, arranged, 0, 2, 2, 3, those of each lateris
arranged 2-1; setae large, longer than lobes; scale dark colored, red-
dish, genacerores in two groups of 4-6 cerores each; second nymphal
female with two pairs of narrow rounded lobes with mesal and lateral
ndtches; pectinae longer than lobes, with single short tooth on each
side, arranged 2, 2, 6, those of each lateris arranged 2-2-2; lamacera-
tubae apparently present, also in first nymphal stage. — Gymnaspis
aberemoae Lindgr. — Brazil on Aberemoa al)eremoae Lindgr.
SPECIES OF POROGMYMNASPIS
a. Body of adult female subcircular in outline, lateral portions of
metathorax not produced; scale of adult female consisting of naked
bright red or reddish second exuvia, broadly oval, highly convex;
pygidium of second nymphal female sharply defined, suberect in
position; eight prominent slender lobes; median and second pairs
obscurely notched; third pair of lobes obscurely indented on distal
end; pectinae broad, three teeth; altaceratubae three on each side;
pygidium of adult female truncate, bearing close fringe of tricuspid
processes, four to six more strongly chitinized, similar in form to
other processes, if lobes, other projections arranged 2, 2, 3, 2; anus
seven times its width from caudal margin and one and one-half
times caudad of vulva; genacerores 1-5(10-15)11-17. — North Australia
on Pandanus. rufa Green.
aa. Body of adult female with lateral portions of metathorax produced
on each side inlto rounded projection bearing minute conical
points; scale of adult female consisting of naked second exuvia
with or without first exuvia, yellow or orange with broad trans-
verse blackish fascia; pygidium of second nymphal female with
three pairs of slender lobes, constricted on proximal portion, distal
portion sharply poinlted; pectinae truncate, arranged 2, 2, 3, 3;
each lateris with seven triangular lobe-like projections, as long as
pectinae; altaceratubae three on each side; pygidium of adult fe-
male with six minute broadly triangular projections or lobes and
fringe of tricuspid processes, each attenuate, arranged 2, 2, 3, 3;
each lateris with three triangular projections; anus about six
TRIBE PARLATORIINI 255
times its width from caudal margin and about once caudad of
vulva; genacerores 8-9(9-12), sometimes arranged in two groups or
connected by a series of three or four isolalted cerores. — North Aus-
tralia on Pandanus angulata Green.
SPECIES OF GE-NAPARLATORIA
a. Pygidium of adult female with five pairs of lobes.
b. Pygidium of adult female with median pair of lobes broad and
deeply notched on lateral and mesal margins; second pair of
lobes much smaller, sides parallel, and deeply notched on lateral
margin; third pair of lobes subequal in size to second and faintly
notched on lalteral margin; fourth and fifth pairs of lobes angu-
lar and pointed; pectinae distinct, narrow in median, second,
and third incisurae and subequal in length to lobes, cephalic
pectinae divided on sides, of unapectina-type, arranged 2, 2, 3, 3,
4, 4; dorsum with numberous brevaceratubae; altaceratubae 8,
nine on each side, mesal wanting; anus distant from vulva,
caudad of middle, much nearer middle of distance between vulva
and caudal margin than caudal margin; mesospiracerores and
metaspiracerores 8-10. — Parlatoria mangiferae Marl. — China, Dis-
trict of Columbia on Mango mangiferae Marl.
bb. Pygidium of adult female with median pair of lobes broad,
notched on mesal and lateral margins; second and third pair of
lobes with parallel sides, mesal margin enltire, lateral margin
notched; fourth and fifth pairs of lobes pointed, broadly trian-
gular; pectinae of median, second, and third incisurae longer
than lobes, toothed only at end, those of fourth and fifth incisurae
broad, toothed also on side, blunltly pointed, arranged, 2, 2, 2, 3,
4, 5; mesospiracerores 10. — Parlatoria pseudaspidiotus Lindgr. —
Singapore on Vanda pseudaspidiotus Lindgr.
aa. Pygidium of adult female with two pairs of entire lobes, first pair
largest, broadly and bluntly rounded, second pair bluntly pointed;
distapectinae broad, filling incisurae, their dis^tal ends irregularly
toothed, some approximating latapectinae ; mesospiracerores 0-2. —
Persea on Ephedra. ephedrae Lindgr.
SPECIES OF GYMNASPIS
a. Pygidium of adult female always with true lobes, never with
pseudolobes or without lobes.
b. Pygidium of adult female with distinct pectinae.
c. Pygidium of adult female with three pairs of lobes; median
pair of lobes short and broad with lateral and mesal notches;
second pair of lobes short and oblique with prominent lateral
notch; third pair of lobes rudimentary with lateral notch;
pectinae of median incisura subequal in length to lobes,
furcapectinae, others unapectinae, longer than lobes, teeth
short; pectinae arranged 2, 2, 3, 3, those of each lateris near
third lobe; anus near caudal margin; scale dull crimson in
color. — This species has habitus of aspidiotid. — Unganda on
unknown shrub. africana Newst.
CO. Pygidium of adult female with two pairs of lobes, lobes nar-
256 THE COCCIDAE
row with subparallel sides and distinct mesal and lateral
notches; unapectinae with teeth much longer than stem of
pectina, arranged 2, 2, 8, those of each lateris approximating
plates in form and spread over entire margin; margin of
segments of preabdomen with furcapenitinae-like projections;
anus near middle and superimposed above vulva; scale bright
yellow, minute, circular, convex. — India on Mesua
spinomarginata Green.
bb. Pygidium of adult female without pectinae or plates.
c. Pygidium of adult female with two pairs of lobes.
d. Pygidium of adult female with median pair of lobes large,
oblique, mesal margins deeply serrate, placed in pygidial
incision; second pair of lobes slender, rounded, rudimentary
projections; setae of median incision large; pygidium of
second nymphal female similar, with prominent altacera-
tubae; second exuvia covering nearly entire scale, naked,
shining black, margined by thin white scale. — Paralatoria
distinctissima Newst. — Aonidia distinctissima Newst. — India
on Nerium. distinctissima Newst.
dd. Pydigium of adult female small with two large clubbed
terminal lobes with fine pointed projections on either side
of lobes, forming irregular fringe; with four irregular de-
pressed areas close to outer margin on either side of ter-
minal lobes; body almost circular, dull yellowish and finely
striate on abdominal segments; scale reddish brown, con-
vex, sometimes covered with thin incrusltation of white se-
cretion.— The figure of scale is that of aspidiotid. — New
South Wales on Acacia acaciae Frogg.
cc. Pygidium of adult female with four pairs of lobes arranged in
transverse row across end of truncated pygidium; lobes small,
subequal, entire, broader than long, distance between median
lobes twice width of lobe; plates and pectinae wanting, with
pair of short pointed projections in each incisura, sometimes
toothed like pectinae; pygidium of second nymphal female
with pygidial margin more convex than that of adult female,
with numerous pseudolobes; pygidium of first nymph with
five pairs of lobes with unapectinae between them and with
prominent altaceratubae; median incisura narrower than any
of others; scale black, cephalic half globular, broader and
larger than caudal portion. — ^Aonidia bullata Green. — Ceylon
on leaves of unidentified tree 'bullata Green.
aa. Pygidium of adult female always without true lobes, either with
pseudolobes or entire and without lobes.
b. Pygidium of adult female entire, always without projections or
pseudolobes; pygidium of second nymphal female with three pairs
of narrow, bluntly rounded, entire lobes; pectinae wanting in
median incisura, median lobes adjacent; each second and third
incisura with two distapectinae and seven or more on each lateris;
scale reddish and subhemispherical. — West Australia on Hakea.__
perpusilla Mask.
TRIBE PARLATORUNI 257
l)b. Pygidium of adult female always with numerous pseudolobes,
never entire.
c. Pygidium of adult female with comparatively broad, bluntly
rounded, crenulate pseudolobes, many of them broader than
long; pygidium of second nymphal female with four pairs of
lobes, median, second, and third pairs bluntly rounded with
mesal and lateral notches, fourth pair subtriangular with lat-
eral notch; unapectinae subequal in length to lobes, arranged
2, 2, 2, 3, 3, those of each lateris near lobe; scale semicircular,
strongly convex, exuviae large, reddish in color, — Aonidia
ebeni Leon. — Aonidia crenulate Green. — Ceylon on Diospyros,
Memecylon. crenulata Green.
cc. Pygidium of adult female with long, slender, plate-like pseudo-
lobes, twice or more as long as wide, never with their mar-
gins crenulate; anus distant from caudal margin, superim-
posed over vulva; pygidium of first nymphal female with two
pairs of lobes, median pair rounded, with lateral and mesal
notches; second pair smaller, bluntly rounded, entire or with
mesal notch; two short distapectinae in median incisura, one
in second, and two just b%ond each second lobe; scale of
adult female consisting entirely of second exuvia, strongly con-
vex, black. — England, Brazil on Aechmea aechmeae Newst.
CHAPTER XXIII
TRIBE LEUCASPIDINI
The White-shield Scales
The body of the adult female is usually longer than broad with the
cephalic and caudal ends bluntly rounded and with the lateral margins
distinct and slightly convex. The lateral margins of the segments of
the preabdomen are never provided with bracteae either in the adult
or second nymphal female. Lobes are present in the second nymphal
female, frequently represented by pseudolobes in adult female. There
are usually two, three, or four pairs of lobes, sometimes unusual in
form and approaching pseudolobes, if wanting, present in the second
nymphal female. Distapectina© are rarely present in the adult female,
frequently with furcapectinae, they may be present and typical in form
in second nymphal female. Plates frequently present in adult female,
very long with sides converging distad. Incisurae are broad and
shallow, usually filled with pectinae or plates. Densariae and par-
aphyses are never present. Altaceratubae are present and usually large,
if wanting in adult female, present in second nymphal female. Breva-
ceratbuae are usually presenit, but not numerous and not arranged in
rows. Genacerores are usually present, five groups are generally rep-
resented, there may be three or a single large group or there may be
additional groups, even as many as twelve. Mesogenacerores are usually
present, when they can be recognized as a separate group, usually con-
sist of less than eight cerores. Scale of adult female is elongate, may
be four or more times as long as broad, pyriform, or with the sides
parallel or subparallel. Exuviae are usually represented by the first
exuvia in the adult female and attached to the end of the scale. Adult
female always enclosed in a puparium, the puparium large, nearly as
long as the scale. Scale of male is elongate, subpyriform or with the
sides parallel, convex, white in color, with exuvia at one end, and never
car in ate.
The tribe Leucaspidini in common with the other tribes of
Diaspidinae, except the Fioriniini, was first recognized as a sepa-
rate subdivision by Leonardi in 1897 under the name of Leucas-
pides. A year later he gave an analytical table for the separation
of the five tribes. In this table the genus Fiorinia, the type of
the tribe Fioriniini, is combined with the Mytilaspides. A snyop-
sis of the species of Leucaspides was published in 1906 by
Leonardi. In this publication the genus Leucaspis and two new
genera, Anamaspis and Actenaspis, are recognized.
258
TRIBE LEUCASPIDINI 259
In all the species of the genus Leucaspis, so far as known^
the adult female completes its existence in a puparium. This
method of development is considered as a mark of the highest
type of specialization. The lines of modification of the pygidium
after the assumption of this method of development is apparent-
ly the same in most of the tribes. One change usually to be noted
is a modification and reduction of the groups of genacerores until
they are finally completely suppressed. An exception to this rule
is found in most of the species of this tribe. Although trans-
forming in a puparium, they have not only retained their gena-
cerores, but they are frequently represented by groups contain-
ing a considerable number of cerores. There may be the typical
number of groups, five, present or there may be additional groups
present until as many as seven additional groups can be counted
or the five groups may be combined into a single large inverted
U-shaped or crescentic mass or omnagenaceroris.
The pygidial fringe of this tribe illustrates well some of the
stages in the modification of lobes and pectinae. The lobes are
modified not only in size and changed in shape, but are lost in
the adult females of certain species. The pectinae, which are
typically distapectinae in the second nymphal females, are great-
ly elongated in the adult female of many species. They are
slender in form with converging sides with blunt ends. These
vary in form from coarsely toothed to finely toothed, to the lack
of teeth, and eventually assume the form of plates. The number
of plates is greatly reduced in certain species while in other
species the pygidium appears with projections, pseudolobes, which
are asjrmmetrical not only in form but in arrangement. Some
of the pseudolobes resemble lobes and others pectinae or plates.
The pygidium of indica is of particular interest as illustrating
an intermediate stage of modification. The genacerores are want-
ing. The lobes, of which there are three distinct pairs, have been
modified into greatly elongated clavate structures. The projec-
tions between the lobes are broad structures with their distal
ends blunt without teeth. That these are pectinae or plates un-
dergoing modification is shown by their number and arrange-
ment, 2, 2, 3, 3, which is typical for the second nymphal stage of
loewi as figured by Newstead and of the large majority of gen-
eralized pygidia of other tribes. The position of the lobes and
the number of projections, pectinae or plates, between them is
more dominant than the form of these structures although the
dominance of the form of the lobes is generally more marked
260 THE COCCIDAE
than that of the projections between them. The descriptions and
figures of certain species would suggest that this tribe contained
species transforming in a puparium that had not only retained
their genacerores but that their pygidia were provided with typi-
cal lobes and pectinae or plates. Many writers content them-
selves with studying only the pygidium of the adult female and it
is not impossible that some have figured and described the pygi-
dium of the second nymphal female as that of the adult. This
may account for the presence of primary lobes and pectinae in
the pygidium of certain adult females or it may be that the pri-
mary lobes and pectinae have not been lost. Lindinger and Green
figure and describe the pygidium not only of the adult female but
also of the second nymphal female. The structure of the
pygidium of this latter stage is more useful in determining the
tribal location and the specific limits of those species transform-
ing in puparia than the pygidium of the adult female.
The common name of The White-shield Scales is derived from
the name of the typical genus, Leucaspis, which is derived from
the Greek words meaning white and shield. The scale of the adult
female is generally light in color, often snowy white. They are
always greatly elongated with the single exuvia, the first nymphal,
at one end. This exuvia bears the remains of the antennae as
porrect structures. The scales vary in outline from those with
parallel sides to those that are pyriform. The puparium or sec-
ond exuvia is large, occupying the greater part of the scale so that
there is only a narrow thin peripheral area of excretion or wax,
which may extend as projections beyond the first exuvia and be-
yond the caudal end as described for cockerelli by Green. The
scale of the male is usually smaller, but similar in form with
a single exuvia. It is white in color, but never longitudinally
carinate as in the white male scales of many Diaspidini.
The body of the adult female is greatly elongated, and re-
sembles superfically many species of Lepidosaphes. The two ends
are bluntly rounded or subtruncated, and the sides vary from
subparallel to slightly convex. The segmentation of the thorax
and abdomen is fairly distinct. The spiracerores are usually dis-
tinct and, when they have been more carefully examined and re-
ported upon, may be found of considerable value in the separa-
tion of species. The mesospiracerores are generally present and
consist of three to twelve cerores, while the metaspiracrores are
frequently wanting. Authors in describing the spiracerores
usually do not mention the metaspiracerores when they are want-
TRIBE LEUCASPIDINI 261
ing. When present they are few in number, rarely with more
than three or four cerores. The pygidium is comparatively small,
not only in the second nymphal females but also in the adult fe-
male. The altaceratubae may be distinct in some adult females,
but so far as noted, are always present in the second nymphal
females. When the pygidia have been more carefully and ex-
tensively studied, they will probably be found characteristic of
the pygdia of all the females of the second nymphal stage. The
number of altaceratubae varies greatly and they are apparently
completely wanting in some adult females. The typical number
of lobes is difficult to determine, because in most species they are
not typical in form, but are greatly reduced and lack their charac-
teristic notches and are greatly elongated. There are usually
two or three pairs present. In the adult female the projections
between the lobes are usually greatly elongated, plate-like in gen-
eral form, but frequently with a few teeth at the distal end or in
the form of furcapectinae. Even when the lobes and pectinae or
plates are greatly modified in form, they are quite constant in
number, usually arranged 2, 2, 3, 3.
The genacerores are more frequently present than wanting
and usually consist of five groups. What is considered as the
primitive type is an arrangement into five groups, such as is found
in bambusae of Kuwana. This pygidium is generalized in many
ways, so as to suggest that the fringe, if it were not that genacer-
ores are present, that it must be that of the second nymphal fe-
male. In stricta, according to Maskell, there are three groups,
the mesogenacerores and pregenacerores form a continuous single
large crescentic group, with the two groups of postgenacerores
distinctly separated from each other and the cephalic group.
There is in pini as figured by Signoret only a single large undi-
vided crescentic group, although Leonardi states that there are
five groups which are more or less distinct among themselves and
arranged in the arc of a circle about the vulva. In cordylinidis
there is a single undivided group of about one hundred cerores.
The pygidium of signoreti shows a very different type, here the
groups of genacerores as figured by Signoret are subdivided into
several secondary groups. These groups are arranged in two
transverse rows. The cephalic row contains five subgroups and
the caudal row seven groups of which each lateral group con-
tains only about three cerores, making twelve groups in all. It
is not possible to homologize among these twelve groups, the meso-
genacerores, the pregenacerores, and the postgenacerores.
262 THE COCCIDAE
Since the adult females of all the species of this tribe are
enclosed in a puparium, it should be a simple matter to study
the pygidium of the second stage female which is that borne by
the puparium. Such a study was made by Lindinger and the re-
sults published in 1906. He figures in this paper and describes
the pygidium of the first and second nymphal stages and from
one to three types of that of the adult female. Nine species are
recognized and figured in this way and it is to be hoped that the
other species may soon receive a similar treatment.
Leonardi in his revision published in 1906, as already noted,
subdivides the tribe into three genera, Leucaspis of which pini
Hartig is the type, Anamaspis of which loewi Colvee is the type,
and Actinaspis of which pusilla Loew is the type. The last two
of these genera he named as new and each includes only its type
species. The differentiation of these genera was not well made.
These names although stated to be subgenera in his analytical table
are designated as genera elsewhere. The following is a transla-
tion of the table by Leonardi: —
GENERA OF LEXJCASPTDINI
a. Pygidium provided with pectinae.
b. Pygidium with lobes Leucaspis Targ.
bb. Pygidium without lobes Anamaspis Leon.
aa. Pygidium without pectinae. Actinaspis Leon.
Lindinger in his synopsis recognizes eleven of the twenty
described species referred to the genus Leucaspis, of which he
figures and describes nine. Four of the species previously re-
ferred to this genus he places in the genus Lepidosaphes and the
remaining ten are referred to the synonomy or considered as
unrecognizable. The classification of Leonardi is discarded as
unten table and the genus is divided into two named sections,
Salicicola and Euleucaspis, which must of necessity be recognized
as subgenera. Salicicola includes the single species kermanensis
Lindgr., which is the type. The pygidium of the adult female
of this species is very similar to that of lowei Colvee, the sulci
Newst. of Lindinger. The remaining eight species are included
in Euleucaspis, for which pini Hartig may be taken as the type.
The section Euleucaspis is divided into two other named groups,
Pusillaspis and Suturaspis, which should also of necessity be con-
sidered as subgenera. Pusillaspifr' includes two species, pusilla
Loew, which may be taken as the type, and sulci Newst. Sutur-
aspis includes the remaining six species, of which pistaceae Lindgr.
TRIBE LEUCASPIDINI 263
may be taken as the type. The following is a translatioir of the
tables of Lindinger: —
SECTIONS OF LEUCASPIS
a. Scale elongate, widened in front and not behind. Pygidium of first
and second nymphal females with comb-like toolthed plates, distinct-
ly separated from lobes. Euleucaspis Lindgr.
aa. Scale elliptical or oval. Pygidium of first and second nymphal fe-
males with lobes and plates formed like lobes Salicicola Lindgr.
SUBDIVISIONS OF EULEUCASPIS
a. First nymph with deep constriction between prothorax and meso-
thorax. Scale gradually widened Siituraspis Lindgr.
aa. First nymph not with deep constriction. Scale suddenly widened.
Pusillaspis Lindgr.
The following tentative tables include most of the described
species. They are based for the most part upon the characters
used by previous authors in describing and differentiating species.
The tables will aid in the identification of the various genera and
species. Attention has already been called to the necessity for
careful extended studies of the pygidium of the second nymphal
female. Such studies are also needed to determine whether the
lobes of the adult female are not frequently true lobes, as their
size, form, and arrangement would suggest. This conclusion is
confirmed by the characteristic number of the projections, 2, 2, 3, 3,
between these so called primary lobes.
GENERA OF LEUCASPIDINI
a. Pygidium of adult female with groups of genacerores.
b. Pygidium of first and second nymphal female with lobes,
c. Pygidium of adult female with lobes, never with pseudolobes.
d. Pygidium of adult female with lobes and distinct pectinae
or plaJtes; pygidium of first and second nymphal females
with lobes and distapectinae ; first nymph, so far as known,
with mesocoria forming distinct constriction
Leucaspis Targ.
dd. Pygidium of adult female without lobes, at most only with
lobe-like projections; pygidium of second nymphal female
with distapectinae; first nymph with mesocoria not forming
distinct constriction. Anamaspis Leon.
cc. Pygidium of adult female without lobes, pectinae, or plates,
but always with pseudolobes; first nymph with mesocoria not
forming distinct constriction Actenaspis Leon.
bb. Pygidium of first and second nymphal females with lobes, with-
out pectinae, plates represented by lobe-like projections; first
nymph with mesocoria not forming distinct constriction
Salicicola Lindgr.
264 THE COCCIDAE
aa. Pygidium of adult female without groups of genacerores
Suturaspis Lindgr.
SPECIES OF LEUCASPIS
a. Pygidium of adult female with five or less groups of genacerores,
b. Pygidium of adult female with five distinct groups of genacerores.
c. Pygidium of adult female with two pairs of lobes; the median
pair of lobes short and broad with distinct lateral and mesal
notches; second pair of lobes broad, incised, each lobelet with
lateral and mesal noitches; distapectinae short and broad, not
as long as lobes, arranged 2, 1, 3; setae long and prominent,
much longer than lobes; altaceratubae and brevaceratubae
present; anus located on line drawn through pregnacerores
and vulva on line between pregenacerores and postgenacerores;
genacerores 5-5(9-11)7-9; scale of adult female snowy-white
with parallel sides. — Japan on bamboo l>ambusae Knw.
cc. Pygidium of adult female with ithree pairs of lobes; lobes
about as broad as long, entire, bluntly rounded; plates short,
stout, triangular, arranged 2, 2, 3, 3-4; mesospiracerores about
10; genacerores in five groups, three cephalic groups some-
times contiguous, forming single group, 19-19(27-32)19-23;
caudal margin with 8 symmetrically placed large setae; sec-
ond nymphal female with two pairs of lobes, broad, deep lat-
eral and mesal notches; distapectinae broad. — Fiorinia gigas
Mask., Fiorinia morrisi (Brittin) Green. — New Zealand on
Astelia, Atherosperma, Coprosoma, Pittosporum gigas Mask.
bb. Pygidium with less than five groups of genacerores.
c. Pygidium of adult female with three groups of genacerores,
mesogenacerores and pregenacerores combined into single
crescentric group, sometimes in five groups, 9-9(15-22)10-14;
pygidium with margin fringed with long slender furcapectinae,
much longer than two pairs of short, blunt, unnotched lobes;
mesospiracerores 10; vulva located between postgenacerores
and anus located at about same distance from caudal margin.
Fiorinia stricta Mask. — New Zealand on Dendrobium, Hedy-
carpus, Phormium, Cordyline, Astelia, Hoheria.- stricta Mask.
cc. Pygidium of adult female with single large horse-shoe-shaped
group of genacerores, 200 cerores, apparently formed by com-
bination of all five groups; pygidial margin with broad mesal
series of thickened, short, sublobe-like projections; on each
side cephalad of this thickened area margin of pygidium to
line drawn through caudal margin of genacerores, bearing
short, sharp, triangular, plate-like projections, much longer
than projections of thickened area; anus and vulva alt about
same level, near transverse line drawn through caudal ends of
genacerores. — Australia on Cordyline cordylinidis Mask.
aa. Pygidium of adult female with more than five groups of gena-
cerores.
b. Pygidium of adult female with three pairs of lobes.
c. Mesospiracerores 9-11, metaspiracerores 2-3; pygidium of adult
TRIBE LEUCASPIDINI
female with three pairs of minute, pointed, triangular lobes;
pectinae and plates numerous, extending around pygidial mar-
gin, slender, bluntly pointed, at least twice as long as lobes;
median and second incisurae each with two plates, each third
incisura with 3-5; genacerores in nine groups, five primary
groups when identifiable 13-13(16-17)10-12; four groups in two
transverse rows cephalad of mesogenacerores, consisting of
3-4 and 2-3 cerores; anus superimposed over mesogenacerores
and vulva located between postgenacerores. — Leucaspis Candida
Targ. — Europe on Pinus. pini Hartig.
cc. Mesospiracerores 12, metaspiracerores 0-1; pygidium with
three pairs of minute bluntly pointed lobes; pectinae want-
ing, plates long and slender, bluntly poinited, twice or more
length of lobes; median incisura with two plates, second in-
cisura with three, third incisura with six or seven; gena-
cerores in about twelve groups arranged in two transverse
curving rows; anus superimposed over pregenacerores and
vulva on transverse line drawn through caudal part of
posltgenacerores. — Leucaspis corsa Lindgr. — Europe on Pinus. __
signoreti Targ.
bb. Pygidium of adult female with two pairs of lobes.
c. Pygidium of adult female with lobes obtusely rounded, all of
them usually entire.
d. Pygidium of adult female with median pair of lobes entire, ob-
tusely pointed lobes and second pair of small reduced denti-
form projections; plates simple, much longer than lobes;
pygidium of second nymphal female with broad pectinae;
genacerores arranged in nine groups, group of mesogena-
cerores much smaller than either of other groups, pregena-
cerores or postgenacerores, four extra groups, small, consist-
ing of only a few cerores, two groups on each side cephalad
of mesogenacerores; scale of adult female pure white,
elongate, attenuate, firslt exuvia brownish. — Algeria on
Ephedra. ephedrae Marchal.
dd. Pygidium of adult female with lobes subequal in size and
similar in form, second pair not or only slightly smaller
than median, entire; median incisura with two distapectinae,
second incisura with two or three pectinae-like projections,
and three or four on each lateris adjacent to a lobe, pec-
tinae with two to four coarse teeth; lobes two-thirds length
of pectinae; mesospiracerores 5-6; genacerores arranged in
single crescentic group of 47 cerores or in three groups,
7-9(29-29), and two small groups on each side cephalad of
the mesogenaceres ; scale of adult female white with blackish
marginal glistening first exuvia. — Leucaspis epidaurica
(Genn.) Lindgr. — France, Greece, Italy, Cyprus on Olea
riccae Targ.
cc. Pygidium of adult female with lobes distinctly pointed, never
obtusely rounded.
266 THE COCCIDAE
d. Pygidium of adult female with median pair of lobes con-
stricted on proximal portion.
e. Exuvia of second nymphal female granulose; pygidium of
adult female with median pair of lobes small, arrow-
shaped, constricted on proximal portion, sharply pointed,
entire; second pair of lobes shorter, triangular; pectinae
longer than lobes, arranged 2, 2, 2-3, blunt, simple, plate-
like projections cephalad of cephalic pectinae; mesospira-
cerores 4-5; anus located on transverse line drawn be-
tween three caudal groups of genacerores; genacerores
20-25 ( )7-8 and a supplementary group, 3-4 cerores,
on each side of two preceding segments; dorsal oracera-
tubae numerous, irregularly arranged; two transverse
series of irregularly quadrate or triangular thickened
patches on caudal portion of pygidium; sinuous series
of minuite ventral conical points extending on each side
from rostrum to pygidium; pygidium of second nymphal
female with two pairs of notched lobes; pectinae dista-
pectinae, arranged 2, 2, 3, also three bluntly rounded dis-
tant projections cephalad of pectinae. — Japan, California
on broom, apple, maple, magnolia, Paeonia.__yapomca Ckll.
68. Exuvia of second nymphal female more strongly granular;
lobes more deeply cleft; body of adult female with ven-
tral series of conical points more extended and pectinae
more deeply fimbriate. — North Australia on Ficus
japonica darwiniensis Green,
dd. Pygidium of adult female with median pair of lobes not con-
stricted on proximal portion.
e. Pygidium of adult female with two pairs of short, minute,
triangular lobes; pectinae two or three times as long as
lobes, sides subparallel, narrow, distal ends with five
short teeth; mesospiracerores 3-4, metaspiracerores want-
ing; group of 16 cerores laterad of each mesospiracle;
genacerores sometimes in five groups 11-11(15-15)11-11
and with two small groups of cerores on each side cephalad
of mesogenacerores ; anus located over mesogenacerores
and vulva between postgenacerores ; scale of adult female
white. — France on pinus affinis Leon.
ee. Pygidium of adult female with two pairs of long, point-
ed, slender lobes with distinct mesal and lateral notches
near middle of their length; pectinae deeply toothed at
distal end, constricted on proximal portion, subequal in
length to lobes, those of each lateris broader and not so
deeply toothed, two in median and second incisurae, three
cephalad of each second lobe and two to four broad
toothed pectinae laterad of others, extending upon ven-
tral surface near to and parallel with lateral margin of
body to head; genacerores in five groups and two on
each side cephalad of mesogenacerores; head with small
stout thorn-like spine cephalad and lalterad of each an-
tenna; pygidium of second nymphal female with two
TRIBE LEUCASPIDINI 267
pairs of lobes with prominent mesal and lateral notches
and with broad deeply toothed distapectinae in incisura
and on each lateris; scale of adult female narrow, taper-
ing to blunt point at each end, excretory portion limited,
extending cephalad of first exuvia as blunt projection
and knob-like constricted area at caudal end. — Fiorinia
cockerelli Charmoy. — Mauritius, Ceylon on Draecaena,
Priitchardia. cockerelli Charmoy.
SPECIES OF ANAMASPIS
Pygidium of adult female with two pairs of small, short, rounded lobes
and third pair frequently wanting and when present asymmetrically
placed or all lobes not sharply defined; genacerores arranged in form
of crescent, sometimes in five groups, arranged 5-10(10-14)9-13; meso-
spiracerores 2-5; anus placed over mesogenacerores and vulva between
postgenacerores ; pygidium of second nymphal female with two dista-
pecitinae in median incisura and in other incisurae. — Fiorinia sulci
Newst. — Europe on Pinus loewi Colvee.
SPECIES OF ACTENASPIS
Pygidium of adult female fringed with numerous long asymmetrically
arranged projections or pseudolobes, distal ends frequently more or
less dilated and finely toothed; genacerores arranged in shape of
crescent, sometimes divisible into five groups, 9-9(10-11)6-10; meso-
spiracerores 0-7; pygidium of second nymphal female with two pairs
of broad truncated lobes with deep mesal and lateral notches; median
and second incisurae each with two distapectinae and four or more
of each lateris adjacent to each second lobe; altaceratubae prominent,
9-11, one mesal and 4-5 on each side. — Leucaspis leonardi (Ckll.)
Lindgr. — Europe, Canary Islands," Algeria, Syria on Pinus
pusilla Loew.
SPECIES OF SALICICOLA
Pygidium of adult female with lobes, plates or pectinae, sometimes with
margin with slight irregular projections, these sometimes forming five
pairs of lobe-like projections; genacerores in single crescentic group
of 28 to 40 or in three distinct groups; mesospiracerores 2; pygidium
of second nymphal female with two pairs of lobes, median pair broad,
subtruncated, rounded, entire: second pair of lobes much smaller, tri-
angular, bluntly pointed; median incisura without pectinae or plates,
second incisurae and each lateris with two short, blunt, lobe-like pro-
jections or plates. — Persia on Salix kermanensis Lindgr.
SPECIES OF SUTURASPIS
a. Pygidium of adult female with three pairs of long spatulate bluntly
rounded lobes, three or four times as long as broad; plates broad
and blunt, about one-half as long as lobes, distal end rounded, ar-
ranged 2, 2, 3, 3. — Leucaspis indica Marl. — India on mango
indica Marl,
aa. Pygidium of adult female never with long spatulate lobes.
b. Pygidium of adult female with two distant pectinae between
median pair of lobes, median pair of lobes distant, rounded, with
268 THE COCCIDAE
slight lateral notch; second and third pairs of lobes successively
smaller and each incised; second and third incisurae with pecti-
nae and each lateris with prominent latadentes. — Leucaspis
kelloggi Golem. — California on Pseudotsuga, Abies
kelloggi Golem,
bb. Pygidium of adult female without pectinae or plates.
0. Pygidium of adult female with lateres entire, with single pair
of broad, very short, rounded, lobe-like projections; second
nymphal female with two pairs of distinct lobes with lateral
and mesal notches; plates short and blunt, shorter than lobes,
two in median incisura and one in each second incisura;
mesogenacerores three or none, usually none; mesocoria of first
■ nymph distinctly transversely constricted. — Leucaspis pistaeae
Lindgr. — Cyprus on Pistacia pistaeiae Lindgr.
cc. Pygidium of adult female with lateres crenulalte, not with
distinct lobe-like projections; second nymphal female with
two pairs of distinct lobes, broad and bluntly rounded, entire;
pectinae broad, two in median incisura and two on each side
of each second lobe; mesogenacerores wanting, — Leucaspis
indiae-orientalis Lindgr. — India on Pinus
J indiae-orientalis Lindgr.
CHAPTER XXIV
TRIBE L.EPIDOSAPHINI
The Oyster-shell Scales
The body of the adult female is always distinctly longer than broad
with the cephalic and caudal ends bluntly rounded. The lateral mar-
gins are parallel or subparallel usualy with distinct indentations mark-
ing the position of the sutures between the thoracic and abdominal seg-
ments. The lateral margins of the segments of the preabdomen are
never provided with bracteae either in the adult female or second
nymphal female. Lobes are usually present in the adult female, usually
three pairs, more or less than this number may be present. Plaltes are
rarely if ever absent, pectinae are rarely present and are then only fur-
capectinae. Incisurae are narrow and the space is usually filled with
plates. The median incisura is typically provided with plates, rarely
without. Densariae are never present and paraphyses only rarely. Alta-
ceratubae, are rarely, if ever absent, usually arranged in pairs, their
oraceratubae oval in outline, often twice as long as broad. Brevacera-
tubae are usually present, frequently numerous, sometimes arranged in
transverse rows. Genacerores are usually present, arranged in five, four,
or two groups. The mesogenacerores, when presenit, usually consist of
more than eight cerores. Scale of adult female is coriaceous or horny,
usually brownish or blackish in color, sometimes white or pale, quite
typical in form, usually much longer than broad, from two to five times,
frequently shaped like an oyster-shell, one margin convex and the other
concave with the surface transversely ridged, sometimes with sides
straight and subparallel. One end is much smaller than the other, small
end bearing two exuviae. The female sometimes transforms in a pupa-
rium and the pygidium of the adult female is provided with pseudolobes.
The scale of the male is similar in form to that of the female, elongate
in form, with a single exuvia at one end.
The tribe Lepidosaphini, as recognized by Leonardi in 1898
contained four genera, one of which was divided into two sub-
genera. This number was increased to eight in his monograph
of the Mytilaspides published in 1903. In the following pages
twenty-eight genera are recognized.
The scale of the adult female is quite constant in form, typi*
cally mussel-shaped or oyster-shell-shaped or pyriform in general
appearance. In certain species there is a departure from this
form, and the two sides or margins of the scale may be straight
and parallel or nearly so. The scales of the female are typically
269
270 THE COCCIDAE
dark in color, blackish or brownish, though in certain species the
color is much lighter, varying from light brown to yellow or
olivaceous. While in certain species the sides are nearly parallel,
in most of the species the sides converge toward one end, the
cephalic end, which bears the two exuviae. The exuviae
overlap each other. The exuvia of the first nymph bears
the remains of the antennae as porrect projections, while the
exuvia of the ;feecond nymphal female is much larger and fre-
quently covers about one-fourth of the outer surface of the scale.
The ventral scale is always complete and continuous with the
dorsal scale. In certain species the central part of the ventral
scale is always thinner, the extent and position of the thinner por-
tion is definite for each species and this thinner portion is rup-
tured when the scale is removed from the host-plant. In those
species where the ventral scale is thicker and tougher, it is not
ruptured when the scale is removed. This results in those species
with a thin area of the ventral scale as being described as having
a divided ventral scale and the others as having the ventral scale
entire or complete. Where the ventral scale is very thin and all
of it remains attached to the host-plant, it may be described as
obsolete or wanting. The wax of the dorsal scale is usually com-
paratively thick and tough and not easily broken.
The scale of the male is elongate, about three times as long
as broad with the single exuvia attached at the cephalic end. The
sides of the scale are usually nearly parallel and in general the
caudal portion is not much wider, as in the second nymphal scale
and adult female scale of many species. The male scale is from
one-half to one-third the length of that of the adult female. They
are quite similar in color and general form to those of the female.
There is frequently a thinner transverse band on the caudal part
of the scale which serves as a hinge and allows the caudal end to
be raised so as to permit of the escape of the adult male.
The species of this group are quite generally designated as
the oyster-shell scales, because of their similarity in shape, color,
and general appearance to the shell of an oyster. Unfortunately
this resemblance is also found in a few species of Diaspidini, but
the name of the oyster-shell scale has been so long applied to the
common scale of the apple and to other species of the tribe that
there is no hopes now of introducing a different name and it would
be unwise to attempt it.
The body of the adult female is placed at the cephalic end
of the scale under the exuviae and the eggs with the pulverulent
TRIBE LEPIDOSAPHINI 271
wax mixed with them are located under the caudal portion of the
scale. In the adult female the body is elongate, typically about twice
as long as broad, but in Ischnaspis it may be over four times as
long as broad. The cephalic end is usually truncate or bluntly
rounded. The antennae are represented by unsegmented tuber-
cles, each bearing one or two long setae. The segmentation of the
body is ordinarily distinguishable, but obscure. In Opuntiaspis
and Mytilella the mesocoria is deep, particularly at each lateral
margin, and the body appears to be divided into two portions.
The segmentation of the preabdomen is usually distinct and
sharply marked, particularly at each lateral margin. The caudo-
lateral angles of these segments are frequently produced as
rounded lobes and in some cases the lobes are prolonged and their
lateral margins are strongly dentate so that each resembles a claw.
The division between the preabdomen and pygidium is gen-
erally distinct. The pygidial fringe is usually quite typical in
form. There are generally two or three pairs of lobes, although
there may be only a single pair, and in rare cases none at all. In
a few species there are four pairs of lobes present, but the fourth
pair is small and rudimentary. One or more of the pairs of lobes
cephalad of the median pair is frequently deeply incised or com-
pletely divided. In the tables it is simply said to be incised.
The second pair of lobes is usually incised only once, forming
two lobelets, but the third pair while often incised only once may
be incised several times in certain species. Each of these lobe-
lets are frequently incorrectly designated and numbered as dis-
tinct lobes. Some species with only three pairs of lobes are said
in such cases to have five or seven pairs. The margins of the
lobes are usually notched, particularly the median lobes, and
usually on both the mesal and lateral margins. The number of
notches may be reduced in size and increased in number until
their margins are crenulate or finely serrate.
Pectinae, if present, are of the type of furcapectinae and
even such reduced pectinae are of rare occurrence. The plates
are long, slender, and pointed. They are usually longer than the
lobes. The number varies somewhat, but is usually 2, 2, 2, 2 or
2, 2, 2-3, 6. The pair of plates in the median incisura are rarely
absent and some of the species referred here and lacking them, with
more study, will probably be found to belong to the Diaspidini. The
greatest variation in the number of plates is on the lateres which
may bear only a few or a considerable number. This region is
272 THE COCCIDAE
usually typical in form, rarely provided with densariae but may
be finely serrate.
The altaceratubae are usually large, with long tubes, their
oraceratubae generally oblique to the tubes and distinctly longer
than wide. In the lateres the altaceratubae are usually arranged
in pairs and their oraceratubae open near together. The laraa-
ceratubae are long and comparatively large. Their tubes can be
traced through the plates to their oraceratubae located in the dis-
tal end. The brevaceratubae are usually numerous and are gen-
erally arranged in crescentic, often interrupted rows. While
there is variation in the length of the tubes of the brevaceratubae,
their tubes are usually longer than those of the brevaceratubae
found in the Diaspidini.
The anal opening is probably more often located near the
cephalic end of the pygidium than near the middle or caudal end.
The vulva is located near the middle of the pygidium. It is sur-
rounded by four or five groups of genacerores in fifteen of the
genera and wanting in the other nine. The genacerores are pres-
ent in much the greater number of species, since they are present
in all the species of the typical genus Lepidosaphes, which con-
tains nearly as many species as all the other genera together.
The number of species with four groups of genacerores is small,
less than ten per cent, and only rarely are the cerores of the
groups combined so that it is not possible to identify their limits.
Much use has been made in the preparation of the follow-
ing tables of the monograph of the Mytilaspides by Leonard!.
The species distributed among the different genera are confined to
two, Lepidosaphes and Ischnaspis, in Fernald's Catalogue and its
supplements. The structures of the pygidium described in the
tables are in every case those of the adult female unless other-
wise specified.
The following species have heen omitted from the tables for various
reasons: — abietis Sign, from Europe on Abies; ceratoniae Genu, from
Isle of Cyprus on Ceratonia; flava Targ. from Europe, Australia, China,
Hawaiian Islands, and North America on olive; juglandis Sign, from
France, Germany on Juglans, this is apparently the same as ulmi
Linn.; myrthi Bouch6 from southern Europe on Myrtus, this is
the same as ulmi Linn.; and Mytilaspis (Coccomytilus) dispar Vays.
from Madagascar on Manihot. The following species are all from New
South Wales: chitinosa Frogg. on Templetonia, cortrioides Frogg. on
Acacia, crassa Frogg. on Melaleuca, eucalypiti Frogg. on Eucalyptus,
lobulatus Frogg. on Casuarina, mulgae Frogg. on Acacia, and recurvata
Frogg. on Acacia. Two species, greeni Charmoy and hibisci Charmoy,
are from the Island of Maritius.
TRIBE LEPIDOSAPHINI 273
GENERA OF LEPIDOSAPHINI _
Pygldium of adult female with groups of generacerores.
b. Pygidium of adult female with lobes arranged in pairs, median
pair never fused into single projection.
c. Pygidium of adult female always with plates cephalad of me-
dian pair of lobes,
d. Pygidium of adult female never with paraphyses.
e. Adult female with mesothorax and metathorax never
separated by deep transverse constriction.
f. Pygidium of adult female without lattice-shaped thick-
enings; usually with four or five large distinct groups
of genacerores; preabdomen and pygidium frequently
with numerous brevaceratubae.
g. Pygidium of adult female with four pairs of lobes;
second, third, and fourth pairs incised; median in-
cisura without plates; scale of adult female with
dense white cottony or flutty covering of wax
Trichomytilus Leon,
gg. Pygidium of adult female never with more than three
pairs of lobes; second and third pairs sometimes in-
cised; median incisura with or without plates; scale
of adult female naked, not with dense white cottony
or fluffy covering of wax.
h. Pygidium of adult female with median pair of
lobes rarely adjacent, if so, only at proximal ends
and not forming single semicircular area.
1. Adult female with pygidium and segments of
preabdomen with distinct, irregular, subtrans-
verse rows of oraceratubae, outlets of breva-
ceratubae; pygidium apparently always with
altaceratubae.
j. Pygidium of adult female always with
plates in median incisura.
k. Pygidium of adult female always with
three pairs of lobes. — Type, Lepidosaphes
bicornis G. & L Triaspidis MacG.
kk. Pygidium of adult female always with
less than three pairs of lobes.
1. Pygidium of adult female always with
two distindt pairs of lobes.
m. Adult female with cephalic end of
body never provided with strongly
chitinized tooth-like projections.
n. Preabdomen with lateral margins
of its segments never produced into
claw-like processes.
0. Adult female with cephalic end
of Ithe body never with a trans-
274 THE COCCIDAE
verse constriction cephalad of
antennae,
p. Adult female with cephalo-lat-
eral angles of body never pro-
duced into ringed horn-like
projections; body always nor-
mal in form
Lepidosaphes Shimer.
pp. Adult female with oephalo-lat-
eral angles of body always
produced into ringed horn-
like projections. — Type, Myti-
laspis ocellata Green
Cornuaspis Mac.
00. Adult female with cephalic end
of body always with transverse
constriction cephalad of antennae,
separating off head-like portion.
— Type, Mytilaspis occuli Green.
Cephalaspis MacG.
nn. Preabdomen with lateral margins
of its segments always produced
into prominent claw-like processes.
— Type, Lepidosaphes ungulata
Green. Ungulaspis MacG.
mm. Adult female with cephalic end of
body always provided with distinct
numerous sitrongly chitinized tooth-
like projections. — Type, Mytilaspis ser-
rifrons Leon ScoMnaspis MacG.
11. Pygidium of adult female always with
single pair of distinct lobes. — Type,
Mytilaspis wilga Leon
Leonardaspis MacG.
jj- Pygidium of adult female always without
plates in median incisura.
k. Pygidium of adult female never with
prominent pygidial incision; median pair
of lobes with proximal ends not adjacent.
1. Pygidium of adult female always with
more than one pair of lobes. — Type,
Mytilaspis intermedia Mask
Scrupulaspis MacG.
11. Pygidium of adult female never with
more than one pair of lobes. — Type,
Mytilaspis spinifera Mask
Berlesaspis MacG.
kk. Pygidium of adult female always with
prominent pygidial incision; median pair
of lobes with their proximal ends united.
TRIBE LEPIDOSAPHINI 275
1. Pygidium of adult female with three
pairs of lobes. — Type, Mytilaspis pliym-
atodidis Mask. Fusilaspis MacG.
11. Pygidium of adult female with two pairs
of lobes. — Type, Mytilaspis nivea Mask.__
Albastaspis MacG.
ii. Adult female with pygidium and segments of
preabdomen without subtransverse rows of ora-
ceratubae, forming outlelts of brevaceratubae ;
pygidium always with altaceratubae
Pinnaspis Ckll.
hh. Pygidium of adult female with median pair of
lobes adjacent, two lobes together forming single
semicircular area. — Type, Lepidosaphes hemichion-
aspiformis Green. Hemiaspis MacG.
ff. Pygidium of adult female with lattice-shaped thicken-
ings and with ithree small indistinct groups of gena-
cerores; preabdomen and pygidium without or with,
only few brevaceratubae Ischnaspis Doug.
ee. Adult female with mesothorax and metathorax separated
by deep transverse constriction; pygidium with two pairs
of lobes, second pair incised; median incisura with two
plates. Mytilella Leon.
dd. Pygidium of adult female with mesal pair of club-shaped
paraphyses and with single pair of lobes. — Type, Lepidosaphes
hawaiiensis Mask. Andaspis MacG.
cc. Pygidium of adult female always without plates cephalad of
median pair of lobes, with three pairs of lobes, second pair in-
cised; plates of median incisura very small
Aonidomytilus Leon.
bb. Pygidium of adult female with median pair of lobes fused into
single projection; sometimes with more than five groups of gena-
cerores. — Type, Mytilaspis uniloba Knw Lepidaspidis MacG.
. Pygidium of adult female without groups of genacerores.
b. Pygidium of adult female always with at least one pair of lobes,
c. Pygidium of adult female always with plates,
d. Pygidium of adult female never with lattice-shaped thicken-
ings.
e. Adult female with mesothorax and metathorax never sepa-
rated by deep transverse constriction.
f. Pygidium of adult female with lateral margins broad-
ly convexly convergent caudad, with caudal end con-
vexly rounded, never truncate, with two, rarely three,
pairs of lobes; median pair of lobes always different
in form and size from the others; scale of male and
female not with prominent transverse constrictions,
g. Body with metacoria located near middle of its
length; portion of body cephalad of meitacoria usually
276 THE COCCIDAE
only slightly longer than portion caudad of it, al-
ways less than twice as long. Coccomytilus Leon.
gg. Body with metacoria located distinctly caudad of
middle of its length; portion of body cephalad of
meltacoria more than twice as long as portion caudad
of it. — Type, Dinaspis ichesii Leon Dinaspis Leon.
ff. Pygidium of adult female with caudal end broadly
truncate and bearing three pairs of pointed, triangu-
lar, equidistant, similar, subequal lobes; scale of male
and female with six or more prominent transverse con-
strictions. Allantomytilus Leon.
ee. Adulit female with mesothorax and metathorax separated
by deep transverse constriction; pygidium with three pairs
of lobes. Optuntiaspis Ckll.
dd. Pygidium of adult female always with distinct lattice-shaped
thickenings; brevaceratubae few in number or wanting. —
Type, Ischnaspis spathulata Lindgr Parischnaspis MacG.
CO. Pygidium of adulit female without plates.
d. Pygidium with median pair of lobes at most never more than
twice width of median lobe apart.
e. Pygidium with two parirs of distinct lobes, median pair
as large as second pair; preabdomen with caudo-laiteral
angles of segments with strong finger-like processes. —
Type, Mytilaspis argentata Ckll Niveaspis MacG.
ee. Pygidium with two pairs of lobes, median pair minute^
indistinct, not more than one-sixth width of second lobe;
preabdomen with caudo-lateral angles not with finger-like
processes. — Pernaldella Leon. — Type, Mytilaspis indentata
Green. Fernaldanna MacG.
dd. Pygidium with median pair of lobes four or more times width
of median lobe apart; median and second pairs of lobes
minute. — Type, Mytilaspis beyeriae Green
Maskellanna MacG.
bb. Pygidium of adult female without lobes or plates, with four stout,
equidistanlt, conical projections extending from surface of pygi-
dium beyond its margin. Phaulomytilus Leon.
SPECIES OF TRICHOMYTILUS
Pygidium with median pair of lobes prominent, bluntly rounded, longer
than broad, entire, distant: second pair of lobes large, incised, mesal
lobelet much smaller than lateral, bluntly pointed, entire; third pair
of lobes broad, deeply and broadly incised, lobelets bluntly pointed,
subequal, entire; fourth pair of lobes deeply incised, similar in size
and form to third pair; plates arranged 0, 3, 3, 3, 2, long and slender,
much longer than lobes; genacerores 5-6(13-18)19-24. — Australia on
Eucalypitus. formosus Mask.
SPECIES OF TRIASPIDIS
a. Pygidium with second pair of lobes not incised, at most only notched,
b. Pygidium with furcapectinae between lobes.
TRIBE LEPIDOSAPHINI 277
c. Pygidium with second and third pairs of lobes denticulate at
distal end; median pair of lobes obliquely truncate at disltal
end; lateres bearing f urcapectinae ; genacerores 8-11(20-22)20-
22. — Australia on Banksia. 'banksiae Mask.
cc. Pygidium with lobes broadly rounded at distal end and notched
on mesal and lateral margins; lateres bearing plates; gena-
cerores in five confused groups, 13-13(20-22)16-18. — Pinnaspis
nitidus Brittin (Green.) — New Zealand on Dysoxylum, Ather-
osperma, Coprosoma, Rhipogonium, Pinus. pyriformis Mask.
bb. Pygidium with plates, never with f urcapectinae between lobes;
lobes notched or coarsely crenulate; second and third pairs much
smaller than first; plates large and prominent, arranged 2, 2, 3,
4-6; genacerores 14-14(23-31)21-21. — New Zealand on Leptosper-
mum. leptospermi Mask.
,. Pygidium with second pair of lobes incised.
b. Pygidium with third pair of lobes incised, at most only deeply
notched on lateral margin.
c. Pygidium with each second and third incisura with two
plates, also median incisura, and four on each lateris arranged
in two groups of two each; median pair of lobes deeply
notched on lateral margin, second and third pairs faintly
notched; genacerores 9-11(19-26)18-25; mesospiracerores 9-14. —
Australia on Eucalyptus. Mcornis Green & Lidg.
cc. Pygidium with each second and third incisura with single
plate.
d. Pygidium with two hair-like plates between median pair of
lobes; plates arranged 2, 1, 1, 6, those of each lateris ar-
ranged in three equidistant groups of two each; median pair
of lobes with two small lateral notches, second with single
deep lateral notch, and third with two small notches; gena-
cerores 7-7(10-12)10-12, five groups adjacenit!. — Lepidosaphes
fulleri Fern. — Australia on Banksia elongata Fuller.
dd. Pygidium with two plates between median pair of lobes as
long as other plates and longer than median pair of lobes;
plates arranged 2, 1, 1, 2, those of each lalteris situated, one
near third lobe and other near middle of lateris; median
pair of lobes oblique, large, finely serrate, set in emargina-
tion; second pair of lobes large, truncately rounded, finely
serrate, mesal lobelet as large or larger than median lobe;
third pair of lobes comparatively large, truncately rounded,
finely serrate, subequal ;to lateral loblet of second pair;
genacerores 0-7(7-7)3-3. — South Africa on Loranthus
kamerunensis Lindgr.
bb. Pygidium with third pair of lobes incised.
c. Pygidium with third pair of lobes not consisting of more than
three lobelets.
d. Pygidium with each third lobe divided into two lobelets.
e. Pygidium witli median pair of lobes with one notch on
mesal and two on lateral margin.
278 THE COCCIDAE
f. Pygidium with median pair of lobes large; second pair
of lobes incised, mesal lobelet with notch on lateral
margin; third pair of lobes with mesal lobelet pointed
and smaller than lateral; plates arranged 2, 1, 1, 3,
always simple, those of median incisura hardly longer
than lobes and shorter than other plaites; genacerores
27-27(28-35)31-33.— Mexico on Ortica mexicana Ckll.
ff. Pygidium with median pair of lobes large and promi-
nent; second pair of lobes small, incised, two lobelets
subequal; third pair of lobes small, deeply incised,
mesal lobelet twice as broad as lateral, lateral lobelet
bluntly pointed; plates arranged 2, 2, 2, 2, those of me-
dian incisura small, shorter than median pair of lobes,
other plates long and stout, those of each lateris near
third lobe; genacerores 5-5(7-10)7-9. — Mexico on Chil-
opsis. chilopsidis Marl.
ee. Pygidium with median pair of lobes large, entire or finely
serrate or slightly crenulate, not notched.
f. Pygidium with single plate in each second incisura.
i g. Pygidium with genacerores 7-7(12-13)10-11; median
pair of lobes large, pointed, finely serrate, distant;
second pair of lobes incised, lobelets bluntly rounded,
finely serrate, mesal lobelet largest; third pair of
lobes smaller than second, similar in form, mesal
lobelet largest; plates arranged 2, 1, 1, 2, those of
each lateris situated, one near third lobe and other
slightly beyond middle; second and third incisurae
prolonged as pointed lobe-like structures bearing Jra-
ceratubae. — South Africa on Macrolobium
meridionalis Lindgr.
gg. Pygidium with genacerores 2-2(5-6)3-3; median pair
of lobes large, bluntly rounded, oblique, serrate; sec-
ond pair of lobes incised, lobelets bluntly rounded,
almost entire, mesal lobelet longer and broader than
lateral; third pair of lobes larger than usual,
smaller than second pair, deeply incised, bluntly
rounded, subentire; plates 2, 1, 1, 2, those of each
lateris situated one near third lobe and other near
cephalic end; median incisura deeply hollowed out,
its plates subequal in length to median lobes; second
and third incisurae with blunt lobe-like projections
bearing oracera-tubae. — South Africa on Crudia
crudiae Lindgr.
ff. Pygidium without plates in median incisura.
g. Pygidium with median pair of lobes distant, at least
not fused at proximal end.
h. Pygidium with genacerores 9-9(16-18)9-10; median
pair of lobes large, entire or slightly crenulate;
second pair of lobes incised, mesal lobelet larger
than laiteral; third pair of lobes broad and very
TRIBE LEPIDOSAPHINI 279
short, deeply incised, lateral lobelet usually mi-
nutely crenulate; plates arranged 2, 0, 2, 5, those
of each lateris in two groups, caudal group of one
or two near third pair of lobes and cephalic group
of two or three some distance cephalad of caudal
group; lateral margin of three caudal segments
of the preabdomen with three plates each. — Mexi-
co on Mimosa. mimosarum Ckll.
hh. Pygidium with genacerores 4-4(6-7)3-4; median
pair of lobes pointed, finely serrate, distant; sec-
ond pair of lobes incised, lobelets bluntly round-
ed, subentire, mesal lobelet larger than lateral,
laiteral different in shape but subequal in size to
median lobe; third pair of lobes large, deeply
incised, subentire, lobelets bluntly rounded, mesal
lobelet much larger than lateral; plates arranged
2, 0, 1, 3, those of each lateris situated, one near
third lobe, one near middle projection, and other
at fcephalic end; plates of median incisura as long
as or longer than median lobes; second and third
incisurae with blunt lobe-like projections bearing
oraceratubae ; each lateris finely serrate with lata-
dentes. — South Africa on Cynometra
uherrans Lindgr.
gg. Pygidium with median pair of lobes fused at proxi-
mal end, the median incisura represented by narrow
deep indentation, distal ends bluntly rounded, en-
tire; second pair of lobes small, incised, lobelets
adjacent to each other and median lobe; third pair
of lobes deeply incised, tooth-like projections;
plates arranged 2, 0, 2, 6, those of each lateris in
three groups, group of two near third lobe, three
equidistant near middle, and one at cephalic end;
plates of median incisura short and slender, much
smaller than other plaJtes; postgenacerores four,
others not determined. — India on Aglaia
travancorensis Lindgr.
dd. Pygidium with each third lobe divided into three lobelets,
median portion largest and lateral portion notched, not
pointed; median pair of lobes notched on lateral and mesal
margins; mesal lobelet of second pair notched on lateral
margin; plates arranged 2, 1, 1, 2, all large and subequal
in size, much longer than lobes, those cephalad of second
pair of lobes sometimes furcapectinae; two plates of each
lateris distant; genacerores 7-8(10-15)14-19. — Brazil on
Baccharis. perlonga Ckll.
cc. Pygidium with third pair of lobes consisting of considerable
number of minute lobelets, always more than five; median pair
of lobes large, rounded, entire; second pair of lobes incised,
two lobelets entire and subequal; plates arranged 2, 2, 1, 2,
those of median and second incisurae short, not at all or only
280 THE COCCIDAE
slightly longer than lobes adjacent to them, plates of each
laterls distant; genacerores 7-7(11-12)8-14. — Australia on Eu-
calyptus. lidgetti Ckll.
SPECIES OF LEPIDOSAPHES
a. Pygidium with second pair of lobes not incised or at most only
deeply notched,
b. Pygidium with median pair of lobes with their proximal ends
distant; plates of median incisura always much longer than
median pair of lobes.
c. Pygidium with median pair of lobes deeply notched on mesal
and lateral margins; second pair of lobes smaller, notched on
mesal and lateral margins, blunitly pointed; plates arranged
2, 2, 3, large and broad, much longer than lobes, three plates
of each lateris near second lobe; lateris deeply incised and
finely serrate; genacerores 3-3(5-8)3-6. — Australia on Mela-
leuca. melaleucae Mask.
cc. Pygidium with median pair of lobes not deeply notched, finely
serrate or entire.
d. Pygidium with median pair of lobes broad, broadly rounded,
finely serrate; second pair of lobes minute, inconspicuous;
plates arranged 2, 2, 2, those of each lateris adjacent near
second lobe; plates long and slender, distinctly longer than
lobes; genacerores 3-3(7-10)7-9. — Japan on Miscanthus
arii Knw.
dd. Pygidium with median pair of lobes subquadrangular, en-
tire; second pair of lobes similar in shape and smaller;
plates arranged 2, 2, 9, long and slender, much longer than
the lobes, those of each lateris arranged in groups of three;
lateres entire, not deeply incised; genacerores 1-7(15-15)15-22.
— Algeria on Ampelodesma. ampelodesmae News.t.
bb. Pygidium with median pair of lobes with their proximal ends
adjacent.
c. Pygidium with four groups of genacerores, (3-3)4-4; median
pair of lobes longest on mesal half, lateral margin longer than
mesal, both sloping and finely serrate; second pair of lobes
much smaller and finger-shaped; plates arranged 2, 1, 1-3, those
of median incisura small, those of each lateris much larger
than others. — Japan on Calicornia,. Elaeagnus, Quercus
cravHi Ckll.
cc. Pygidium with five groups of genacerores, 2-3(8-11)5-6. — This
was described as variety of crawil Ckll. The figures of
Maskell and Leonardi and description of Cockerell are more
different than is implied by description by Maskell. Japan on
Quercus. canaliculata Mask.
aa. Pygidium with second pair of lobes incised.
b. Pygidium with proximal ends of median pair of lobes distinctly
separated.
c. Pygidium with only single plate or without plates in each
second incisura.
TRffiE LEPIDOSAPHINI 281
d. Pygidium with single plate in each second incisura.
e. Pygidium. with furcapectinae on each lateris; plates and
pectinae arranged 2, 1, 6, those of each lateris situated,
two plates adjacenit to each second lobe, four pectinae in
two groups of two each equidistant from each other and
caudal group of plates; median pair of lobes small, round-
ed, notched; second pair of lobes rounded, incised, lobe-
lets entire or finely notched; genacerores 4-4(6-7)4-5. —
Ceylon on Lasianthus. lasianthi Green.
ee. Pygidium with plates on each lateris, never with furca-
pectinae.
f. Pygidium with plates located on each lateris at caudal
end, at cephalic end, and near middle, one of groups
sometimes wanting.
g. Pygidium with genacerores 6-6(10-12)8-10; median
pair of lobes not especially large, rounded at ends,
with minute la;teral lobelets; second pair of lobes in-
cised, mesal lobelet larger than lateral; plates ar-
ranged 2, 1, 3, those of each lateris situated, one
near each second lobe, one near middle, and one at
cephalic end. — Mexico on "Laurel-tree", Yucca,
Agave. nigra Ckll.
gS- Pygidium with genacerores 3-4(6-6)4-4.
h. Pygidium with median pair of lobes oblique, fair-
ly large, bluntly rounded, margins finely serrate;
second pair of lobes large, incised, each mesal
lobelet larger than median lobe, bluntly rounded,
margin finely serrate; plates arranged 2, 1, 3,
those of each lateris large and broad, situated one
near each second lobe, one near middle, and one
at cephalic end; plates of median and second in-
cisurae large, as long as or longer than lobes. —
South Africa on Macrolobium tenuior Lindgr.
hh. Pygidium with median pair of lobes broadly sepa-
rated, not oblique, margins with mesal and lat-
eral notches, with a median pointed process;
second pair of lobes incised, mesal lobelet much
larger than lateral and one-third width of median
lobe; plates arranged 2, 1-0, 4, those of each
lateris large and broad, in two groups of two
plates each, one group near cephalic end and
other midway between cephalic group and second
lobe; plates of median incisura slender, longer
than lobes; each second incisura with blunit pro-
jection bearing an oraceratuba; anus located ce-
phalad of mesogenacerores. — Ceylon on Hevea.
fasciata Green.
ff. Pygidium with plates on each lateris at caudal end
only; plates arranged 2, 1, 1-2, long, as long as lobes,
broad at proximal end; median pair of lobes large.
282 THE COCCIDAE
symmetrical, broadly spatulate, indistinctly three-
notched; second pair of lobes similar, much smaller,
unsymmetrical, incised, lateral lobelet longer than
broad; genacerores 3-3(7-8)7-8. — Turkey in Asia on
Juniperus. juniperi Lindgr.
dd. Pygidium without plates in second incisurae.
6. Pygidium with median pair of lobes broad, broadly round-
ed, notched on lateral and mesal margins; second pair
of lobes small, incised, mesal lobelet larger than lateral;
plates arranged 2, 0, 4, those of each lateris arranged in
two groups of two each, one near middle, other at the
cephalic end; median incisura with pointed projection,
second incisurae with blunt projection bearing oracera-
-tuba and seta; genacerores 3-4(6-6)4-4. — Philippine Is-
lands, Ceylon on Eugenia, Hevea rubrovitatus Ckll.
ee. Pygidium with median pair of lobes bluntly pointed, en-
tire or inconspicuously notched, distant; second pair of
lobes incised, mesal lobelet much larger than lateral;
plates arranged 2, 0, 3, those of each lateris placed, one
near each second lobe, one near middle, other at cephalic
end; median incisura with spine-like projection between
plates, each second incisura with lobe-like projection bear-
ing oraceratuba; genacerores 4-7(6-7)4-4. — Seychelles
auriculata Green.
cc. Pygidium with two plates in each second incisura.
d. Pygidium with median pair of lobes distinctly notched or
coarsely crenulate on disltal portion, never entire.
e. Pygidium never with more than four genacerores, nine
pregenacerores, and nine postgenacerores, usually within
following limits: 2-5(3-9)3-9.
f. Pygidium with median pair of lobes broader than long.
g. Pygidium with plates of median incisura subequal
in length or shorter than median lobes; median pair
of lobes broad and rounded, lateral margins usually
notched, widely separated; second pair of lobes in-
cised, entire; plaites arranged 2, 2, 4-6, those of sec-
ond incisurae long and slender, twice as long as
median lobes; genacerores 2-5(3-6)3-6. — Algeria on
Ficus. minima Newst.
gg. Pygdium with plates of median incisura always dis-
tinctly longer than median lobes.
h. Preabdomen with lateral margins of its caudal
segment with four plates on each side; pygidium
with median pair of lobes broad, subtruncate,
notched on mesal and lateral margins; second
pair of lobes incised, margins subtruncate; plates
arranged 2, 0, 6, those of each lateris arranged
in three equidistant groups of two each, one near
second lobe, one near middle, and one at cephalic
TRIBE LEPIDOSAPHINI 283
end; lateris irregular, dentate; genacerores 4-4
(6-6)5-6. — Philippine Islands on Cocos
macgregori Banks,
hh. Preabdomen with lateral margins of its caudal
segments with two plates on each side.
i. Genacerores 4-4(4-4)4-4. — Italy on Ficus.
ficifolii Berl.
ii. Genacerores 3-5(5-8)4-7. — Italy on Ulmus
ficifolii ulmicola Leon,
ff. Pygidium with median pair of lobes moderately long
and narrow, always longer than broad; plates of median
incisura always distinctly longer than median lobes.
g. Scale of adult female reddish brown; ventral scale
with two margins subadjacent, division between
them linear; pygidium with median pair of lobes
distant, notched on mesal and lateral margins; sec-
ond pair of lobes incised, margins of lobelets entire
or notched; plates arranged 2, 2, 2, 6, those between
lobes shorter and smaller than others which are
long and slender; genacerores 3-5(6-11)4-6. — United
States, Mexico, South America, West Indies, Asia.
Ceylon, Australia, South Africa on orange, lime,
palms, magnolia. gloverii Pack.
gg. Scale of adult female pale straw color or fulvous;
ventral scale with two margins straight but distant,
division between them not linear; genacerores
3-3(7-8)4-4. — Ceylon on unidentified shrub.
gloverii pallida Green,
ee. Pygidium never with less than five mesogenacerores, nine
pregenacerores, and nine postgenacerores, usually within
following limits: 5-14(9-24)9-23.
f. Pygidium with conspicuous double transverse series of
oraceratubae extending on each side from level of anus
almost to margin; median pair of lobes conspicuous,
bluntly pointed, irregularly crenulate or dentate; sec-
ond pair of lobes small, incised, mesal lobelet larger;
plaites of usual number; genacerores 6-7(9-10)8-9. —
Java on Coftea. corrugata Green.
ff. Pygidium not with conspicuous double transverse series
of oraceratubae.
g. Pygidium with median pair of lobes crenulate at
distal end, but not deeply notched, frequently pointed ;
mesogenacerores arranged in single transverse row.
h. Pygidium with median pair of lobes pointed and
finely serraite or crenulate.
1. Body never with series of four or five stout
spines on each side of metathorax and segments
of preabdomen; pygidium with second pair of
lobes incised, lateral lobelet much smaller than
284 THE COCCIDAE
mesal, mesal lobelet sometimes with lateral
notch; plates arranged 2, 2, 6, those of each
la;teris arranged in three equdistant groups of
two each.
j. Genacerores 5-8(9-17)7-12; mesospiracerores
6-9. — Lepidosaphes beckii Newm. — America,
Europe, Africa, Asia, Australia on orange,
lemon, citron, fig, grape-fruit, oak, croton,
Elaeagnus, Banksia, Taxus, Pomaderris, etc.
pinnaeformis Bouche.
jj. Genacerores 11-11 (10-13) 11-11. — Italy on
olive. pinnaeformis oleae Leon.
ii Body with series of stout spines on each side
of metathorax and each segment of preabdo-
men; pygidium with two distinct pairs of lobes
and broad inconspicuous slightly projecting
area in position of third pair; median pair of
lobes large, margins minutely denticulate,
sloping each side to blunt point; second pair
of lobes smaller than median, incised, mesal
lobelet much larger than lateral, lobelets blunt-
ly pointed; plates slender, arranged 2, 2, 6, those
of each lateris arranged in three equidistant
groups of two each; mesospiracerores numer-
ous, metaspiracerores wanting; genacerores
4-7(11-14)12-15; anus located cephalad of meso-
genacerores and vulva between postgenacerores.
— Australia on Cassinia cassiniae Green.
hh. Pygidium with median pair of lobes bluntly
pointed.
i. Genacerores 4-4(11-13)7-8; pygidium with me-
dian pair of lobes bluntly pointed and finely
serraite; second pair of lobes incised, lateral
lobelet much smaller than mesal; plates ar-
ranged 2, 2, 6, those of each lateris arranged
in three equidistant groups of two each; me-
dian incisura with marginal projecting point. —
India on Piper. piperis Green.
ii. Genacerores 6-6(7-7)9-9; pygidium with me-
dian pair of lobes well developed with crenu-
late margins; second pair of lobes incised,
margins of lobelets crenulate; plates large and
prominent, arranged 2, 2, and several on each
lateris. — Japan on vine huzenensis Knw.
gg- Pygidium with median pair of lobes blunt at distal
end, notched, not crenulate or finely serrate, some-
times with smaller indentations within notches;
mesogenacerofes always arranged in two or more
transverse rows.
h. Pygdium with six plates on each lateris; median
TRIBE LEPIDOSAPHINI 285
pair of lobes large, broad, blunt, deeply notched
on mesal and lateral margins; second pair of
lobes incised, mesal lobelet much larger than
lateral, their margins entire or irregularly
notched; plates arranged 2, 2, 6, those of each
lateris arranged in three equidistant groups of
two each; genacerores 5-14(9-24)8-18; mesospira-
cerores 3-6. — United States, Brazil, Europe, Japan,
Australia on elm, apple, pear, plum, willow, dog-
wood, poplar, etc. ulmi Linn.
hh. Pygidium with five plates on each lateris; me-
dian pair of lobes large, broad, blunt, deeply
notched on mesal and lateral margins; second
pair of lobes incised, mesal lobelet much larger
than lateral; plates arranged 2, 2, 5, those of
each lateris arranged in three equidistant groups,
two at caudal end, two at cephalic end, and one
near middle; genacerores 10-10(14-14)21-23; meso-
spiracerores 11. — New Zealand on Pittosporum
multipora Leon,
dd. Pygidium with median pair of lobes entire and rounded at
distal end, at most, only finely notched.
6. Pygidium with mesal lobelet of each second lobe never
approximating median lobe in size, always much smaller.
f. Pygidium with median pair of lobes broader than long,
subtriangular in outline; second pair of lobes incised,
two lobelets subequal in length, margins entire; plates
arranged 2, 2, 6, those on each side of each second
pair of lobes distinctly shorter than others, each lat-
eris with plates arranged in three equidistant groups
of two each, two cephalic groups in deep indentations;
genacerores 4-4(7-9)7-8. — Japan on Eurya
euryae Knw.
ft*. Pygidium with median pair of lobes as long as or
longer than broad, sometimes faintly or irregularly
notched, distal end broadly rounded or truncated; sec-
ond pair of lobes incised, two lobelets with entire mar-
gins, mesal distinctly longer than lateral; plates ar-
ranged 2, 2, 4, those on each side of each second pair
of lobes subequal in length to ithose of median incisura,
each lateris with four plates in three equidistant
groups, two at caudal end, one near middle, and one at
cephalic end, those of two cephalic groups much smaller
than plates of caudal group; genacerores 1-7(7-13)7-12;
mesospiracerores 2-3. — Bohemia, Japan on Pinus
newsteadi Sulc.
ee. Pygidium with mesal lobelet of each second lobe equal to
median lobe in size; median pair of lobes as broad as long,
lateral and distal margins convex, converging, distal end
rounded, entire; second pair of lobes incised, mesal lobe-
let as large as median lobe, broader than long, subquad-
286 THE COCCIDAE
rangular, distal margin oblique, entire, lateral lobelet tri-
angular, minute; plates of incisurae subequal in length
to lobes, others longer, arranged 2, 2, 1-2-2-2; each lateris
with broad cristate prominence in position of third lobe;
altaceratubae four on each side, arranged 1-2-1; dorsal
oraceratubae few in number, arranged in rows; anus lo-
cated cephalad of mesogenacerores; genacerores 3-5(6-9)
7-9. — Seychelles on cocoanut palm duponti Green.
bb. Pygidium with proximal ends of median pair of lobes contiguous
or nearly so; median pair of lobes large, with one mesal and
two lateral notches; second pair of lobes minute, incised, mesal
lobelet larger than lateral, with lateral notch; plates arranged
2, 2, 6, those of second inciscurae much smaller than others, each
lateris with six subequidistant plates in three groups of two
each; genacerores 7-12(9-15)7-12; mesospiracerores about 3. —
Europe on fig. ficus Sign.
SPECIES OF CORNUASPIS
a. Body with cephalic end rounded with well defined eye-spots sur-
rounded by fine concentric lines on a leval with rudimentary anten-
nae; pygidium with median pair of lobes broad, distant; second pair
of lobes incised, each lobelet distinctly less than one-half size of each
median lobelet; plates arranged 2, 2, 6; genacerores 2-4(9-10)7-8. —
Seychelles on Davallia. ocellata Green.
aa. Body with cephalic end truncate, its lateral angles produced into
ringed horn-like pappillae; pygidium with median pair of lobes
rounded and notched on lateral and mesal margins; second pair of
lobes incised, entire or nearly so; furcapectinae arranged 2, 1, 5,
projection on one side of furcapectinae small and distant from dis-
tal end; genacerores consisting of twenty-four cerores arranged in
a crescent, 4-4(6-7)4-6. — Philippine Islands on Cocos
unicolor Banks.
SPECIES OF CEPHALASPIS
Pygidium with median pair of lobes bluntly pointed and finely serrate;
second pair of lobes incised, lateral lobelet much smaller than mesal,
entire; plates arranged 2, 2, 6, those of each lateris arranged in three
equidistant groups of three each; two caudal segments of preabdomen
on each side with four or five plates; genacerores 1-6(8-13)6-8; meso-
spiracerores 3-4. — Ceylon on Cocculus cocculi Green.
SPECIES OF UNGULASPIS
Preabdomen with margins of its segments strongly produced and pro-
duced portions with claw-like processes; pygidium with median pair of
lobes faintly notched on lateral and mesal margins; second pair of
lobes incised, mesal lobelet much larger than lateral; plates arranged
2, 2, 6; each lateris with three indentations, each with pair of plates;
genacerores 3-3(6-9)4-6. — Java on Syzygium. ungulata Green.
SPECIES OF SCOBINASPIS
Head with cephalic and ventral aspects in region of antennae with num-
ber of tooth-like projections; pygidium with median pair of lobes
short and broad, faintly notched on mesal and lateral margins; sec-
ond pair of lobes incised, lateral lobelet much smaller than mesal;
TRIBE LEPIDOSAPHINI 287
plates arranged 2, 2, 5, slender in form and longer than lobes, those
of each lateris arranged In three equidistant groups of one, two, and
two; genacerores 4-4(5-6)4-6. — Italy on Croton serrifrons Leon.
SPECIES OF LEONARDASPIS
Pygldium with only single pair of distinct lobes, the median pair short,
about as broad as long, distal margin broad, notched on mesal and lat-
eral margins; plates arranged 2, 3, those of median incisura short and
small, hardly longer than median pair of lobes, those of each lateris
much longer and arranged singly and distant from each other; margin
of lateris thickened, toothed; genacerores 5-5(24-27)29-35. — Australia
on an unknown plant. wilga Leon.
SPECIES OF SCRUPULASPIS
a. Pygidium with second pair of lobes not incised.
b. Pygidium with median and second pairs of lobes distant, dis-
tance between them greater than width of one of second pair of
lobes.
c. Pygidium with median pair of lobes as broad as long.
d. Pygidium with median pair of lobes broad, lateral margin
slightly crenulate, projecting very slightly.
e. Pygidium with genacerores 9-15(13-17)10-17; second pair
of lobes small, rounded, entire; plates arranged 2, 4, 12,
those of median incisura largest, resemble large setae,
those of second incisura short, subdivided, pectinae-like,
may be typical number or toothed prolongations of mar-
gin, those cephalad of each second pair of lobes six or
seven, subdivided, and two distant groups cephalad of
these on each lateris. — New Zealand on Leptospermum...
intermedia Mask,
ee. Pygidium with genacerores 1-3(6-9)4-7; median pair of
lobes narrower than in typical form, other lobes obsoles-
cent; plates very small and inconspicuous; lateral mar-
gins of segments of preabdomen not markedly produced. —
Australia on Acacia intermedia victoriae Green.
dd. Pygidium with median pair of lobes small, only slightly
projecting, rounded, truncate, entire; second pair of lobes
small, finger-shaped, bluntly pointed; plates arranged 0, 1, 3,
those of each second incisura filling space between lobes,
as broad as median lobe, with spine-like projection on mesal
corner, hardly longer than lobes, those of each lateris situ-
ated, small one near second lobe, one at middle, third at
cephalic end; caudal segment of preabdomen with two
plates on each side; median pair of lobes adjacent; gena-
cerores 4-4(12-16)12-13. — Java on Persea longula Leon.
cc. Pygidium with median pair of lobes triangular or subtriangu-
lar, longer than broad, margins entire; second pair of lobes
similar in general form to median but smaller, entire; plates
arranged 0, 2, 3, those of second Incisurae about as long as
288 THE COCCIDAE
second pair of lobes, those of each lateris in two distant
groups, two and one; genacerores 2-4(6-9)1-6. — Australia
defect a Mask,
bb. Pygidium with median and second pair of lobes close together,
not as far apart as width of one of second lobes.
c. Pygidium with second pair of lobes distinctly narrower than
median pair,
d. Pygidium with median pair of lobes large, broadly rounded,
not notched, finely serrate; second pair of lobes minute, tri-
angular, adjacent to first lobe, margin irregular; plates
large, arranged 0, 0, 4, those of each lateris arranged two
adjacent to each second lobe, shorter than other plates, one
near middle, and fourth at cephalic end; genacerores 8-10
(14-15)16-24. — ^Australia on Banksia grandilobis Mask.
dd. Pygidium with median pair of lobes triangular.
6. Pygidium with median pair of lobes with margin broad-
ly crenulate, triangular in outline with distal end bluntly
rounded; second pair of lobes broadly rounded, subtrian-
gular, near but not adjacent; plates apparently wanting;
genacerores 4-6(6-8)8-10. — Japan on Machilus
macMli Mask,
ee. Pygidium with median pair of lobes with margin entire,
triangular in outline, apex nearer mesal margin, mesal
and lateral margins forming unbroken curve with distal
margin, lobes separate but close together, lateral mar-
gins oblique; second pair of lobes with mesal half al-
most as large as median lobe and of much same shape,
lateral half about one-half as broad, shorter and rectan-
'gular; plates arranged 0, 1, 2-2-2, plates of median in-
cisura diverging caudad; altaceratubae arranged 1-2-2-1;
genacerores present; body with triangular process on
each side projecting cephalad of mouth-parts and margin
slightly bulged out. — Ceylon on Mesua amMgua Ruth.
CO. Pygidium with second pair of lobes broader or subequal in
length with median pair, broadly rounded at distal end, mesal
margin notched; plates arranged 0, 1, 3, those of each lateris
placed equidistant from each other, all large; genacerores
3-4(7-11)7-10. — ^Australia on Casuarina easuarinae Mask.
aa. Pygidium with second pair of lobes incised.
b. Pygidium with median pair of lobes broadly rounded, entire;
second pair of lobes incised, mesal lobelet largest, both lobelets
notched; third pair of lobes rudimentary, incised, notched, irregu-
lar in outline; plates arranged 0, 2, 2, 1, situated on each lateris
near third lobe; genacerores 4-5(10-13)11-15. — ^Australia on Eu-
calyptus, Acacia grisea Mask.
bb. Pygidium with median pair of lobes with single mesal notch
and several lateral notches, bluntly pointed between notches;
second pair of lobes incised, entire, mesal lobelet twice as large
as lateral; third pair of lobes very small, entire, not incised;
TRIBE LEPIDOSAPHINI 289
plates arranged 0, 2, 2, 4, those of each lateris arranged in two
groups of two each, two near third lobe and others near cephalic
end; genacerores 4-4(6-7)5-5. — Italy on Phillyrea
destefanii Leon.
SPECIES OF BERLESASPIS
a. Pygidium with median pair of lobes with distal end truncated or
with sides parallel and distal end pointed.
b. Pygidium with median pair of lobes longer than broad, large,
distant, projecting, two sides of each lobe parallel, distal portion
pointed; other lobes apparently obsolete; plates apparently want-
ing in median incisura, each lateris with six to eight; genacerores
2-6(8-10)10-15; mesospiracerores 4. — Australia on Acacia
spinifera Mask,
bb. Pygidium with median pair of lobes short, broader than long,
distal ends truncate, not distant; plates wanting in median in-
cisura and each lateris with eight to ten; genacerores 4-6(12-16)
14-16. — New Zealand on Astelia. epiphytidis Mask.
aa. Pygidium with median pair of lobes conical, not notched; plates
arranged 2, 3, all minute; genacerores 2-3(7-9)1-15. — Australia on
Melaleuca. spinosa Puller.
SPECIES OF FUSILASPIS
a. Pygidium with second pair of lobes incised two or three times.
b. Pygidium with each lateris with five or six plates; median pair
of lobes irregular, not projecting beyond pygidial incision; sec-
ond pair of lobes small, incised, forming two subequal lobelets;
third pair of lobes short and broad, divided by incisions into three
lobelets, median largest; plates arranged 0, 1, 1, 5-6, plates of
each lateris grouped one near third lobe, one near middle of
lateris, others near cephalic end; genacerores 7-8(10-15)16-18;
mesospiracerores 3-7. — Australia on Phymatodes
phymatodidis Mask.
bb. Pygidium with two plates on each lateris.
c. Pygidium with second and third pairs of lobes each divided
into three lobelets, median lobelet in each case largest, mar-
gins of all entire; median pair of lobes fused to form crescen-
tic area with caudal ends bluntly rounded; plates large and
prominent, arranged 0, 1, 1, 2, those of each lateris far apart;
genacerores 7-14(14-20)17-24. — ^Australia on fan-palm
pollens Mask.
cc. Pygidium with second pair of lobes divided into four lobelets
and third pair into three, margins entire or irregularly cren-
ulate; median pair of lobes fused to form crescentic area with
lateral ends prolonged and broadly rounded; plates large and
prominent, arranged 0, 1, 1, 2, those of each lateris far apart;
genacerores 7-9(14-23)20-28. — New Zealand on Cordyline,
Asplenium, Phormium, Gahnia, Drimys, Astelia, Eucalyptus
cordylinidis Mask,
aa. Pygidium with second pair of lobes simple, not incised, margin
290 THE COCCIDAE
rounded and entire, located near to fused median pair of lobes;
median pair of lobes strongly produced, broadly rounded, finely ser-
rulate; third pair of lobes small, incised; plates large, arranged
0, 1, 1, 4, those of each lateris placed separately and equidistant;
genacerores 6-8(20-25)25-35. — New Zealand on Fuchsia... lactea Mask.
SPECIES OF ALBATASPIS
Pygidium with two pairs of lobes; median pair with lateral ends pro-
longed and crenulate; second pair of lobes large, incised, mesal lobe-
let longest, bluntly rounded, lateral lobelet pointed; plates long and
broad at proximal end, arranged 0, 1, 3, those of each lateris placed
singly and equidistant; lateris irregular and finely serrate; genacerores
2-5(5-14)15-24. — Australia on Melaleuca. nivea Mask.
SPECIES OF PINNASPIS
a. Scale of adult female with a median keel and inclined to be tri-
carinate; pygidium with median pair of lobes closely approximated,
notched on lateral margin, broadly rounded; second pair of lobes
small; third pair rudimentary; plates arranged 0, 1, 1, 2, those of
second incisurae small, those of third large, those of each lateris
large and one located near each end; each lateral margin of caudal
segment of preabdomen with two plates. — Jamaica on bamboo
hamhusae Cntll.
aa. Scale of adult female not with median keel and never tricarinate.
b. Pygidium with second pair of lobes incised, mesal lobelet larger
than lateral; median pair of lobes prominent, almost contigu-
ous, rounded, with notch on lateral margin; plates arranged 0, 1,
each lateris with a few plates; lateres serrate; genacerores
4-4(10-10)9-11.— Philippine Islands on Celtls, Sandoricum, Si-
phonodon. siphonodontis Ckll. & Rob.
bb. Pygidium with second pair of lobes not incised.
c. Body without groups of mesospiracerores ; pygidium with me-
dian pair of lobes adjacent, distal end rounded, notched on
lateral margin; second pair of lobes bluntly rounded, short,
curved mesad; plates arranged 0, 1, 3, those of each lateris
situated one near each second lobe, one near middle, one near
cephalic margin; genacerores 5-5(13-15)12-13. — Java on Persea.
romMca Leon.
cc. Body with groups of mesospiracerores.
d. Pygidium with median pair of lobes rounded, adjacent,
deeply notched on lateral margin; second pair of lobes
small, rounded, spatulate; plates arranged 0, 1, 3, those of
each lateris situated, one near second lobe, one near middle,
third at cephalic end; each lateral margin of caudal seg-
ment of preabdomen with two plates; genacerores 4-4(9-12)
11-13; mesospiracerores about 3. — Mytilaspis pandani Comst.
— Massachusetts, New York, West Indies, Panama, Europe
on Buxus, Dictyosperma, Areca, Pandanus, Anthurium,
Thrinax, Dracaena, cocoa-palm buxi Bouche.
TRIBE LEPmOSAPHINI 291
dd. Median pair of lobes uiformly rounded, entire, not notched,
scarcely adjacent, mesal margins divergent; second pair or
lobes similar to first, entire, not notched; small rounded
lobelet caudad of second lobe may be lateral lobelet of sec-
ond lobe; plates arranged 0, 1, 3, those of each lateris sit-
uated, one near second lobe, one at middle, one at cephalic
end; genacerores 6-6(16-17)11-12; mesospiracerores 7-10. —
Java on Ilex. javanica Leon.
SPECIES OF HEMIASPIS
Pygidium of median pair of lobes adjacent, semicircular in form, mar-
gins entire, lateral margin entire or with faint notch; other lobes
wanting; plates flattened, caudal one not as long as lobes, arranged
0, 1-1-1-1, cephalic plate very broad; minute paraphyses, two pairs,
or thickenings located between caudal, first and second, and second
and third plates; dorsal octaceratubae few; genacerores 4-4(6-6)4-4. —
Lepidosaphes hemichionaspiformis Green. — North Australia on Mela-
leuca. hemichionaspiformis Green.
SPECIES OF ISCHNASPIS
a. Pygidium with two plates in each second incisura; plates arranged
0, 2, 6, those of each lateris two, distant, at caudal end, two sub-
adjacent one-third from caudal end, and two subadjacent near
cephalic end; median pair of lobes broadly rounded, widely sepa-
rated, not oblique, finely serrate; second pair of lobes incised, each
lobelet similar to median, entire, broadly rounded, mesal lobelet
larger than lateral; large oraceratubae in each second incisura
laterad of its plates and another laterad of each second lobe; mar-
gin of pygidium broadly concave between caudal oraceratubae;
genacerores in three groups, median group 3-4 and each lateral
group 4-6. — Lepidosaphes ritsema-bosi Leon. — United States, West
Indies, Brazil, Great Britain, Japan, Australia on Pandanus, Latania,
Magnolia, Jasminum, Monstera, palms, coffee, etc
longirostris Sign.
a,a. Pygidium with single plate in each second incisura; plates arranged
0, 1, 3, those of each lateris situated, one near second lobe, one near
middle, and third near cephalic end; median pair of lobes large,
broadly rounded, strongly oblique, distinctly separated, finely ser-
rate; second pair of lobes small, incised, mesal lobelet much larger
than lateral, bluntly rounded, serrate; oraceratubae in each second
incisura adjacent to second lobe, two adjacent ones cephalad of
caudal plate of each lateris and one cephalad of each of other
plates; pygidial margin deeply concave between plates of second
incisurae and median lobes situated in concavity; genacerores ar-
ranged in five groups, 2-3(6-12)4-4; mesospiracerores 0-1. — South
Africa on Cyclostomum, Strychnos. bipindensis Lindgr.
SPECIES OF MYTILELLA
Pygidium with median pair of lobes inconspicuous, tooth-shaped, faintly
notched on lateral margin, distant; second pair of lobes incised,
mesal lobelet slightly larger than lateral, notched on lateral margin,
292
THE COCCIDAE
lobelets similar in form, bluntly rounded; plates arranged 2, 1, 7, those
of each lateris situated, one adjacent to each second lobe, three equi-
distant groups of two each; each lateris between caudal and next
group of plates with four or five adjacent oraceratubae ; genacerores
(4-4) 4-4; mesospiracerores two. — Central America on Anthurium-like
plant. carinata Ckll.
SPECIES OF ANDASPIS
a. Pygidium with two mesal club-shaped paraphyses connected with
mesal margins of median pair of lobes; median pair of lobes dis-
tinct, triangular, lateral margin long, minutely serrate, mesal notch
small; other lobes wanting; plates longer than lobes, arranged 2,
1-2-2-1; margin with six altaceratubae arranged 1-2-2-1; anus super-
imposed over mesogenacerores ; vulva located between pregena-
cerores and postgenacerores; mesospiracerores 1-2, metaspiracerores
0; dorsal oraceratubae small, in two rows; genacerores 4-5(5-6)4-5.
— Lepidosaphes flava hawaiiensis Mask., Lepidosaphes erythrinae
Rutherford, Lepidosaphes moorsi Doane & Ferris. — Ceylon, Hawaiian
Islands, Samoan Islands on Erythrinia, orange hawaiiensis Mask.
aa. Pygidium with two pairs of paraphyses, none connected with lobes;
median pair of lobes large, triangular, adjacent, distal ends meet-
ing like pair of pincers; other lobes wanting; altaceratubae open-
ing in latadentes, arranged 1-(1-1)-(1-1)-1-(1-1) ; plates small and
inconspicuous, shorter than lobes, arranged 0, 2-1-11; anus nearly
over mesogenacerores; dorsal oraceratubae very few; mesospira-
cerores 2-3; genacerores 3-5(7-12)8-12. — Lepidosaphes incisor Green.
— North Australia on Melaleuca. incisor Green.
SPECIES OF AONIDOMYTILUS
Pygidium with three pairs of lobes, median pair large, mesal and lat-
eral notches deep; second pair of lobes incised, mesal lobelet pointed
with deep mesal and lateral notch; third pair of lobes triangular, not
incised; plates arranged 2, 0, 0, 0, limited to minute plates in median
incisura, shorter than median lobes; genacerores 3-3(4-4)2-3. — New
Mexico, Colorado on Chenipodium, Atriplex. concolor Ckll.
SPECIES OF LEPIDASPIDIS
Pygidium with single, large, mesal, crenulate lobe; four distant plates
on each side, cephalic two much larger than caudal two; genacerores
arranged in five groups, 4-4(10-12)11-14, some individuals possess four
groups arranged in transverse row cephalad of mesogenacerores, each
group with two to three cerores; caudal segment of preabdomen with
two large plates, segments not deeply constricted; scale of adult fe-
male narrow, straight or curved, sides subparallel, with whitish, nar-
row flattened border; color dark brown with pale margin, older speci-
mens pale white. — Japan on Osmanthus. uniloba Knw.
SPECIES OF COCCOMYTILUS
a. Preabdomen with lateral margin of each segment with row of
short, thick, tubular bristles, "some with a circular top, some forked,
TRIBE LEPIDOSAPHINI 293
and some appearing like bundles of parallel fibres (pectinae and
plates or setae);" pygidium with two pairs of subsimilar, not in-
cised lobes. — New Zealand on Drimys drimydis Mask.
aa. Preabdomen not with lateral margin of each segment fringed with
row of pectinae or plates or setae.
b. Pygidium with plates between median pair of lobes.
c. Pygdium with three pairs of lobes.
d. Pygidium with second pair of lobes incised.
e. Pygidium with third pair of lobes incised.
f. Pygidium with median pair of lobes short and broad,
distinctly broader than long, bluntly rounded, faintly
notched on mesal and lateral margins, distant; second
pair of lobes incised, mesal lobelet much larger than
lateral, bluntly pointed, faintly notched on mesal and
lateral margins; third pair of lobes incised and notched;
plates arranged 2, 2, 2, 4, those of each lateris situated,
one adjacent to third lobe, two near middle, distant
from each other, one at cephalic end; each lateris more
or less scalloped and indented. — Lepidosaphes cocker-
elliana Kirkaldy. — Florida, New Mexico, Mexico, Ba-
hama Islands, Jamaica on Solanum albus Ckll.
ff. Pygidium with median pair of lobes large, rounded,
entire, not far apart; second pair of lobes incised, mesal
lobelet larger than lateral, lateral margin of mesal
lobelet minutely crenulate; third pair of lobes incised,
margins crenulate, lateral lobelet broadest; each lateris
irregularly serrate, with one or two pointed processes.
— Mexico on "Garabatillo." townsendiana Ckll.
ee. Pygidium with third pair of lobes not incised; median
pair of lobes broad, as broad as long, sloping on both
margins to blunt point, minutely dentate, proximal half
of margins subparallel; second pair of lobes incised,
mesal lobelet much larger than lateral, similar to median
lobe, more or less notched on each side, lateral lobelet
bluntly pointed, entire; third pair of lobes short, broad at
proximal end, rounded; plates subequal in length to lobes,
arranged 2, 2, 3, 3; dorsal oraceratubae prominent. —
Philippine Islands on Ixora ixorae Ckll. & Rob.
dd. Pygidium with second pair of lobes not incised; median
pair of lobes large, margins strongly sloping, converging
caudad, distal end bluntly pointed; second pair of lobes mi-
nute, subtriangular in outline; third pair of lobes similar
in size and shape to second pair; plates of median incisura
shorter than lobes, those of other incisurae and lateris vari-
able in length, arranged 2, 2, 3-4, 3, those of each lateris near
a lobe; brevaceratubae few in number. — Australia on Hymen-
anthera. hymenantherae Green.
cc. Pygidium with two pairs of lobes.
294 THE COCCIDAE
d. Pygidium witli two plates in each second incisura; second
pair of lobes not incised.
e. Pygidium with median pair of lobes broader than long,
short, bluntly rounded, notched on mesal and lateral mar-
gins, distant; second pair of lobes not incised, rounded,
faintly serrulate, two rounded lobe-like projections ce-
phalad of each second lobe; median and second incisurae
each with two plates, each lateris with four or five plates,
those of median incisura large, as long as lobes; lateres
with small serrulations. — Australia on Acacia
convexus Mask.
ee, Pygidium with median pair of lobes entire, broader than
long, distal end truncated with rounded angles, margin
finely crenulate, distant; second pair of lobes not incised,
bluntly rounded, faintly notched; plates arranged 2, 2, 8,
those of incisurae short and stout, shorter than lobes and
plates of lateres, plates of each lateris arranged in three
groups, three near second lobe, three near middle, two at
cephalic end; lateres irregular and serrate. — Australia on
Acacia. acaciae Mask.
dd. Pygidium with single plate in each second incisura; median
pair of lobes pointed, finely serrate, sides strongly roundly
converging; second pair of lobes large, incised, mesal lobe-
let twice as large as lateral, distal end subtruncate, margins
of lobelets finely serrate; plates arranged 2, 1, 6, those of
median incisura minute, not one-half length of median lobe,
all others stout and much longer than lobes, those of each
lateris arranged, one adjacent to second lobe, two near
middle, three at cephalic end; oraceratubae located, one in
each second incisura, two between two caudal groups of
plates of each lateris and adjacent, three adjacent ones be-
tween the two caudal groups of plates. — South Africa on
Berlinia. chitinosus Lindgr.
bb. Pygidium without plates between median pair of lobes.
c. Pygidium with two pairs of rounded, produced, very distinct
lobes; median pair of lobes largest, distant; median incisura
with short bifid process; plates arranged 0, 1, 4, those of each
second incisura long, nearly twice as long as lobe, those of
each lateris arranged, one adjacent to second lobe, three equi-
distant from each other and caudal one; lateris with obtuse
projections; mesal forked organ may be median pair of lobes. —
Brazil on bamboo. hambusicola Ckll.
cc. Pygidium with median pair of lobes large and broad, adjacent,
mesal margin with single notch and lateral with two, distal
portion of lobes bluntly rounded; second pair of lobes much
smaller than median, incised, entire, each mesal lobelet larger
than lateral, adjacent to median lobe; third pair of lobes al-
most obsolete; plates arranged 0, 0, 3, those of each lateris
very large, distal portions elongate and finely attenuate, one
TRIBE LEPIDOSAPHINI 295
placed near third lobe, two adjacent near cephalic endr — ^Ja-
maica on Croton. crotonis Ckll.
SPECIES OF DINASPIS
a. Pygidium with three pairs of lobes; median pair short, bluntly-
rounded, entire, less than width of one of them apart; second pair
of lobes similar in size and form to the first pair, entire; third pair
of lobes small, inconsipicuous, distant from second pair; plates ar-
ranged 2, 1, 1, 0, small, only slightly longer than lobes, those of
each second incisura adjacent to second lobe; oraceratubae situated
one in each second incisura, two in each third incisura and adjacent,
one adjacent to cephalic margin of each third lobe, fifth slightly far-
ther cephalad. — Argentina on Ephedra. ichesii lieon.
aa. Pygidium with two pairs of lobes; median pair of lobes short, blunt-
ly rounded, entire, twice width of one of median pair of lobes apart;
second pair of lobes small, not incised, entire, bluntly pointed, dis-
tant from median pair of lobes; plates arranged 2, 1, 3, those of
each lateris situated one near second lobe, two short distance ce-
phalad; oraceratubae situated, one in each second incisura, five
others equidistant from each other and one in second incisura. —
Argentina on Bulvesia. lahillei Leon.
SPECIES OF ALLANTOMYTILUS
Pygidium with caudal margin truncate and bearing three pairs of simi-
lar, very distinct, subequal, triangular lobes with mesal and lateral
margins faintly notched; plates arranged 2, 2, 2-3, 1, all subequal in
size and much longer than lobes; segments of preabdomen with their
lateral margins bearing three or four broad f urcapectinae ; scale of
male and female with six or more deep transverse constrictions.— Aus-
tralia on Litsea. maideni Mask.
SPECIES OF OPUNTIASPIS
a. Pygidium with three pairs of lobes; median pair prominent, distant,
broadly rounded, slightly notched; second pair of lobes twice in-
cised, cephalic and median lobelets subequal, bluntly pointed, small-
er than median lobelet, entire; third pair of lobes equally promi-
nent, once incised, lobelets bluntly pointed, entire; plates arranged
2, 1, 1, 2, 6-7, those of each lateris in two groups, two situated near
each third lobe, four or five spaced on cephalic portion; mesospira-
cerores 5-6; each lateris indented and finely serrate. — Mexico on
Yucca, Agave. philococcus Ckll.
aa. Pygidium with two pairs of lobes; median pair of lobes long, prom-
inent, bluntly pointed, slightly constricted at proximal end; second
pair of lobes incised, mesal lobelet twice as large as lateral, bluntly
pointed, each subsimilar to median lobe; plates arranged 2, 1, 2;
each lateris cephalad of second lobe with two latadentes, each with
an oraceratuba, each second incisura with latadententis with oracera-
tuba between plate and second lobe. — Java on Agave
javanensis Green.
SPECIES OF PARISCHNASPIS
Pygidium of adult female without genacerores; median pair of lobes
broadly rounded, strongly oblique, distinctly separated, crenulate; sec-
296 THE COCCIDAE
ond pair of lobes large, Incised, mesal lobelet much larger than lateral,
margin finely serrate, truncate; third pair of lobes as large as second,
incised, similar in form; plates arranged 0, 1, 1, 4, those of each lateris
situated, one adjacent to third lobe, one near middle, two at cephalic end,
these plates much larger than others; oraceratubae situated one on me-
son, one in each second incisura, one in each third incisura, one between
lobelets of each third pair of lobes, two cephalad of first plate of each
lateris, one cephalad of second; spiracerores wanting. — Ischnaspis
spathulata Lindgr. — India on Atica. spathulata Lindgr.
SPECIES OF NIVEASPIS
Pygidium with median pair of lobes hardly projecting, margin entire,
caudal end truncated, angles rounded; each second lobe with margin
more oblique, hardly projecting; plates wanting, but with spine-like
projections, these projections in median and second incisurae and
cephalad of third pair of lobes; preabdomen with lateral margins of
segments straight, but with strong finger-like process near each caudo-
lateral angle. — Lepiodsaphes argentata Ckll. — Brazil, Mexico
argentata Ckll.
SPECIES OF FERNALDANNA
Pygidium with median pair of lobes minute, entire, rounded, only slight-
ly projecting; second pair of lobes large, short and broad, much
broader than long, entire, distal margin convex, each six or eight times
as broad as median lobe; plates wanting; oraceratuba in each second
incisura, another cephalad of each second lobe, three on each lateris
cephalad of second lobe, equidistant from each other and oraceratuba
adjacent to this lobe; each lateris apparently with densariae in
which oraceratubae open; mesospiracerores 0-1. — Fernaldella indentata
Green. — Australia. indentata Green.
SPECIES OF MASKELLANNA
Pygidium with median pair of lobes minute, narrow, rounded, entire,
constricted on proximal portion, very distant; second pair of lobes mi-
nute, subequal in size to median and similar in form, distant from
median; plates wanting; altaceratubae arranged 1, 1, 2, 2, 1, one in
median incisura; setae minute; lateres with slight projections near
oraceratubae of altaceratubae, otherwise entire; dorsal oraceratubae
numerous, minute, promiscuously arranged; body of adult female
strongly chitinized, no differentiation between pygidium and other
parts. — Mytilaspis (Ferneldella) beyeriae Green. — Australia on Bey-
eria. 'beyeriae Green.
SPECIES OF PHAULOMYTILIUS
Pygidium without lobes or plates; margin irregularly crenulate; with
four small, conical, widely separated, spine-like projections arising
from surface of pygidium; large oraceratuba located on each side of
each cephalic projection; scale of female pyriform, marked with
alternate transverse bands of silvery white and dark gray. — Australia
on Casuarina. striata Mask.
CHAPTER XXV
TRIBE DIASPIDINI
The Double-shielded Scales
The body of the adult female is usually longer than broad, rare-
ly broader than long, frequently two or three times as long as broad.
The cephalic end is usually bluntly rounded and the caudal end is
often bluntly pointed and frequently provided with a pygidial incision.
The lateral margins are convex, the body is broadest through the
prothorax or the first abdominal segment, and the sides usually con-
verge caudad. The indentations marking the position of the coriae
between the thoracic and abdominal segments are usually distinct. The
lateral margins of the segments of the preabdomen are never provided
with bracteae either in the adult female or in the second nymphal
female, but often with plates. Lobes are usually present, rarely entire-
ly wanting. Plates are rarely absent, usually only in case the lobes
also are wanting, pectinae are rarely present and, if present, are repre-
sented by simple furcapectinae. The incisurae, except the median, are
usually narrow and filled by the plates. The median incisura is fre-
quently a pygidial incision and is usually without plates. Densariae are
never present, paraphyses are only rarely present and, when present,
are not typical in form. Altaceratubae are rarely, if ever, wanting;
they are arranged singly, rarely in pairs, their oraceratubae oval in out-
line, rarely twice as long as wide. Brevaceratubae are usually pres-
ent, generally numerous, frequently arranged in interrupted transverse
rows. Genacerores are usually present, usually arranged in five groups,
sometimes with seven or more and sometimes with only two, three,
or four groups. Mesogenacerores, when present, consisting of more than
eight cerores. Scale of the adult female coriaceous or filamentous,
usually whitish, sometimes grayish or brownish. Scale variable in form,
either round or nearly round with the two exuviae placed just within
the periphrey or projecting slightly or considerably beyond it; the scale
oval or pyriform or mussel-shaped in outline with the exuviae placed
at one end; or greatly elongated, six or more times as long as broad,
with the sides parallel. If pyriform, much longer than broad with both
sides convex or with one side convex and the other concave, and with
one end much smaller than the other. The two exuviae overlapping,
one on the end of the other, the first exuvia bearing the remains of
the antennae as porrect projectons. The female is rarely enclosed In a
puparium, but in a few species lives in galls and the lobes and plates
may be represented by pseudolobes. The scale of the male is elongate,
two or three times as long as wide, with the single exuvia projecting
beyond one end. The wax is usually opaque, white, or chalky in ap-
297
298 THE COCCIDAE
pearance, rarely glassy and transparent. The surface is often more or
less distinctly tricarinate.
There is a vague similarity about the general appearance
of the species of the tribe Diaspidini by which one who is familiar
with the species of the Diaspidinae is able to recognize them.
This appearance is common to the scales of the adult female and
of the male, the general form of the body of the adult female^
the pygidium, and the pygidial fringe. While there is this similar-
ity in appearance, yet it is too vague to define. The species that
belong to the tribe Diaspidini as listed in Fernald 's Catalogue are
about thirty. These are distributed among nine genera.
The scale of the male is probably more typical for the tribe
than that of the female. They are generally quite similar in size,
shape, and appearance, elongate oval in outline with the two
sides parallel or subparallel with the caudal end bluntly rounded
and with the cephalic end rounded and bearing the single exuvia
which projects beyond the waxy portion of the scale and bears
the porrect remains of the antennae of the first nj^mph. The
dorsal surface of the secretionary portion is slightly or strongly
convex and may be uniformly convex or flat and faintly or strongly
carinate. The wax of the male scale is almost invariably opaque
white or chalky white. This chalky white appearance of the
male scales of many species is characteristic. It is a feature that
is emphasized from a habit of the males of certain species. At
certain times they congregate in immense numbers, in such num-
bers that a branch an inch in diameter for several inches or the
surface of an entire leaf may be completely covered by them. The
grouping together of such numbers of the chalky white scales
makes the branch or leaf very conspicuous even at a considerable
distance.
The scales of the adult female of the different species are very
unlike in size, shape, and appearance, sometimes even among the
species of the same genus. They are of four types which inter-
grade more or less. In most species they are pyriform with the
two exuviae at the small end, the caudal and larger end is
broadly rounded with the sides slightly convex and strongly di-
vergent caudad. In a very few species the scale is very long,
eight or more times as long as wide, with parallel sides. When
the scale is mussel-shaped, which occurs in only a limited number
of species, it is curved with one margin convex and the other con-
cave. The surface of the mussel-shaped scales is usually more
or less transversely ridged as in an oyster shell. A third type
TRIBE DIASPIDINI 299
of scale is circular or subcircular. The exuviae overlap in the
circular scales just as in the pyriform scales or mussel-slrapefi-
scales, but they are frequently placed so that they do not project
beyond the periphery of the scale. There may be in fact a nar-
row area of wax between the cephalic end of the first exuvia and
the periphery or the exuviae may project very slightly beyond the
periphery, only the first exuvia protruding, or the exuviae may
protrude for a considerable distance, for as great a distance as
in the pyriform scales. In the location of the exuviae, even in
different individuals of the same species, there may be consider-
able variation, sometimes being placed parallel or nearly so with
the margin and at others perpendicular to it. When the exuviae
project a considerable distance beyond the periphery of the scale,
the wax of the scale adjacent to the exuviae is sometimes extend-
ed so that the scale becomes slightly prolonged and approaches
the pyriform type. The scales, while they are more often white
than any other color, are often grayish or blackish, particularly
in the genus Chionaspis. The texture of the scale in many species
as in those of the genus Diaspis and its allies is very thin, semi-
transparent, tough, and parchment-like. In the darker colored
mussel-shaped scales, they are more coriaceous and leathery like
the scales of the genus Lepidosaphes. The exuviae overlap each
other and the first nymph bears the remains of the antennae
as porrect projections. The exuvia of the second nymph is
about three or four times as long as that of the first. It is
always much less in extent than that of the wax of the scale,
covering one-fourth or less of the area of the scale. Only a very
few of the species transform in a puparium which explains why
the second exuvia is generally not very large. The ventral scale is
extremely variable in extent and texture, it may be very thin and
delicate and remain attached to the host-plant when the dorsal
scale is lifted or it may be stout and thick and retain its con-
nection with the dorsal scale when removed.
The scales of certain species of Chionaspis resemble closely
the scales of most of the species of Lepidosaphes, some individ-
uals resembling them so closely that it is necessary to examine
the body of the female to be sure of their correct generic position.
The Latin word aspis from which so many of the generic names
in this subfamily are compounded means a shield. This may
have been applied originally to the scale as a whole, but evident-
ly in the case of the generic name Diaspis from which the name
of this subfamily and tribe is derived, refers to the exuviae of
300 THE COCCIDAE
which there are two. The word Diaspis would mean literally
then two shields and since the genus is the typical one of the
tribe, the common name of The Double-shielded Scales has been
applied to them. The name of a subfamily and tribe according
to rule is derived from the stem of the genative which in the
case of Diaspis is Diaspidis, hence, the proper form for the
names of these groups is Diaspidinae and Diaspidini and not
Diaspinae and Diaspini as they are generally written.
The body like the scale varies considerably in size. Where
the scale is pyriform, mussel-shaped, or elongate, the body if
elongate, is usually two to four times as long as broad; but in
those species with a circular scale, the body is shorter and broader
and the greatest width is through the thoracic region which is
always much wider than the preabdomen. In dried specimens
the segments of the preabdomen are crescentic and retracted into
the caudal part of the thorax. The pygidium in turn is retract-
ed into the caudal end of the preabdomen so that the preabdomen
and the pygidium seem to constitute only a small portion of the
area of the body. The cephalic part of each lateral margin of
the thorax sometimes bears a short bluntly rounded tubercle. In
those species with elongate bodies the preabdomen and pygidium
are larger and constitute, even in dried retracted individuals, a
much larger and more prominent portion of the body. The ce-
phalic end of the body is usually broad and convexly rounded.
The width of the cephalic end varies with the species, in some it
is the broadest part of the body while in others it may be only
one-half or less of the greatest width. The body, while it is al-
ways depressed, is never decidedly flattened as in some other
tribes of Diaspidinae. The antennae are usually represented by
minute distant tubercles bearing one to three stout setae.
The preabdomen is generally distinctly segmented and easily
distinguished from the thorax and pygidium. The segments are
separated by deep constrictions marking the position of the coriae.
The portions of the segments between the coriae are generally
strongly convex, which generally emphasizes the depth of the
coriae. The lateral margins of the segments, particularly the
caudal ones, are frequently provided with plates or short blunt
plate-like projections, but never with bracteae.
The pygidium is comparatively large and well differentiated.
It is usually circular or oval in outline, both the cephalic and the
TRIBE DIASPIDINI 301
caudal margins are convex. The lateral margins are slightly
convex and are rarely if ever almost straight and converge caudad.
The mesal part of the caudal margin of the pygidium in a num-
ber of species is concave or emarginate and provided with a pygi-
dial incision. This is variously described as a deep median cleft
or the pygidium as semicircularly hollowed out.
The lobes vary in number as in the other tribes. There are
typically about three pairs, the median pair is always the largest.
The second or third pairs may be divided into two or three lobe-
lets or deeply incised. When the caudal margin of the pygidium
is concave or provided with a pygidial incision, the first pair of
lobes is usually located in the concavity, oblique in poseition and
the greater part of their lateral margin is continuous with or
fused to the pygidium. Their mesal margins, which are usually
convexly rounded and continuous with the rounded distal ends,
are usually finely serrate or crenulate. Where the median pair
of lobes is free on the lateral margin, their distal ends are
usually notched. In certain species the proximal ends of the
median lobes are fused, in others their mesal margins are straight
and adjacent, the two lobes appearing to form a single semicircu-
lar area, while in others the adjacent mesal margins have fused
until there is only a notch on the meson marking their point of
separation. The margins of the second and third pairs of lobes
may be serrate, crenulate, or notched, if notched, the number
and distinctness of the notches vary with the species. In a few
species all the lobes have been lost and the margin of the pygidium
is entire, in some others pseudolobes which are very lobe-like in
appearance have replaced the true lobes.
The plates are only rarely wanting except in the median in-
cisura where they are only rarely present. They are usually long
and slender, as long as or longer than the lobes, spine-like in form,
with lamaceratubae opening in their pointed ends. The plates
are usually arranged, 0, 2, 2, 2, 2, or 0, 1, 1, 1, 2. There is not
much variation in the number in the different incisurae and on the
lateris. The plates are in extreme cases represented by apically
furcate furcapectinae. The group of four or five plates near the
cephalic end of the pygidium is located on the caudal segment of
the preabdomen.
The altaceratubae are prominent, large with comparatively
long tubes. They are generally located singly but in some species
302 THE COCCIDAE
there are two adjacent ones in each third or third and fourth in-
cisura. They are never as large as those of the Lepidosaphini
and their oraceratubae are generally shorter and more nearly cir-
cular in outline. The oraceratubae are generally located in lata-
dentes. The presence of latadentes seems to be quite constant
for all the species.
The brevaceratubae are more prominent and more numerous
in many of the species of this tribe than in the species of other
tribes. In some species their oraceratubae are irregularly dis-
tributed over the dorsal surface while in others they are arranged
in crescentic or semicircular transverse rows. Where the breva-
ceratubae are arranged in rows, each row usually consists of a single
line of oraceratubae, but the row may consist of two or three
lines or of a number of oraceratubae irregularly arranged,
forming a broad band. The rows are always so far as observed
interrupted on the meson, forming right and left halves. The
halves of the rows in many species are also interrupted near the
middle of each lateral half so that each complete crescentic trans-
verse row is generally subdivided into four parts or groups. The
crescentic rows of oraceratubae are located on lines connecting
the median lobes, the second incisurae, the third incisurae, the
fourth incisurae, the fifth incisurae, and the cephalic margin of
the pygidium. The lines connecting the points named on the
two sides of the pygidium extend cephalad in each case around
the anus. Green has recognized six of these lines as present,
while Cooley identifies only five. Both Green and Cooley includ-
ing Morgan consider the lines of oraceratubae as indicating, at
least approximately, the lines of the suppressed segments of the
pygidium. These authors differ, however, somewhat as to the
way in which these lines should be traced. Green considers the
rows of oraceratubae as marking the posterior or caudal border
of the segment and it would seem as if there was evidence for
his conclusion. The lines or rows of oraceratubae are numbered
on each side, beginning at the meson, as the first, second, third,
fourth, fifth, and sixth rows. In order to be able to distinguish
between the two groups of each half row, the group near the
meson may be known as a mesal group, the anterior group of
Cooley, and the group near the margin of the pygidium as the
lateral group, the posterior group of Cooley. There are rarely
oraceratubae present in the first row. They are also of rare oc-
currence in the second row, if present, they are only represent-
ed by a mesal group or a lateral group. The other rows are gener-
TRIBE DIASPIDINI 30^
ally represented by both mesal and lateral groups, sometimes by
only a single group, mesal or lateral. There is considerable varia--
tion in the number of oraceratubae in the various groups in differ-
ent species and some use has been made of this difference in number
in differentiating species.
The lamaceratubae are never very prominent in the pygidia
of this tribe. They are slender and extend through each plate,
their oraceratubae located at the distal end, as in the plates found
in the pygidia of other species.
The vulva is a transverse slit, which is usually located near
a transverse line drawn through the pregenacerores and post-
genacerores, that is, it is approximately central in position. There
is a slight variation from this position, in some it is a short dis-
tance cephalad and in others a short distance caudad of this line.
While the vulva is fairly constant in position, the anus seems
always to be extremely variable. It may be located caudad of a
line drawn through the caudal margin of the postgenacerores or
in the cephalic portion of the pygidium, usually cephalad of a
line drawn through the cephalic ends of the pregenacerores or
it may be located anywhere between these lines. In a number
of species it is superimposed over the vulva.
The vulva is typically surrounded by five groups of gena-
cerores. In a few genera the number may be increased to seven
or more groups. The number of cerores in each group is vari-
able. There is usually about eight mesogenacerores, the number
varying from two to thirty or more. The groups of pregena-
cerores may contain a few cerores, twelve or a large number,
sixty. There is also considerable variation among the postgena-
cerores, the number varying from one to forty. The mesogena-
cerores are sometimes wanting and in several genera all the gena-
cerores are wanting. Green records the presence of two more
or less definite glandular scars. These are reticulate and located
cephalo-laterad of each group of pregenacerores. They are con-
sidered as homologous with the groups of genacerores, in fact he
describes the occurrence of cerores in one of these glandular scars.
The marginal setae of the pygidium are frequently compara-
tively short and stout, so that they extend as rigid projections
and, for this reason, are easily mistaken for plates and have
been described as such. This is particularly true of the two
prominent setae located in the median incisura. These setae
instead of being located on the dorsal side of the median pair
304 THE COCCIDAE
of lobes as is usual are located between and project between the
lobes as rigid extensions or like plates.
The parts of the pygidial fringe that in this tribe occur
with only rare exception are beginning at the meson arranged
in the following order. There is a well marked median incisura
sometimes containing one or two latadentes, usually without
plates, and bounded on each side by a median lobe. The second
incisura cephalad of each median lobe contains a latadentis and
one or two plates. This incisura is limited by a second lobe.
The third incisura also contains a latadentis and one or two
plates and is limited by a third lobe. The fourth pair of lobes
may be present but is only rarely so and the area cephalad of
the third lobe belongs to a lateris and usually bears four groups
of plates, the groups usually separated by latadentes. The fourth
or cephalic group generally contains more plates than the others.
The genera and species of this tribe are difficult to separate.
Many of the structures available in the other tribes for separat-
ing genera and species are not available here. Some writers
have not always been careful to differentiate the characters present
so that it is sometimes difficult to decide whether a given species
has lobes, plates, or what is more likely latadentes bearing ora-
ceratubae.
The following species have heen omitted from the tables, either be-
cause the description was not available or because it was not possible
to place the species: — Chionaspis bambusae Ckll. from Japan on bam-
boo; depressa Zehnt. from E. Java on Saccharum; myrthi Bouche from
France on Myrtus; natalensis Mask, from Natal on grass; nitida
Mask, from Australia on Daviesia; planchonii Sign, from southern
Europe on Quercus; rosae Goethe from Germany; sacchari-folii Zehnt.
from Java on sugar cane; tegalensis Zehnt. from Java, Mauritius;
Diaspis blankenhorni Targ from Italy; celtidis Ckll. from Texas on
Celtis; chilensis Ckll. from Chili; colvei Penzig from Italy; cordiae
Rubs, from Brazil on Cordia; euphoriae Charm, from Mauritius on
Nephelium; minima Targ. from Europe on arbor-vitae; monserati Colv.
from Spain on Corynocarpus(?) ; olea Colvee from Spain on olive;
sen tali Mask from New Zealand on Santalum, pear, plum; squamosus
Newst. & Theobald from Egypt on peach, pear; taxicola Vays. from Al-
geria on Taxus; trinacis Colvee from Spain on Trinax, Strelitzia; vici
Schr. from Europe on mistletoe; Aspidiotus (Diaspis) vitiensis Mask.
(Leon.) from Fiji; Epidiaspis subterranea Lindgr. from France on grass,
evidently an aspidiotid; Poliaspis casuarinae Lidg. from Australia on
Casuarina; Pseudoparlatoria ostreata Ckll. from Jamaica on Acalypha.
The tables offered for separating the genera and species should be used
with care and wherever possible specimens should be compared with the
original descriptions and figures.
TRIBE DIASPIDINI 305
GENE-RA OF DIASPIDINI
a. Pygidium always with one or more groups of genacerores.
b. Female never transforming in puparium; scale rarely without
distinct second exuvia.
c. Pyigidum never with more than five groups of genacerores.
d. Pygidium always with four or five groups of genacerores.
e. Pygidium always with plates in median incisura.
f. Pygidium with plates of median incisura separate, prox-
imal ends, even if adjacent, not fused and never form-
ing fish-tail-shaped projection.
g. Pygidium with two or three pairs of lobes, second
pair incised; genacerores in five groups. — Type,
Diaspis juniperi Bouche Carulaspis MacG.
gg. Pygidium with series of fifteen or more pairs of
lobe-like projections or latadentes; genacerores in
four groups. : Diaspidistis Hemp.
ff. Pygidium with pair of plates in median incisura adja-
cent and fused at proximal end and with distal portions
divergent, forming fish-tail-shaped projection
Pseudoparlatoria Ckll.
ee. Pygidium never with plates in median incisura, frequent-
ly with pair of large setae that are plate-like in appear-
ance.
f. Scale of adult female round or oval with exuviae usually
located just within or projecting slightly beyond the
periphery of the scale.
g. Pygidium with plates in incisurae or on lateres or on
both.
h. Pygidium with median pair of lobes, even if adja-
cent, never fused.
i. Pygidium with rounded latadentes or plates in
incisurae other than median, never with trun-
cate lobe-like projections.
j. Pygidium with oraceratubae of brevacera-
tubae arranged in crescentic transverse rows.
k. Pygidium with caudal margin convex,
never concave with median pair of lobes
placed in concavity. — Type, Aulacaspis
pentagona Targ Pseudaulacaspis MacG.
kk. Pygidium with caudal margin deeply con-
cave on meson, forming pygidial incision
with median pair of lobes located in con-
cavity. Aulacaspis Ckll.
jj. Pygidium with oracertubae of brevaceratubae
arranged promiscuously, never in crescentic
transverse rows.
•306 THE COCCIDAE
k. Pygidium with caudal margin convex,
never concave with median pair of lobes
located in concavity; brevaceratubae always
few in number.
1. Pygidium with anus located about its
own width from vulva, distant from
caudal margin. Epidiaspis Ckll.
11. Pygidium with anus located about mid-
way between vulva and caudal margin. —
Type, Diaspis montana Ckll
Cocker ellaspis MacG.
kk. Pygidium with caudal margin deeply con-
cave on meson with median pair of lobes
in concavity; brevaceratubae usually nu-
merous; anus located nearly midway be-
tween vulva and caudal margin than to
vulva. Diaspis Costa.
ii. Pygidium with truncate lobe-like projections
located in incisurae other than median. — Type,
Diaspis regularis Newst JJm'baspis MacG.
hh. Pygidium with median pair of lobes fused into
single area. — Type, Chionaspis malloti Ruth
Rutherfordia* MacG.
gg. Pygidium without plates in incisurae or on lateres.
h. Pygidium with two pairs of lobes, five groups of
genacerores, anus located near caudal margin, and
large altaceratubae Protargionia Leon.
hh. Pygidium without lobes, genacerores forming
U-shaped group, anus located distant from caudal
margin, and altaceratubae small or wanting. — Type,
Protodiaspis agrifoliae Essig Essigaspis MacG.
ff. Scale of adult female never circular, always pyriform,
mussel-shaped, mytilaspiform, or linear, twice or more
as long as wide, rarely with poorly defined scale and
living in gall,
g. Insects always living exposed, never producing ab-
normal growths or galls upon plants,
h. Pygidium of adult female always with plates in
incisurae or on lateres.
i. Pygidium with caudal margin convex, never
with deep angular or semicircular pygidial in-
cision.
j. Pygidium with mesal margins of median
pair of lobes convexly rounded and diverg-
ent, if straight, two margins usually distant,
if straight and adjacent, proximal ends fused,
*This genus is named for my former student, the late Andrew Rutherford, who died
from enteric fever while in service as Government Entomologist of Ceylon.
TRIBE DIASPIDINI 307
two lobes together never forming single com-
pact semicircular area.
k. Pygidium with median pair of lobes nor-
mal in form, never minute and plate-like.
1. Pygidium with oraceratubae of breva-
ceratubae arranged in crescentic trans-
verse rows, sometimes few in number,
m. Pygidium with rows of oraceratubae
of brevaceratubae of each side of
meson divided into two prominent
subgroups,
n. Pygidium with median pair of lobes
sometimes adjacent, but never fused,
o. Pygidium with three or four pairs
of lobes.
p. Pygidium with four pairs of
lobes. — Type, Chionaspis lani-
ger Newst.-Sclopetaspis MacG.
pp. Pygidium with three pairs of
lobes. Chionaspis Sign.
00. Pygidium with one or two pairs
of lobes,
p. Pygidium with two pairs of
lobes. — Type, Chionaspis gram-
inis Green
Duplachionaspis MacG.
pp. Pygidium with one pair of
lobes. — Type, Chionaspis cole-
mani Knw
Unachionaspis MacG.
nn. Pygidium with median pair of lobes
fused on proximal portion, remain-
der of lobes divergent or adjacent,
rarely completely fused; combined
lobes not semicircular,
o. Pygidium with three pairs of lobes.
— Type, Chionaspis americana
Johns. Fundaspis MacG.
00. Pygidium with one pair of lobes.
— Type, Diaspis chionaspiformls
Newst. Africaspis MacG.
mm. Pygidium with rows of oraceratubae
of brevaceratubae few in number and
not divided into two subgroups.
n. Body with head and thorax together
distinctly more than twice as long
as abdomen. — Type, Chionaspis
elongata Green Greenaspis MacG.
308 THE COCCIDAE
nn. Body with head and thorax not ab-
normal in length, together only
slightly longer than abdomen. —
Type, Chionaspis minuta Green
Tenuiaspis MacG.
11. Pygidium with oraceratubae of breva-
ceratubae arranged promiscuously, nev-
er in crescentic transverse rows. — Type,
Chionaspis acuminata Green
Unaspis MacG.
kk, Pygidium with median pair of lobes minute
and plate-like, second pair of lobes large
and apparently with six plates between
them. — Type, Chionaspis striata Newst
Lineaspis MacG.
jj. Pygidium with mesal margins of median pair
of lobes straight and adjacent, rarely if ever
fused, two lobes together always forming
single compact semicircular area
Hemichionaspis Ckll.
. Pygidium with caudal margin deeply concave,
forming deep angular or semicircular pygidial
incision.
j. Pygidium with median pair of lobes located
within pygidial incision,
k. Body with preabdomen symmetrical, dextral
and sinistral margins of segments similar, if
bearing setae on spine-like projections,
these confined to thorax and located on both
sides.
I. Body with thorax at most never twice
as long as wide, varying from as wide
as long to one and one-half times as long
as wide.
m. Thorax normal, not bearing one or
two small tubercle-like bracteae with
ceratubae on mesothorax and meta-
thorax, mesospiracerores usually pres-
ent. Phenacaspis Ckll.
mm. Thorax bearing one or two small con-
ical tubercle-like bracteae with cerat-
ubae on mesothorax and metathorax;
mesospiracerores wanting. — Type, Chi-
onaspis arundinariae Green.
Canaspis MacG.
II. Body with thorax with parallel sides and
two and one-half times as long as wide. —
Type, Chionaspis praelonga Newst
Cooleyaspis MacG..
TRIBE DIASPIDINI 309
kk. Body with preabdomen asymmetrical, sini-
stral margin of four segments of preabdo^
men bearing large spine-like setae on
tubercle-like swellings of margin, wanting
dextral margin. — Type, Chionaspis unilater-
alis Newst. Sinistraspis MacG.
jj. Pygidium with median pair of lobes located
on ends of projections forming lateral limits
of pygidial incision. — Type, Chionaspis nudata
Newst. Augulaspis MacG.
hh. Pygidium of adult female always without plates,
i. P*ygidium always with one or more pairs of dis-
tinct lobes,
j. Pygidium with three pairs of lobes. — Type,
Chionaspis africana Newst
Gramenaspis MacG.
jj. Pygidium with one pair of lobes. — Type, Chi-
onaspis subnudata Newst. --Contigaspis MacG.
ii. Pygidium without lobes, margins undulate. —
Type, Chionaspis simplex Brain
Natalaspis MacG.
gg. Insects always living in abnormal growths or galls
upon plants; scale of adult female lining cavity of
gall, exuviae placed in orifice of gall or wanting;
pygidium of adult female with triangular, median,
lobe-like projection. — Type, Chionaspis galliformens
Green. Parachionaspis MacG.
dd. Pygidium with a single transverse group of genacerores
Parlatoreopsis Lindgr.
cc. Pygidium always with more than five groups of genacerores,
seven or more.
d. Pygidium with oraceratubae of brevaceratubae arranged in
crescentic transverse rows; with at least one pair of lobes;
scale of adult female pyriform, of male not carinated
Poliaspis Mask.
dd. Pygidium with oraceratubae of brevaceratubae promiscuous-
ly arranged, not in rows; lobes wanting. — Type, Chionaspis
simplex Green. PoUaspoides MacG.
bb. Female always transforming in puparium; scale of adult female
not with second exuvia.
c, Pygidium of adult female with plates between median pair of
lobes. Cryptaspidus Lindgr.
cc. Pygidium of adult female never with plates between median
pair of lobes, with deep pygidial incision with oblique distant
lobes. Cryptodiaspis Lindgr.
aa. Pygidium always without groups of genacerores.
310 THE COCCIDAE
b. Body of adult female never enclosed in puparium; scale of adult
female usually with second exuvia.
c. Pygidium with plates or pectinae cephalad of median pair of
lobes,
d. Body never with deep transverse constriction.
6. Pygidium of adult female always without paraphyses.
f. Pygidium with furcapectinae or plates in median in-
cisura.
g. Pygidium with furcapectinae in median, second, and
third incisurae; caudal margin with pygidial incis-
ion.— Type, Diaspis zamiae Morg. Furchadaspis MacG.
gg. Pygidium with plates in median, second, and third
incisurae; caudal margin convex, not with pygidial
incision. — Type, Chionaspis funtumiae Newst
Mitulaspidis MacG.
ff. Pygidium without furcapectinae or plates in median in-
cisura.
g. Pygidium with furcapectinae on lateres; median pair
of lobes incised and smaller than second pair. — Type,
Chionaspis capensis Newst Dentachionaspis MacG.
gg. Pygidium never with furcapectinae on lateres; me-
dian pair of lobes never incised.
h. Body with rostrum normal in position, distant
from cephalic margin.
i. Body with rudimentary antennae normal in
form, consisting of minute projection bearing
one to three setae, never located on chitinized
area bearing corona of four to six spine-like pro-
jections.
j. Pygidium with three or four pairs of lobes;
scale usually elongate.
k. Pygidium with four pairs of lobes; caudal
margin with distinct pygidial incision ;
median pair of lobes rudimentary. — Type,
Chionaspis amaniensis Lindgr
Inchoaspis MacG.
kk. Pygidium with three pairs of lobes; me-
dian pair of lobes always distinct.
1. Pygidium with anus and vulva superim-
posed.
m. Pygidium with caudal end truncate or
convex; median pair of lobes distant
and strongly divergent caudad. — Type,
Chionaspis permutans Green
Graphaspis MacG.
I
TRIBE DIASPIDINI 311
mm. Pygidium with caudal margin with
deep pygidial incision, median pair of
lobes located in incision and not or
scarcely projecting beyond its sides.
— Type, Chionaspis flava Green
Ametrochaspis MacG.
11. Pygidium with anus distinctly farther
caudad than vulva.
m. Pygidium with caudal margin with
deep pygidial incision; each median
lobe as large as or larger than mesal
lobelet of second lobe; median pair of
lobes subadjacent. — Type, Chionaspis
citri Comst. Prontaspis MacG.
mm. Pygidium with caudal margin convex;
each median lobe distinctly smaller
than mesal lobelet of second lobe; me-
dian pair of lobes distinctly farther
apart than width of median lobe. —
Type, Chionaspis dentilobis Newst
Remotaspis MacG.
jj. Pygidium with one or two pairs of lobes.
k. Pygidium with two pairs of lobes.
1. Pygidium with anus located distinctly
cephalad of vulva, near cephalic margin.
m. Pygidium with two pairs of large
prominent lobes; oraceratubae of
brevaceratubae in rows. — Type, Chion-
aspis distorta Newst
Asymmetraspis MacG.
mm. Pygidium with two pairs of minute
inconspicuous lobes; oraceratubae of
brevaceratubae not in rows, promiscu-
ously arranged. — Type, Protodiaspis
lobata Ferris OMuctaspis MacG.
11. Pygidium with anus located distinctly
caudad of vulva, near caudal margin. —
. Type, Pseudodiaspis condaliae Ferris
Situlaspis MacG.
kk. Pygidium with single pair of lobes.
1. Pygidium with caudal margin uniformly
convex. — Type, Chionaspis (?) hikosani
Kuwn. Kuwanaspis MacG.
11. Pygidium with caudal margin with dis-
tinct pygidial incision. — Type, Diaspis
pugionifera Lindgr Inciaspis MacG.
312 THE COCCIDAE
ii. Body with rudimentary antennae borne on circu-
lar strongly chitinized area fringed with four
to six stout spine-like projections. — Type,
Chionaspis coronifera Green. ..Goronaspis MacG.
hh. Body with rostrum abnormal in position, located
quite close to cephalic margin. — Type, Chionaspis
substriata Newst. Dentaspis MacG.
ee. Pygidium with at least one pair of paraphyses.
f. Thorax usually wider than long, never four times as
long as wide; two plates in median incisura
Howardia Leon,
ff. Thorax four or more times as long as wide; plates want-
ing in median incisura. — Type, Chionaspis cinnamomi
Green. Proceraspis MacG.
dd. Body of adult female with deep transverse constriction be-
tween mesothorax and metathorax. — Type, Chionaspis vuilleti
Marchal. Marchalaspis MacG.
cc. Pygidium never with plates cephalad of median pair of lobes,
d. Pygidium with at least one pair of distinct lobes, never with
pseudolobes.
e. Pygidium with caudal margin convex, not with pygidial
incision.
f. Pygidium with anus located distinctly cephalad of mid-
dle of pygidium, near its cephalic end. — Leucaspis
cupressi Colem Cupidaspis MacG.
ff. Pygidium with anus located distinctly caudad of middle
of pygidium, near its caudal margin. -Pseudodiaspis Ckll.
ee. Pygidium with deep angular pygidial incision, median pair
of lobes inserted in incision. — Type, Chionaspis (Dinaspis)
diosmae Brain. Versiculaspis MacG.
dd. Pygidium with caudal margin either entirely without lobes
or with serrate or dentate pseudolobes.
e. Pygidium with anus located distinctly cephalad of middle
of pygidium, near its cephalic end.
f. Pygidium with caudal margin entire, without lobes or
pseudolobes; insects never producing galls upon plants.
Protodiaspis Ckll.
ff. Pygidium with caudal margin with pseudolobes, six or
seven subequal serrate pairs; insects producing galls
upon leaves of plants. — Type, Chionaspis fodiens Green.. _
Nudachaspis MacG.
ee. Pygidium with anus located distinctly caudad of middle of
pygidium; three pairs of lobe-like projections with setae at
their distal end; lobe-like projections and lateres distinctly
continuously crenulate; insects living exposed, never pro-
ducing galls. Xanthopthalmus Ckll. & Parr.
TRIBE DIASPIDINl 313
bb. Scale of adult female never with a second exuvia; body always en-
closed in a puparium.
c. Pygidium of adult female with single pair of distinct lobes.— ^
Type, Diaspis (Cryptodiaspis) limuloides Lindgr
Credodiaspis MacG.
cc. Pygidium of adult female without \ohes..Adiscodiaspis Marchal.
SPECIES OF CARULASPIS
a. Pygidium of adult female with two pairs of lobes; caudal margin
truncately rounded; median pair of lobes with distal end rounded,
entire or slightly notched; second pair of lobes incised, mesal lobe-
let much larger than lateral, entire; plates stout, slightly longer than
lobes, arranged 2, 2, 7, those of each lateris arranged 1-1-2-3; lata-
dentes of each second incisura triangular and as long as lobes, two
adjacent to plate at cephalic margin of second pair of lobes, round-
ed, with oraceratubae ; genacerores 5-8(10-16)7-9; mesospiracerores
usually two. — Diaspis carueli Targ., Diaspis juniperi Bouche. — Italy,
England, New York, Massachusetts, District of Columbia on Juni-
perus, arbor-vitae. juniperi Bouche.
aa. Pygidium of adult female with three pairs of lobes,
b. Pygidium with third pair of lobes incised.
c. Pygidium with lobes all entire; median pair of lobes broader
than long, indistinct mesal and lateral notches, bluntly point-
ed between notches; second and third pairs of lobes incised,
lobelets subequal in size and similar in form to lobelets of
second pair; plates slender, arranged 2, 1, 1, 4, those of each
lateris arranged 1-1-1-1; lateres irregularly broadly indented,
crenulate; latadentes bearing oraceratubae in second and third
incisurae; genacerores in four groups. — Pseudoparlatoria
crlstata Lindgr. — Brazil on Gnetum cristata Lindgr.
cc. Pygidium with third pair of lobes minutely serrate, wide
apart, striated, produced, but not conspicuously beyond margin;
second and third pairs of lobes incised; plates spine-like, ar-
ranged 2, 1-2, 1-2, 5, those of lateres arranged 1-1-1-1-1; lata-
dentes caudad of each second and third lobe and two cephalad
of third pair of lobes representing fourth pair of lobes; gena-
cerores 11-11(18-18)8-8. — Diaspis miranda Ckll. — Mexico on
"Cherimoya," "Zapote." miranda Ckll.
bb. Pygidium with third pair of lobes not incised, rounded; lateral
margin oblique, intermediate in size between lobelets of second
pair of lobes; mesal lobelet of second pair of lobes nearly twice
as large as lateral, bluntly rounded, lateral irregularly rounded;
median pair of lobes slightly larger than mesal lobelet of second
pair, bluntly rounded; lobes all entire; plates longer than lobes,
arranged 2, 1, 1, 4, those of each lateris arranged 0-1-2-1; lateres
crenulate and irregularly finely notched; latadentes large in sec-
ond incisurae, smaller in third; anus superimposed over vulva;
genacerores 5-7(14-21)20-25; mesospiracerores 2-1, metaspiracerores
2. — Pseudoparlatoria chilina Lindgr. — Chile on Saxegothaea,
Podocarpus. chilina Lindgr.
314 THE COCCIDAE
SPECTES OF DIASPIDISTIS
Pygidium of adult female with median pair of lobes largest, crenulate;
second and third pairs of lobes incised, lobelets subequal, crenulate;
fourth pair of lobes deeply incised into five subequal crenulate lobe-
lets; each lateris with seven or eight subequal projections or lata-
dentes; plates arranged 2, 1, 1, 0; second, third, and fourth incisurae
with broad lobe-like latadentis bearing oraceratuba; genacerores (15-
28)28-27. — Brazil on Myrtaceae multiloMa Hemp.
SPECIES OF PSEUDOPARLATORIA
a. Pygidium of adult female with median pair of lobes rounded,
minutely serrate, not notched; second pair of lobes divided into
three lobelets; plates of median incisura hardly projecting beyond
lobes; genacerores 2-2(12-15)11-15. — Mexico on Myrtus
serrulata Towns. & Ckll.
aa. Pygidium with median pair of lobes not serrulate.
b. Pygidium with five groups of genacerores, 7-7(20-20)16-18; median
pair of lobes rounded at distal end, sides sloping, subbasal
notches wanting; second and third pairs of lobes incised; plates
of median incisura subparallel, longer than lobes, — Brazil on
Nectandra. noacki Ckll.
bb. Pygidium usually with four groups of genacerores, 0-2(9-15)7-10;
median pair of lobes quadrangular, deep mesal and lateral
notches; second and third pairs of lobes similar, incised, mesal
lobelet largest, lobelets triangular, bluntly pointed, often with
lateral notch; plates arranged 2, 1, 1, 0, those of median incisura
subequal to lobes, those of other incisurae longer than lobes;
each lateris with four slight projections, arranged in pairs,
caudal pair distant from third lobe; anus midway between vulva
and caudal margin, about three times its width from each. —
Florida, Mexico, Brazil on Persea, Drimys, Psidium, Oncidium,
. Magnolia, peach. parlatoroides Comst.
SPECIES OF PSEUDAULACASPIS
a. Pygidium always with two or more pairs of distinct lobes, median
pair large and prominent, second pair always small.
b. Pygidium with more than two pairs of lobes.
c. Pygidium with second and third pairs of lobes incised,
d. Pygidium with median pair of lobes rounded, radiately
striate, moderately large, margins irregularly crenulate; sec-
ond pair of lobes consisting of two large subequal lobelets;
third pair of lobes consisting of two lobelets, mesal small-
est; fourth pair of lobes consisting of two very broad, low
lobelets, mesal low and conical, lateral gently convex and
serrate with five teeth; fifth pair of lobes represented by
two broad low processes; plates arranged 0, 1, 1, 1, 0, 0;
most of described lobes probably latadentes; latadentes of
each second and third incisurae as large as mesal lobelet, of
each third incisura smaller; genacerores 29-30(49-58)21-31. —
China, Japan on Eleagnus. crawii Ckll.
TRIBE DIASPIDINI 315
dd. Pygidium with median pair of lobes with mesal margins
long and straight near proximal end, broadly rounded,
finely crenulate; second pair of lobes incised, mesal lobelet
larger than lateral, distal end of mesal lobelet rounded, of
lateral lobelet pointed; third pair of lobes similar to sec-
ond, smaller; plates long and slender, much longer than
lobes, arranged 0, 1, 1, 8, those of second incisurae furca-
pectinae, those of each lateris arranged 1-7; genacerores
27-31(55-57)31-39.— Natal on Melia, Ricinus fulleri Ckll.
cc. Pygidium with none of lobes incised; median pair of lobes
distant, broadly rounded, entire; second and third pairs of
lobes subquadrangular, margins substraight, distal ends ser-
rate; fourth pair of lobes smaller, subquadrangular, angles of
distal end prolonged, spine-like; plates slightly longer than
lobes, arranged 0, 0, 0, 3, 6, those of each lateris arranged 3-3;
anus located caudad of vulva near caudal margin; setae as
long as lobes; genacerores 8-8(17-20)9-10. — Greece on Pistacia.—
gennadii Leon,
bb. Pygidium with two pairs of lobes.
c. Pygidium with median pair of lobes with proximal end dis-
tinctly wider than distal, never constricted on proximal portion.
d. Pygidium with second pair of lobes short, distinct, incised,
e. Pygidium with genacerores 6-25(14-46)12-38.
f. Pygidium with second pair of lobes small, incised,
mesal lobelet twice or more as long as lateral lobelet;
median pair of lobes much larger, subtriangular, both
margins crenulate, distal end entire or margins rarely
subentire and notched; each lateris with two groups
of bluntly rounded or toothed latadentes, each with an
oraceratuba; minute latadentis in each second incisura;
plates of furcapectinae arranged 0, 1, 10-12, those of
each lateris arranged 1 or 2-3-6 ; anus and vulva super-
imposed; genacerores 6-25(14-46)12-38; mesospiracerores
15-17. — Europe, south eastern United States, West
Indies, Mexico, South America, China, Japan, India,
south Africa, Ceylon, New Zealand, Australia, etc. on
peach, plum, prune, cherry, apricot, walnut, grape, per-
simmon, geranium, etc. pentagona Targ.
ff. Pygidium with second pair of lobes doubly incised, me-
dian lobelet much longer than mesal or lateral; plates
long, one in each second incisura, two cephalad of each
second lobe, about fourteen on each lateris; genacerores
9-9(26-26)16-19. — Japan, California on Osmanthus
pentagona auranticolor Ckll.
ee. Pygidium with genacerores 76-76(51-51)40-50; median pair
of lobes well developed, triangular, rounded at distal end,
slightly serrate; second pair of lobes only slightly de-
veloped; plates strong, arranged 0, 1, 12-14, those of each
lateris arranged 1-2-4-6-6-7; brevaceratubae arranged in
316 THE COCCIDAE
two transverse interrupted rows; latadentes small; meso-
spiracerores present, metaspiracerores wanting. — Ceylon on
Flacourtia. flacourtiae Ruth.
dd. Pygidium with second pair of lobes obscure, rounded, not
incised; median pair of lobes extremely large, widely sepa-
rated, strongly divergent, rounded, finely crenulate; plates
arranged 0, 0, 12, those of each lateris arranged 2-1-1-2-1-2-
(1-1-1) ; anus five times its width from caudal margin and
over six times from vulva; genacerores 24-24(36-36)25-25. —
Arizona on Holocantha. toumeyi Ckll.
CO. Pygidium with median pair of lobes prominent, divergent,
broadly flabelliform, constricted on proximal portion, broadly
rounded and slightly and irregularly notched; second pair of
lobes incised, mesal lobelet prominent, slightly dilated toward
extremity, truncately rounded, lateral lobelet triangular, much
shorter than mesal; plates short on caudal and long on
cephalic portion of pygidium, arranged 0, 1, 10, those of each
lateris arranged 1-2-2- ( 2-2 ) ; anus and vulva probably near
together; genacerores 6-9(18-23)19-23; dorsal oraceratubae
large, numerous, conspicuous. — India on Loranthus
harteri Green.
aa. Pygidium with single pair of distinct, not particularly large lobes,
edges slightly crenulate, lobes almost adjacent at proximal end;
second pair of lobes represented by pair of minute tubercles; three
furcapectinae in each second incisura and about twelve on each
lateris; genacerores 25-25(15-15)7-12. — Mexico on "Chico Sapote."
persimilis Ckll.
SPECIES OF AULACASPIS
a. Body oval or trigonal in outline, cephalic and lateral margins form-
ing continuous broadly rounded curve.
b. Pygidium with vulva and anus superimposed.
c. Pygidium with plates or furcapectinae of second and third
incisurae projecting almost their entire length caudad of
caudal ends of median pair of lobes,
d. Pygidium with seven pairs of altaoeratubae.
e. Pygidium with median pair of lobes serrate and with
plates in incisurae cephalad of median pair of lobes,
f. Pygidium with three continuous series of brevacerat-
ubae; median pair of lobes long, widely divergent,
finely serrate, proximal ends almost adjacent; second
and third pairs of lobes subequal, incised; plates large
and spine-like, arranged 0, 1, 1, 4, those of each lateris
arranged 1-1-1-1; genacerores 11-33(17-40)14-40; meso-
spiracerores 8-12, metaspiracerores 4-6. — Europe, Aus-
tralia, Japan, Hawaiian Islands, Chile, West Indies,
eastern United States, Canada, California on rose, rasp-
berry, strawberry, blackberry, pear, myrtle, Ailanthus,
Cycas, mango, etc. rosae Bouche.
TRIBE DIASPIDINI 317
ff. Pygidium with single irregular series of brevaceratubae,
consisting of about four oraceratubae on each side; me-
dian pair of lobes strongly divergent, free margins ser-
rate; second and third pairs of lobes incised, lobelets
subequal in length, distal ends truncate or slightly
oblique, slightly serrate; plates long and slender, ar-
ranged 0, 1, 1, 4, those of each lateris arranged 1-1-1-1;
genacerores 15-15(16-19)16-17; body with prominent
tubercle at each cephalo-lateral angle. — Aulacaspis cin-
namomi Newst. — Java on Cinnamomum
tuhercularis Newst.
ee. Pygidium with median pair of lobes entire and with fur-
capectinae in incisurae; median pair of lobes oblique,
distant, broadly rounded, entire; second and third pairs
of lobes entire, incised, lobelets similar in form, entire,
broadly rounded, mesal lobelet larger than lateral; plates
and furcapectinae arranged 0, 1, 1, 4, those of incisurae
and one adjacent to each third lobe furcapectinae, cephalic
ones much longer than caudal, plates of lateris long, they
and furcapectinae arranged 1-1-1-1; genacerores 15-15(26-
29)16-17. — Diaspis (Aulacaspis) cinnamomi mangiferae
Newst. — Ceylon, Egypt on mango mangiferae Newst.
dd, Pygidium with nine pairs of altaceratubae, sometimes eight,
extra ones located on cephalic part; median pair of lobes
widely divergent, free margins convex and finely serrate;
second and third pairs of lobes incised, lobelets of each pair
subequal, those of second larger than those of third, distal
margins truncate, margins entire; plates stout, arranged
0, 1, 1, 5-6, those of each lateris arranged l-l-l-2or3; caudal
segment of preabdomen with 4-5 plates on each side; meso-
genacerores 15-16, pregenacerores and postgenacerores forming
continuous group on each side of 18-22 cerores. — Java
javanensis Newst.
cc. Pygidium with plates of second and third incisurae not pro-
jecting caudad beyond caudal end of median pair of lobes or
at most projecting only slightly; median pair of lobes large,
long, projecting strongly, distant, distal end broadly rounded,
margins minutely serrate; second pair of lobes incised, mesal
lobelet twice as long as lateral, elongate, narrow, rounded,
both lobelets entire; third pair of lobes incised, mesal lobelet
long, narrow, twice as long as broader lateral lobelet, margin
of mesal lobelet serrate; plates long, slightly longer than mesal
lobelet of second pair of lobes, arranged 0, 1, 1, 5-6, those of
each lateris arranged l-l-l-3or4; genacerores 8-16(14-25)11-26;
anus and vulva superimposed; thoracic and cephalic abdominal
segments with one and sometimes two pairs of conspicuous
depressed dorsal areas. — Ceylon on Lioranthus. -.loranthi Green,
"bb. Pygidium with vulva located some distance caudad of anus.
c. Pygidium with genacerores 8-10(15-18)20-30; median pair of
lobes comparatively long, divergent, minutely crenulate, fused
318 THE COCCIDAE
at proximal end; second pair of lobes elongate, Incised, lobe-
lets spatulate, mesal lobelet larger than lateral; plates long,
particularly those of lateres, arranged 0, 1, 11, those of each
lateris arranged 1-2-2-6; altaceratubae about seven, their ora-
ceratubae located in latadentes, those of second and third in-
cisura elongate, bluntly pointed, none of the oraceratubae
adjacent; each lateris with three pairs of adjacent, short,
broad, serrate latadentes; rows of oraceratubae of brevacera-
tubae interrupted on meson and divided into mesal and lateral
groups; mesospiracerores forming large cluster. — Ceylon on
Fagraea. fagraeae Green.
CO. Pygidium with genacerores 14-17(28-37)26-56; median pair of
lobes narrowed at proximal end, expanded at disto-lateral
angles and broadly rounded; second pair of lobes incised,
lateral lobelet conspicuous, distinctly rounded at distal end,
similar in shape to mesal lobelet; brevaceratubae in two rows;
mesospiracerores 19-20, metaspiracerores wanting; plates ar-
ranged 0, 1, 11, those of each lateris arranged 1-2-5-3; latadentes
distinct. — Ceylon on Myristica myristicae Ruth.
aa. Body pentagonal in outline, never oval or trigonal, cephalic margin
consisting of three subequal portions, mesal transverse portion and
on each side an oblique portion which joins straight lateral thorac-
ic margin at an angle; pygidium with median pair of lobes oblique,
not or slightly projecting beyond pygidial incision, lateral margins
not free, mesal margin slightly convex and serrate; second and third
pairs of lobes incised, lobelets bluntly rounded, mesal lobelet slightly
longer than lateral; plates of incisurae short, longer than lobes,
distinctly shorter than long plates of lateres, arranged 0, 1, 1, 5,
those of each lateris arranged 1-1-1- (1-1) ; latadentes in second and
third incisurae nearly as large as lateral lobelets; genacerores in
five groups. — Java on Ilex penzigi Leon.
SPECIES OF EPIDIASPIS
Pygidium with median pair of lobes large, broad, rounded, faint lateral
notch; second, third, and fourth pairs of lobes small, rudimentary,
incised, mesal lobelet largest, all probably latadentes; dorsal setae
cephalad of second pair of lobes surrounded by densaria-like thicken-
ings; brevaceratubae few in number, elongate, irregularly arranged;
plates short, broad, arranged 0, 0, 1, 1, 6, those of each lateris ar-
ranged 1-1-1-2-1; anus eight times its width from caudal margin and
once caudad of vulva; genacerores (12-16)9-10. — Diaspis leperii Sign.,
Epidiaspis piricola Del Guer. — Italy, France, Portugal, Germany, Eng-
land, California, New York on apple, plum, peach, pear, currant
pyri Colvee.
SPECIES OF COCKERELLASPIS
Pygidium with median pair of lobes distant, rounded, very low, not
oblique or divergent, height above margin less than half their breadth;
second and third pairs of lobes small, rounded, nearly obsolete; plates
small, spine-like, arranged 0, 0, 1, 7, those of each lateris arranged
l-(l-l)-l-3; anus longer than broad, located sixteen times its width
TRIBE DIASPIDINI 319
from caudal margin and about five times from vulva; genacerores
5-7(13-14)8-8. — New Mexico on Quercus montanus Ckll.
SPECIES OF DIASPIS
a. Body of adult female with large, blunt, button-like projection or
tubercle on each side on level with rostrum.
b. Pygidium with an altaceratubae opening in mesal latadentis, with
four marginal altaceratubae opening in latadentes on each side
and two submarginal altaceratubae; median pair of lobes strong-
ly divergent, oblique, proximal ends not fused, finely serrate;
second and third pairs of lobes small, short, incised, lateral
lobelet usually larger than mesal; plates comparatively long, spine-
like, arranged 0, 1, 1, 7, those of each lateris arranged 1-1-1-3-1,
cephalic four much broader at proximal end than others; gena-
cerores 8-12(6-25)7-20; mesospiracerores about 18, subdivided into
groups. — Diaspis cattleyae Ckll. — Europe, Australia, Hawaiian
Islands, America on Acacia, Cattleya, palms, wattle, various hot-
house plants. doisduvalii Sign.
bb. Pygidium not with an altaceratuba opening in mesal latadentis,
with five marginal altaceratubae opening in latadentes on each
side and two submarginal latadentes, two large adjacent breva-
ceratubae located cephalad of median incisura; median pair of
lobes subtriangular, unsymmetrical, divergent, mesal margins
serrate; second and third pairs of lobes incised, lobelets sub-
equal with truncated ends with rounded angles, entire, mesal
lobelet slightly larger than lateral; cephalad of each third lobe
a lobe-like latadentis and cephalad of this a triangular projection,
long oblique side, serrate; plates caudad of serrate projection
much shorter and narrower on proximal portion than those
cephalad, arranged 0, 1, 1, 7, those of each lateris arrangedl-(l-l)-
l-(l-l-l); genacerores 0-8(5-12)4-8.— South Africa on Connarum,
Syzygium. africana Lindgr.
aa. Body of adult female not with large, blunt, projection on each side
on level with rostrum.
b. Pygidium with median pair of lobes distinctly larger than sec-
ond pair of lobes or at least than second lobelet, if incised,
c. Pygidium with median pair of lobes entire or at most only
slightly crenulate, never with mesal margins serrate,
d. Pygidium with plates in second and third incisurae.
e. Pygidium with median and second pairs of lobes wholly
entire.
f. Pygidium with genacerores arranged 9-15(12-23)10-24;
median pair of lobes with lateral margins free, mesal
and lateral margins entire, not strongly oblique; sec-
ond and third pairs of lobes subequal, incised, mesal
lobelet always much larger than lateral, lobelets point-
ed, rounded, entire; plates short, arranged 0, 1, 1, 8,
those on each side of third pair of lobes longest, those
on each lateris short and broad at proximal end, all
320 THE COCCIDAE
distant; oraceratubae of lataceratubae located in lata-
dentes; brevaceratubae small and numerous; anus lo-
cated nearly midway between vulva and caudal margin,
nearer vulva; mesospiracerores about 6. — Diaspis
echinocacti Bouche, Diaspis echinocacti cacti Comst.,
Diaspis echinocacti opuntiae Ckll. — Europe, northern
Africa, India, eastern United States, New Mexico,
Arizona, Mexico, West Indies, Brazil on Opuntia,
Echinocactus, Cereus. calyptroides Costa.
ff. Pygidium with genacerores arranged 11-11(10-10)3-4;
median pair of lobes close together, low, evenly round-
ed, entire; second and third pairs of lobes represented
by extremely small, colorless, pointed prominences;
plates large, two on each side especially large and
broad with fimbriated ends, pectinae; anus distant from
vulva; latadentes bearing oraceratubae at proximal
ends of lobes, closely adjacent pair beneath median pair
of lobes, and three or four other pairs at intervals along
margin. — Mexico on mistletoe phoradendri Ckll.
ee. Pygidium with lateral margin of median pair of lobes
and margin of mesal lobelet of second pair serrate; me-
dian pair of lobes with mesal margins straight, entire,
lateral margins feebly rounded, oblique, only small por-
tion free, finely serrate; second pair of lobes incised,
mesal lobelet broader than long, strongly rounded, very
finely serrate, lateral lobelet elongate, rounded, rudimen-
tary; third pair of lobes incised, mesal lobelet large,
unsymmetrical, straight inner and oblique outer mar-
gins, lateral lobelet similar, small, rudimentary; plates
of second and third incisurae only slightly longer
than lobes, others larger, arranged 0, 1, 1, 6, those of each
lateris arranged 1-2-3, distal portion of plates suddenly
reduced in size; genacerores 10-20(14-19)8-9. — Syria on
Pistacia. syriaca Lindgr.
dd. Pygidium with furcapectinae in second and third incisurae;
median pair of lobes wide apart, large, mesal margins oblique,
entire or slightly crenulate, lateral margins only partly
free; second pair of lobes incised, short; third pair of lobes
incised; latadentis cephalad of each second and third lobe
and two caudad of third lobe; plates and pectinae arranged
0, 1, 1, 22, those of each lateris arranged 2-3 and others
uniformly distributed cone-like plates, furcapectinae adja-
cent to each third lobe and about twenty plates cephalad of fur-
capectinae; caudal segment of preabdomen with twenty-two
plates on each side; oraceratubae numerous; mesospira-
cerores and metaspiracerores 20-25; genacerores 15-28(17-45)
17-32. — Brazil on Myrtaceae australis Hemp.
cc. Pygidium with median pair of lobes with mesal margins serrate.
d. Pygidium with second pair of lobes not incised.
TRIBE DIASPIDINI 321
e. Pygidium with second pair of lobes represented by long
bluntly pointed projections; median pair of lobes distant,
strongly divergent, mesal margins serrate, caudal end
projecting far beyond caudal margin, lateral margin free
and entire; third pair of lobes wanting or represented
by two low rounded protuberances; pointed projection
between median and second lobe probably latadentis;
plates shown arranged 0, 0, 1, 1; anus located distinctly
caudad of vulva; genacerores 10-10(20-20)9-9. — Mexico on
Prosopis. townsendi Ckll.
ee. Pygidium with second pair of lobes subround, each rather
less than one-half width and length of median lobe, dis-
tant by about its own width from median lobe; median
pair of lobes largest, almost imperceptibly excavated on
distal portion of lateral margin, very slightly notched on
distal portion of mesal margin; third pair of lobes about
one-half size of second pair, appearing as tubercles on
margin, each distant from second lobe rather more than
twice its width; with projections identified as fourth and
fifth pairs of lobes; plates not large, not long, equaling
in length median pair of lobes; anus distant from vulva;
genacerores 15-15(30-30)20-20. — Mexico on Baccharis
baccharidis Towns. & Ckll.
dd. Pygidium with second pair of lobes always deeply incised.
e. Pygidium with one plate in each second incisura, each
side with eight plates.
f. Pygidium with one plate in each third incisura.
g. Pygidium with five marginal, one mesal, and two
submarginal oraceratubae of altaceratubae on each
side; median pair of lobes with lateral margins not
free, mesal margins serrate, oblique; second and
third pairs of lobes subequal, incised, entire, distal
ends of lobelets broadly rounded; plates short, ar-
ranged 0, 1, 1, 6, those of each lateris distant, ar-
ranged l-( 1-1) -1-1-1, cephalic plates much broader
than caudal; marginal oraceratubae of altaceratubae
located in latadentes, two latadentes on each lateris,
located between groups of plates; genacerores
8-10(14-20)13-17. — Europe, eastern United States,
California, Mexico on Bromelia, Hibiscus, Canna,
Bilbergia, Olea,, pineapple, ivy bromeliae Kern.
gg. Pygidium with three or four marginal and sub-
marginal rows of oraceratubae of altaceratubae on
each side, two caudal longitudinally elongate and
three cephalic small and round; median pair of
lobes distant, oblique, mesal margin long and con-
vex, with five distinct serrations including caudal
one, lateral margin with single serration; second
pair of lobes very small and low, incised; third pair
of lobes low, broad, trifid, similar to three mar-
322
THE COCCIDAE
bb.
ginal serrations; plates of second incisurae sub-
equal in length to median pair of lobes, others larger,
arranged 0, 1, 1, 6, those of each lateris arranged
1-1-1- (1-1-1); genacerores 8-8(16-16)10-10.— Texas on
Sophora. texensis Ckll.
ff. Pygidium without plates in each third incisurae, plates
arranged 0, 1, 0, 4, those of each lateris arranged
1-(1-1)-1, subequal in length to lobes; latadentes lo-
cated in median, second, and third incisurae, and three
on each lateris; median pair of lobes longer than broad,
serrate, mesal margin oblique; second and third pairs
of lobes incised, lobelets entire, mesal lobelet larger
than lateral, distal end of mesal round, of lateral sub-
truncate; anus located nearly midway between vulva
and caudal margin; each lateris with lobe-like projec-
tions, caudal one serrate; genacerores (18-23)16-20. —
Senegal, Africa on Khaya senegalensis Vays.
ee. Pygidium not with plate in each second incisura, plates
short, hardly longer than lobes, arranged 0, 0, 1, 9-13, those
of each lateris arranged l-3or4-6or9; median pair of
lobes largest, oblique, serrate; second and third pairs of
lobes subequal and subsimilar, incised, mesal lobelet three
or more times as large as lateral; five altaceratubae on
each side and similar mesal one; lateris irregular and
lobe-like opposite cephalic oraceratubae of fourth and
fifth altaceratubae; anus about midway between caudal
margin and vulva, over six times its width from caudal
margin; genacerores 6-10(9-16)6-10. — California on Man-
zanita, Acrostaphylos. manzanitae Whitney.
Pygidium with median pair of lobes distinctly smaller than mesal
lobelet of second pair of lobes or of second pair if not incised,
if apparently subequal, cephalic end of body with blunt mesal
plate-like structure.
c. Pygidium with second pair of lobes not incised, as broad as
long, broadly rounded, entire, projecting farther caudad than
median pair; median pair of lobes small, subtriangular, distal
portion notched; third pair of lobes wanting; plates longer
than lobes, arranged 0, 1, 8, those of each lateris arranged
2-1-1-4; oraceratubae in three groups, located between groups
of plates of each lateris; dorsal setae large; genacerores 8-8
(12-13)8-8. — Canary Islands on Juniperus. atlantica Lindgr.
cc. Pygidium with second pair of lobes deeply incised.
d. Body with cephalic margin with single, short, conical, trun-
cate, plate-like, mesal projection; median pair of lobes dis-
tant, divergent, lateral margin straight, mesal margin
oblique and serrate, subequal to or slightly larger than
mesal lobelet of second pair; second pair of lobes incised,
entire, truncate with rounded angles, mesal lobelet slightly
larger than lateral; third pair of lobes incised, similar in
form to second pair, smaller, lateral lobelet sometimes with
TRIBE DIASPIDINI 323
lateral notch; plates and pectinae arranged 0, 1, 1, 7, each.
lateris with caudal and cephalic projections, plates, others
furcapectinae, arranged 1-1-1-1-1-1-1; marginal altaceratubae
four, opening in latadentes, located in second and third in-
cisurae and cephalad of caudal plate and between third and
fourth from caudal end; margin where fourth pair of lobes
would be located, not produced but serrate; genacerores
4-4(10-10)8-8; pygidium of second nymphal female similar to
adult. — South Africa on Strychnos stilosa Lindgr.
dd. Body with meson of cephalic margin never with mesal plate-
like projection; pygidium with median pair of lobes minute,
subtriangular, divergent, mesal margin sometimes notched;
second pair of lobes incised, mesal lobelet larger than lat-
eral, both broadly rounded, entire; third pair of lobes simi-
lar in form and size, incised, entire; plates stout, longer
than lobes, arranged 0, 1, 1, 6, those of each lateris broader
at proximal end, arranged l-( 1-1) -1-1-1; anus located caudad
of vulva; altaceratuba opening in mesal latadentis, four
marginal altaceratubae opening in latadentes; mesospira-
cerores 0-1; genacerores 7-9(13-16)11-14; pygidium of second
nymphal female with two pairs of lobes, median pair plate-
like, strongly divergent. — German East Africa on Loranthus.
parva Lindgr.
SPECIES OF UMBASPIS
Scale of adult female circular, yellowish or creamy white in color,
exuviae tilted; body pyriform; pygidium with median pair of lobes
distant, small, projecting, meso-distal angles rounded, entire; second
and third pairs of lobes incised, lobelets small, truncately rounded,
subequal, entire; one lobe-like projection in each second incisura and
three in third; single plate in third incisurae adjacent to third lobe
and two cephalad of it; prominent spine-like projection just cephalad
of cephalic plates, two plates adjacent to its cephalic margin and
three others equidistant from each other and two plates adjacent to
spine-like projection; altaceratubae arranged, one in median and each
second incisura, two in each third incisura, and two associated with
each third pair of lobes, and one with each spine-like projection; meso-
spiracerores 0-1; genacerores 8-11(16-19)13-16. — Uganda, Africa
regularis Newst.
SPECIES OF RUTHERFORDIA
Pygidium with median pair of lobes fused, forming single lobe, promi-
ment, broader than long, sides parallel on proximal portion, converg-
ing on distal portion, rounded at distal end; incisura cephalad of me-
dian lobe short, heavily chitinized, two sides fused; plates numerous,
long, those of each lateris in three groups, separated by minute, point-
ed, hyaline, lobe-like projections, arranged 11-12-11, cephalad of these
plates, four groups of similar plates, decreasing in size toward cephalic
end; genacerores in jfive groups, very numerous, 100 mesogenacerores ;
mesospiracerores and metaspiracerores present; brevaceratubae nu-
merous.— Ceylon on Mallotus malloti Ruth.
THE COCCIDAE
SPECIES OF PROTARGIONIA
Scale of adult female oval, exuviae located on one side near periphery;
pygidium with median pair of lobes short, projecting only slightly,,
bluntly pointed, faint notches or few crenulations; second pair of lobes
hardly projecting, lateral margin oblique; described third pair of lobes
not projecting, bearing seta; altaceratubae large, their oraceratubae
one in each second incisura, two in third, and two, more distant,
cephalad of rudimentary lobes; plates wanting; spiracerores wanting;
genacerores 2-2(9-12)7-11; anus three times its width from caudal
margin and distant from vulva. — Argentina on Larrea larreae Leon.
SPECIES OF ESSIGASPIS
Pygidium with lobes, pectinae, and plates wanting; margin broadly
undulate; oraceratubae of brevaceratubae very small, numerous, par-
ticularly near caudal margin, promiscuously arranged; anus twelve
times its width from caudal margin and four times cephalad of vulva;
genacerores arranged in almost continuous U-shaped arch, consisting
of 20-30 cerores. — California on Quercus agrifoliae Essig.
SPECIES OF SCLOPETASPIS
a. Pygidium with second pair of lobes not incised; median pair of
lobes short, projecting, notched, distant; second pair of lobes simi-
lar in form, broader than median pair; third pair of lobes incised,
lateral lobelet broader than mesal, lateral margin oblique, dentate,
disto-mesal angle prolonged; fourth pair of lobes incised, similar to
third pair, broader, distal margin dentate; plates small, inconspic-
uous, arranged 0, 1, 2, 2, 3; brevaceratubae in broad interrupted
bands; oraceratubae numerous on preabdomen; anus longer than
broad, about twenty-four times its width from caudal margin, seven
times cephalad of vulva, vulva located on the line drawn between pre-
genacerores and postgenacerores; genacerores 15-15(30-30)60-60; meso-
spiracerores 10, metaspiracerores 4-5; scale covered with white felt-
ed woolly wax. — Uganda, Africa on Loranthus laniger Newst.
aa. Pygidium with second pair of lobes incised, lobelets subequal in size,
prominent, projecting, distal ends rounded, entire; median pair of
lobes prominent, distant, similar to mesal lobelet of second pair,
larger; third and fourth pairs of lobes incised, similar to each
other and to second pair, entire; plates small, arranged 0, 1, 1, 1, 5,
those of each lateris arranged 1-4; mesospiracerores and metaspira-
cerores numerous, about twenty; brevaceratubae arranged in three
prominent interrupted rows; margin with three large altaceratubae
on each side; genacerores arranged in five large groups; anus oppo-
site vulva, vulva located slightly caudad of middle of pygidium;
lateral margin of body dilated, forming tubular extension, bearing
eye-like spot; cephalic portion of body with numerous chitinized
patches. — Java, Uganda, Africa on sugar-cane
madiunensis Zehntner.
SPECIES OF CHIONASPIS
a. Pygidium with third pair of lobes not incised.
b. Pygidium with second pair of lobes not incised, broader than
long, with or without slight notches, entire; median pair of lobes
TRIBE DIASPIDINI 325
broad, broader than long, entire, distant; third pair of lobes tri-
angular, almost obsolete, lateral margin long, oblique, entire;
plates longer than lobes, arranged 0, 2, 3, 11, those of each lateris
arranged (2-2-2) -4-1; setae small; dorsal oraceratubae arranged in
three broad uninterrupted bands; latadentes in second and third
incisurae; genacerores 16-19(17-33)13-29. — Algeria on Nerium,
Oleander. nerii Newst-
bb. Pygidium with second pair of lobes incised, mesal lobelet elongate,
broadly bluntly rounded, one-half width of median lobe, lateral
lobelet shorter and narrower than mesal; median pair of lobes
adjacent at proximal end, divergent caudad, longer than broad,
distal end bluntly rounded; third pair of lobes not incised, simi-
lar in form to mesal lobelet of second pair; plates about as long
as or shorter than median pair of lobes, arranged 0, 2, 2, 8-9,
those of each lateris arranged 2-2-4to5; anus located between pre-
genacerores; each lateris with seven altaceratubae ; dorsal ora-
ceratubae small, arranged in rows; genacerores 12-13(20-29)18-22.
— China on Populus. micropori Marl.
aa. Pygidium with both second and third pairs of lobes incised.
b. Pygidium with two plates in each second incisura, only rarely
with one, if so, each lateris with plates arranged in two groups.
c. Pygidium with genacerores arranged 15-20(30-40)25-30; median
pair of lobes triangular, pointed, not adjacent, divergent, en-
tire; second pair of lobes large, smaller than median, incised,
mesal lobelet larger than lateral, bluntly rounded; third pair
of lobes small, rudimentary; plates twice as long as median
pair of lobes, mesal plate of each group shortest, arranged
0, 2, 2, 6, those of each lateris arranged 2-2-2; submarginal
setae distinct, marginal setae prominent; anus and vulva su-
perimposed, seven times its width from caudal margin. — Massa-
chusetts, California on Spartina spartinae Comst.
cc. Pygidium with genacerores always much less in number than
15-20(30-40)25-30.
d. Pygidium with median pair of lobes finely serrate, large,
longer than broad, bluntly pointed, proximal ends adjacent,
divergent caudad; second and third pairs of lobes incised,
mesal lobelet largest, distal ends bluntly rounded and
toothed; latadentes in second and third incisurae; plates
slender, longer than median pair of lobes, not twice as long,
arranged 0, 2, 2, 8-9, those of each lateris arranged 2-2or3-4;
caudal preabdominal segment with four or five plates on
each side; anus located distinctly cephalad of vulva; gena-
cerores 4-6(5-8)2-7. — New York, Virginia, Georgia, Ohio, Eng-
land, France on Euonymus euonymi Comst.
dd. Pygidium with median pair of lobes entire, divergent, dis-
tance between them nearly equal to width of median lobe,
broadly rounded; second pair of lobes incised, lobelets
large, mesal larger than lateral, slightly longer than broad,
entire, broadly rounded; third pair of lobes incised, lobelets
subequal in width, mesal larger than lateral, distinctly long-
326 THE COCCIDAE
er than broad, entire, lateral margin oblique; plates muchi
longer than lobes, arranged 0, 1-2, 1-2, 4-5, those of each
lateris in two groups, arranged 1 to 2, 3; dorsal oraceratubae
numerous, arranged in rows; genacerores 7-12(18-24)15-17. —
South Africa on Stanotophrum stanotophri Cooley.
bb. Pygidium always with less than two plates in each second in-
cisura; each lateris with plates arranged in two groups.
c. Pygidium with one plate in each second incisura.
d. Pygidium with median pair of lobes not adjacent at proximal
end.
e. Pygidium with median pair of lobes as broad as long,
broadly truncately rounded, entire, separated by distance
equal to about one-fourth width of median lobe; second
and third pairs of lobes incised, mesal lobelets largest,
lobelets about as broad as long, broadly rounded, entire;
latadentes in second and third incisurae; plates one-fourth
longer than lobes, arranged 0, 1, 1, 4-7, those of each lat-
eris arranged l-l-lor2-lor3; dorsal oraceratubae not
numerous; anus located slightly cepahlad of vulva; gena-
cerores 7-13(12-21)14-19.— United States, Canada on Pinus,
Pseudotsuga, Abies, Torreya pinifoUae Fitch.
ee. Pygidium with median pair of lobes distinctly longer than
broad.
f. Pygidium with median pair of lobes subquadrangular,
mesal and lateral margins subparallel, distal ends
bluntly pointed at middle with mesal portion minutely
serrate and lateral portion notched; second and thir^*
pairs of lobes incised, mesal lobelet slightly larger than
lateral, entire, bluntly pointed; plates long, arranged
0, 1, 1, 7, those of each lateris arranged 1-1-5; lateres
notched, each with four latadentes bearing oraceratubae;
dorsal oraceratubae arranged in rows; anus its own
width cephalad of vulva; genacerores 12-19(35-40)27-35.
— Ceylon on Polygonum polygoni Cooley.
ff. Pygidium with median pair of lobes triangular or coni-
cal, mesal and lateral margins convergent caudad and
pointed.
g. Pygidium with seven altaceratubae on each side;
lobes crowded onto subtruncate caudal portion;
median pair of lobes pointed, distant, not oblique;
second and third pairs of lobes incised, lobelets sub-
equal, pointed, nearly as large as median pair of
lobes; plates stout, arranged 0, 1, 1, 6-7, those of
each lateris arranged l-l-4or5, plates between lobes
much shorter and smaller than others; genacerores
15-25(18-37)14-22; mesospiracerores crowded groups;
anus superimposed over vulva or slightly caudad. —
Ceylon on Vitis, Elaeagnus, Loranthus vitis Green.
TRIBE DIASPIDINI 327
SS' Pygidium with eight altaceratubae on each side;
lobes normal in position, not crowded onto l3ub-
truncate caudal portion; median pair of lobes
slightly divergent, bluntly pointed or rounded, long;
second and third pairs of lobes incised, second pair
slightly larger than third, lobelets all similar in
form, distal end bluntly rounded, mesal lobelet larg-
er than lateral; plates long, much longer than lobes,
^ arranged 0, 1, 1, 5-6, those of each lateris arranged
l-l-3or4; genacerores 14-16(22-28)15-20; mesospira-
cerores and metaspiracerores present; anus superim-
posed over vulva or slightly cephalad. — Ceylon on
Hedyotis. hedyotidis Green.
dd. Pygidium with median pair of lobes adjacent at least at
proximal end.
e. Pygidium with median pair of lobes as long as or longer
than broad.
f. Pygidium with four to nine plates on each lateris, rare-
ly with only three.
g. Pygidium with mesal margins of median pair of
lobes straight, parallel, adjacent, distal margins
rounded, sides obscurely serrate; second and third
pairs of lobes incised, mesal lobelet of each lobe
three times size of lateral lobelet, entire or obscurely
serrate; plates not long, arranged 0, 1, 1-2, 7-9, those
of each lateris arranged 2-2-4-5; anus located dis-
tinctly cephalad of vulva; dorsal oraceratubae ar-
ranged in transverse rows; genacerores 10-25(18-35)
16-24. — Massachusetts, Ohio, Missouri, Kansas, Ne-
braska, New Mexico, southern California on willow,
Cottonwood, poplar, honey-locust, butternut
ortholoMs Comst.
gg. Pygidium with mesal margins of median pair of
lobes distinctly divergent distad.
h. Pygidium with genacerores arranged 37-37(41-43)
43-43; median pair of lobes large, adjacent at
proximal end, strongly divergent caudad, mesal
and lateral margins of each lobe convergent, meet-
ing in right angle, entire or slighly notched; sec-
ond and third pairs of lobes incised, mesal lobe-
lets of third pair larger than lateral; plates of
second incisura shorter than lobes, others longer,
arranged 0, 1, 2-3, 11-12, those of each lateris ar-
ranged 3-4-4or5; each lateris with three small
pyramidal projections between two caudal groups
of plates; dorsal oraceratubae conspicuous, ar-
ranged in rows. — Japan on Elaeagnus
difficilis Ckll.
hh. Pygidium with genacerores never more than
19(44)30.
328 THE coccroAE
i. Pygidium with brevaceratubae of first and sec-
ond rows wanting, those of third and fourth
rows numerous; median pair of lobes elongate,
tapering, bluntly pointed, serrate; second pair
of lobes incised, mesal lobelet elongate, more
than twice as long as lateral, lateral margin
oblique, serrate, lateral lobelet bluntly round-
ed; third pair of lobes rudimentary, incised,
lobelets broader than long, lateral margins
' oblique, serrate; plates stout, arranged 0, 1, 1,
5-6, those of each lateris arranged l-l-3or4;
anus located slightly cephalad of vulva; gena-
cerores 4-10(12-21)8-14.— Ohio on Gleditsia
gleditsiae Saunders.
ii. Pygidium with brevaceratubae of first or second
rows or both rows present.
j. Pygidium with median pair of lobes with
distal portion bluntly pointed or obscurely
trilobed, faintly serrate; second and third
pairs of lobes incised, mesal lobelet much
larger than lateral, bluntly rounded, faintly
serrate ; plates long, slender, arranged 0, 1,
1-2, 9-14, those of each lateris arranged 2-1
or3-6or9, usually with one plate in each
third incisura and two on lateris adjacent
each third lobe; brevaceratubae 3-6 in mesal
group of second row, 4-6 in mesal group and
5-7 in lateral group of third row, 6-8 in mesal
group and 8-10 in lateral group of fourth row;
anus located cephalad of vulva; genacerores
11-19(25-42)19-28. — Canada, Massachusetts.
New York on Alnus, Salix, Viburnum, Cornus,
Benzoin, Amelanchier, Betula, Dirca, etc
lintneri Comst.
jj. Pygidium with median pair of lobes with dis-
tal portion broadly bluntly rounded.
k. Pygidium with brevaceratubae with mesal
group of second row 7, third row in two
groups, 9 lateral and 7 mesal, fourth row
in three groups, 13 lateral, 4 median and
3 mesal, fifth row in two groups, 11 lat-
eral and 5 mesal; median pair of lobes
prominent, sides subparallel, distal ends
bluntly rounded; second pair of lobes in-
cised, mesal lobelet longer and broader
than lateral, less than one-half width of
median lobe; third pair of lobes incised,
mesal lobelet distinct, lateral lobelet small,
obscure; plates slightly longer than median
pair of lobes, arranged 0, 1, 2, 9-10, those
TRIBE DIASPIDINI 329
of each lateris arranged 2-2-5or6; anus
located near middle of length of pregena-
cerores; genacerores 13-13(24-25)16-22.—
California on orange. --sassceri Ckll. & Robs.
kk. Pygidium with brevaceratubae with first
row wanting, second row with one group,
mesal 2-6, third row with two groups, lat-
eral 5-9 and mesal 4-9, fourth row in two
groups, lateral 3-9 and mesal 3-8; median
pair of lobes as broad as long, serrate, dis-
tal end bluntly rounded; second and third
pairs of lobes incised, mesal lobelet much
larger than lateral, serrate; plates long,
slender, arranged 0, 1, 1-2, 5-11, those of
each lateris arranged lor2-lor2-3or7, usually
with two plates in each third incisura
and with two on each lateris adjacent to
third lobe; anus located about its own
width cephalad of vulva; genacerores 11-23
(21-44) 18-33.— Europe on Salix, Populus,
Alnus, Cornus, Acer, Vaccinium, Tilia,
Sorbus, Fraxinus, Sarcolthamnus
salicis Linn,
ff. Pygidium with two to three plates on cephalic part of
each lateris.
g. Pygidium with median pair of lobes with mesal mar-
gins divergent from proximal end, lobes large,
broadly bluntly rounded, serrate; second pair of
lobes incised, lobelets broadly bluntly rounded,
mesal lobelet more than twice as long as lateral,
margins serrate; third pair of lobes incised, mesal
lobelet much longer than lateral, oblique, lateral
lobelet inconspicuous, margins serrate; plates long
and stout, arranged 0, 1, 2, 4-7, those of each lateris
arranged Ior2-lor2-2or3, cepahalic plates longer
than caudal; dorsal oraceratubae arranged in rows;
genacerores 10-16(20-24)10-17. — Texas on cotton wood.
longiloba Cooley.
gg. Pygidium with median pair of lobes with mesal mar-
gins contiguous on proximal half and divergent on
distal half, large, prominent, distal and lateral mar-
gins broadly rounded, crenulate; second and third
pairs of lobes incised, lobelets wedge-shaped, broad
end directed distad, mesal lobelets larger than lateral,
margins crenulate; plates of incisurae only slightly
longer than lobes, arranged 0, 1, 1, 4-5, those of each
lateris arranged l-l-2or3 ; anus its own width caudad of
vulva; dorsal oraceratubae arranged in rows; alta-
ceratubae six on each side; genacerores 8-18(15-20)8-
18. — Ceylon on Elaeagnus. elaeagni Green.
m
THE COCCIDAE
ee. Pygidium with median pair of lobes distinctly broader than
long.
f. Pygidium with genacerores arranged 18-36(31-47) 28-39.
g. Pygidium with median pair of lobes strongly oblique,
adjacent only for short distance at proximal end,
divergent, entire, bluntly pointed at caudal end; sec-
ond pair of lobes incised, mesal lobelet largest, each
bluntly pointed; third pair of lobes smaller than sec-
ond, incised, mesal lobelet largest; plates about
twice as long as lobes, arranged 0, 1-2, 1-2, 5-6, those
of each lateris arranged 1-1- ( 1-1-1-1 ) ; dorsal oracera-
tubae conspicuous, arranged in rows; altaceratubae
opening in latadentes, 7 on each side; anus about its
width cephalad of vulva; spiracerores conspicuous;
genacerores 18-21(33-47)31-39. — Ceylon on Panicum,
Ischaemum, Ophismenus, other grasses, herdae Green,
gg. Pygidium with median pair of lobes not strongly
oblique, mesal margins adjacent for proximal half of
length, broad, short, broadly rounded, entire or
faintly serrate; second pair of lobes large, incised,
mesal lobelet twice as large as lateral, margins
broadly rounded, entire or faintly serrate; third pair
of lobes distinct, incised, mesal lobelet larger than
lateral, both broader than long, margins broadly
rounded, entire or faintly serrate; plates long and
slender, arranged 0, 1, 1-2, 7-11, those of each lateris
arranged lor2-lor3-5or6; dorsal oraceratubae con^
spicuous, arranged in rows; genacerores 21-36(31-
45)28-32. — United States, Canada on willow, poplar,
Cornus, Amelanchier, Ceanothus, Liriodendron
salicis-nigrae Walsh.
If. Pygidium with genacerores arranged 7-16(11-32)7-22.
g. Pygidium with genacerores arranged 2-7(7-11)6-13,
usually about 4-4(10-10)6-8; median pair of lobes
spatuliform, adjacent, scarcely divergent, entire;
second pair of lobes incised, mesal lobelet like
median, lateral smaller; third pair of lobes incised,
mesal lobelet broad, notched, oblique, margin round-
ed, rudimentary, lateral lobelet similar, small;
plates of second incisurae one-half length of median
pair of lobes, others longer, arranged 0, 1, 1, 5, those
of each lateris arranged 1-1-3, one and one-half times
length of caudal plates; lateres with prominent
latadentes. — Austria on Pinus austriaca Lindgr.
gg- Pygidium with at least some of groups of genacerores
more numerous than 7(10)8.
h. Pygidium with second row of brevaceratubae
wanting, lateral group sometimes represented by
about two pores, third and fourth rows repre-
sented; median pair of lobes well developed, en-
TRIBE DIASPIDINI 331
tire, broadly rounded, mesal margins straight and
adjacent; second pair of lobes entire, incised, mesal
lobelet much larger than lateral, lateral mar-
gin oblique, lateral lobelet triangular; third pair
of lobes entire, incised, lobelets similar in shape,
smaller than second pair; plates long, slender,
arranged 0, 1, 1, 5-6, those of each lateris ar-
ranged l-l-3or4; genacerores 7-9(11-16)7-20. —
New Zealand on Dysoxylon, Hoheria, Melicytus.__
dysoxyli Mask,
hh. Pygidium with second row of brevaceratubae pres-
ent, lateral group with 2-5, mesal group sometimes
wanting.
i. Pygidium with median pair of lobes with proxi-
mal portion of mesal margins straight, adja-
cent, distal portion longer than proximal,
straight and strongly oblique, lateral margin
rounded, margins entire or finely serrate; sec-
ond pair of lobes incised, mesal lobelet much
larger than lateral, more than twice as long,
both broadly rounded, entire; third pair of
lobes incised, similar but smaller than second
pair; plates long, slender, arranged 0, 1, 1,
6-10, those of each lateris arranged lor2-lor
2-4or6; dorsal oraceratubae arranged in rows;
anus located about its own width cephalad of
vulva ; genacerores 9-15 ( 17-28 ) 9-17. — Massachu-
setts, New York, Ohio on Cornus.- -corni Cooley.
li. Pygidium with median pair of lobes broadly
rounded, proximal portion of mesal margin near
but not adjacent and caudal portion strongly
divergent and forming with distal and lateral
margins oroad curve; second pair of lobes in-
cised, mesal lobelet three times as long as lat-
eral, both broadly rounded, margins entire or
finely serrate; third pair of lobes incised, short,
lobelets much broader than long, distal mar-
gins truncate or roundly truncate, serrate;
plates long, slender, arranged 0, 1, 1, 6-11, those
of each lateris arranged l-l-4or9, plate in
each second incisura small, shorter than median
lobe; dorsal oraceratubae arranged in rows;
anus located its own width cephalad of vulva;
genacerores 7-16 ( 22-32 ) 16-22. — United States,
Canada, England on apple, pear, cherry,
quince, peach, choke-cherry, walnut, mountain
ash, elm, Pyrus, currant, hawthorn
furfura Fitch.
cc. Pygidium without plates in each second incisura, plates ar-
ranged 0, 0, 2, 7, those of each lateris arranged 2- (1-2) -(1-1),
THE COCCIDAE
plates only slightly longer than median pair of lobes; median
pair of lobes not adjacent, quadrangular, entire; second pair
of lobes incised, lobelets small, lateral lobelet twice or more
as large as mesal; third pair of lobes incised, loblets subequal,
smaller than lateral lobelets of second pair; anus located
slightly caudad of vulva; mesospiracerores 3, metaspiracerores
2; genacerores 15-17(23-30)10-16.— Italy on Tamarix
etrusca Leon.
SPECIES OF DUPLACHIONASPIS
a. Pygidium with second pair of lobes not incised.
b. Pygidium with median pair of lobes distinctly divergent.
c. Pygidium with each median pair of lobes twice as broad as
second lobe, caudal portion expanded, malleiform, appearing
notched on middle of mesal margin, entire; second pair of
lobes simple, promient, narrow, bluntly pointed, entire;
plates well developed, plates arranged so far as shown, none in
median incisura, one in second, one cephalad of each second
lobe, another about as far cephalad as distance between two
caudal plates; anus located cephalad of vulva; dorsal oracera-
tubae arranged in conspicuous rows; genacerores 6-8(11-17)17-
20. — ^Australia on Callistemon Candida Green.
CO. Pygidium with each median lobe distinctly narrower than sec-
ond lobe; median pair of lobes strongly divergent, twice as
long as wide, mesal and lateral margins subparallel, distal
end bluntly rounded, entire, proximal ends distant; second pair
of lobes large, truncately rounded, as long as median pair;
plates longer than lobes, arranged 0, 2, 5-6, those of each lateris
arranged 3-lor2-l, described plates of each second incisura
probably a latadentis; genacerores 7-8(14-19)11-14. — Chionaspis
berlesii Leon. (Lindgr.) — Canary Islands on Plocama, Micro-
meria, Ruta, Salsola, Cneorum, Cytisus canariensis Lindgr.
bb. Pygidium with median pair of lobes quadrangular, not divergent.
c. Pygidium with median and second pair of lobes distinct.
d. Pygidium with median pair of lobes crenulate at distal end.
e. Pygidium with genacerores arranged 6-10(8-12)10-14;
median pair of lobes subquadrate, short, conspicuous be-
cause of their dark color, distal end crenulate; second
pair of lobes indicated as short rounded structures, trian-
gular; plates long and spine-like, arranged 0, 1, 2, those
of each lateris distant; anus distant from caudal margin;
submarginal region with transverse elongate groups of
oraceratubae ; second nymphal female with median pair
of lobes large, quadrate, distant; lateres strongly serrate;
plates long and spine-like. — Natal on Capparis
retigera Ckll.
ee. Pygidium with genacerores arranged 1-7(8-13)23-30;
median pair of lobes quadrangular, mesal and lateral
margins straight, caudal end irregularly and coarsely ser-
TRIBE DIASPIDINI 333
rate; second pair of lobes rudimentary, sometimes obso-
lete, broader than long, broadly rounded, entire; plates
long, slender, longer than median lobes, arranged 0, 1, 3,
those of each lateris arranged l-(l-l) ; dorsal oraceratubae
in large, well marked groups or bands; anus between pre-
genacerores; mesospiracerores 4-5; lateres with irregular
projections. — Southwest Africa on Rhus
mytilaspiformis Newst.
dd. Pygidium with median pair of lobes quadrangular, distal
margin with mesal and lateral notches, rounded between
notches; second pair of lobes longer than broad, bluntly
rounded, one-half width of median lobe; altaceratubae in
nine pairs, arranged 1-2-2-2-2, their oraceratubae very large;
plates arranged 0, 1, 7, those of each lateris arranged 1-2-2-2,
placed between altaceratubae; brevaceratubae numerous,
prominent; genacerores 2-4(12-16)15-23; anus located about
eleven times its width from caudal margin. — Australia on
Eucalyptus. frenchi Green.
cc. Pygidium with median and second pair of lobes small and in-
conspicuous, delicate; median pair of lobes with sides sub-
parallel, distal end rounded, entire; second pair of lobes simi-
lar in form to median; plates longer than lobes, sharply point-
ed, proximal portion broad, arranged 0, 1, 2, one on each lat-
eris adjacent second lobe, other short distance cephalad; lat-
eres irregular, dentate; brevaceratubae apparently few; gena-
cerores 8-14(13-18)6-14. — South Africa on aloe exalMda Ckll.
aa. Pygidium with second pair of lobes incised.
b. Pygidium with median pair of lobes with proximal portion of
their mesal margins contiguous, sometimes apparently fused.
c. Pygidium with median pair of lobes smaller than mesal lobe-
let of second pair, broad, not prominent; second pair of lobes
entire, incised, mesal lobelet large, broadly rounded, broader
than long, projecting, three times as large as lateral lobelet;
plates on caudal portion about one-half size of those on cephalic
portion, arranged 0, 2, 9, those of each lateris arranged 2-2- (1-1) -
1-2; tooth-like latadentis near middle of each lateris; anus
distant from caudal margin; genacerores 10-10(14-14)10-15. —
Algeria on Deverra biloMs Newst.
cc. Pygidium with median pair of lobes distinctly larger than
mesal lobelet of second pair, subtriangular in outline, bluntly
pointed, crenulate, fused at proximal ends, mesal margins di-
vergent, lateral margins oblique; second pair of lobes incised,
mesal lobelet much larger than lateral, lobelets triangular, en-
tire; plates or furcapectinae, arranged 0, 0, 1; genacerores
5-7(8-14)19-25. — Transvaal, Africa on Acacia caffra Brain.
bb. Pygidium with median pair of lobes not with proximal portion of
mesal margins fused for short distance.
c. Pygidium with two plates in each second incisura.
334 THE COCCIDAE
d. Pygidium with plates short and slender, arranged 0, 2, 7^
those of each lateris arranged 2-2-1-2; median pair of lobes
entire, long, narrow, bluntly pointed, spatulate; second pair
of lobes incised, mesal lobelet much larger than lateral,
slightly smaller than median lobe, similar in shape; gena-
cerores 13-13(18-24)14-17; mesospiracerores and metaspira-
cerores present; anus and vulva superimposed. — Ceylon on
Andropogon. divergens Green.
dd. Pygidium with plates slender, arranged 0, 2, 10, those of each
lateris arranged 2-2-l-5or6; median pair of lobes elongate,
subparallel, distal ends rounded or bluntly pointed, entire;
second pair of lobes incised, entire, mesal lobelet much larger
than lateral, nearly as wide as median lobe, rounded; gena-
cerores 14-16(30-43)20-27; mesospiracerores and metaspira-
cerores present; anus superimposed over vulva. — Ceylon on
Andropogon. graminis Green.
cc. Pygidium with one plate in each second incisura.
d. Pygidium with distance between median pair of lobes never
more than width of median lobe.
e. Pygidium with fifteen to sixteen plates on each lateris,
plates arranged 0, 1, 15-16, those of each second incisura
small, those of each lateris arranged 3-2or3-10; median
pair of lobes large, rounded, broader than long, distinct
lateral notch, narrowly separated; second pair of lobes
incised, lobelets subequal, triangular, together much small-
er than median lobes; thorax, preabdomen, and pygidium
with numerous brevaceratubae ; mesospiracerores repre-
sented by large compact groups; genacerores 2-8(23-35)
42-65; anus located near mesogenacerores, caudad of vulva.
— British East Africa on wild fig tree fici Newst.
ee. Pygidium with ten or eleven plates on each lateris.
f. Pygidium with two or three plates on each lateris.
g. Pygidium with postgenacerores 36-46; median pair
of lobes as broad as long, broadly rounded, entire;
second pair of lobes incised, mesal lobelets nearly as
long as median lobe, longer than broad, bluntly
pointed, entire, lateral lobelet similar in form but
smaller; plates long, much longer than lobes, ar-
ranged 0, 1, 3, those of each lateris arranged 1-2;
lateres irregular; genacerores 4-9(16-23)36-46. —
South Africa on Chaetachme chaetachmae Brain.
gg. Pygidium with postgenacerores 8-22.
h. Pygidium with three plates on each lateris, ar-
ranged 1-1-1, plates long, stout, two or three times
as long as lobes; median pair of lobes subtriangu-
lar, pointed, lateral and mesal margins convex
at middle, serrate; second pair of lobes incised,
mesal lobelet subquadrangular, distinct mesal and
lateral notches, nearly as large as median lobe,
TRIBE DIASPIDINI 335
lateral lobelet small, much smaller than mesal,
bluntly pointed, lateral notch; lateres irregular;
genacerores 3-4(4-9)8-13. — South Africa on Cap-
paris. capparisi Brain.
hh. Pygidium with two plates on each lateris.
i. Pygidium with mesal lobelet of second pair of
lobes distinct, never inconspicuous; median pair
of lobes triangular, pointed, one or two mesal
notches, lateral margin oblique; second pair of
lobes incised, mesal lobelet entire, distal end
bluntly rounded, nearly twice as large as lat-
eral; plates small, slender, longer than lobes,
arranged 0, 1, 2, those of each lateris distant;
lateres irregular; genacerores 2-4(14-20)19-22. —
South Africa on Euphorbia.. _et*p7ior&iae Brain.
ii. Pygidium with mesal lobelet of second pair of
lobes inconspicuous, narrowed toward proximal
end, spatulate, lateral lobelet smaller than
mesal, tapering; median pair of lobes conspicu-
ous, short, wide, divergent, truncate; plates ar-
ranged 0, 1, 2, those of each lateris arranged
one near second lobe and the other short dis-
tance cephalad; genacerores 4-8(13-15)19-21. —
West Australia on Agonis agonis Fuller.
ff. Pygidium with four or more plates on each lateris.
g. Pygidium with second pair of lobes small and in-
conspicuous, mesal lobelet never subequal in size to a
median lobe.
h. Pygidium with median pair of lobes large and
conspicuous.
i. Pygidium with second pair of lobes incised,
lobelets subequal, similar, triangular, or slight-
ly truncate; median pair of lobes conspicuous,
apparently diverging, wide and short, distant,
entire; latadentes present cephalad of second
pair of lobes, three groups of two each on each
lateris; mesal margin of median pair of lobes
and median incisura thickened, horse-shoe-
shaped; plates long, slender, much longer than
lobes, arranged 0, 1, 3, those of each lateris ar-
ranged 1-1-1 and placed adjacent to and cephalad
of each group of latadentes, genacerores 5-5(11-
15)15-21. — West Australia on Eucalyptus
ethelae Fuller,
ii. Pygidium with second pair of lobes incised,
mesal lobelet three or four times size of lateral,
subtriangular, distal end bluntly rounded;
median pair of lobes large, projecting, mesal
margins oblique, undulate; plates large, dagger-
336 THE COCCIDAE
shaped, arranged 0, 1, 6; genacerores in five
almost contiguous groups, arranged 4-7(11-18)
26-32. — South Africa on Leucadendron
leucadendri Brain.
hh. Pygidium with median pair of lobes short, conic-
al, separated by slight wedge-shaped depression;
second pair of lobes incised, mesal lobelet longer
and larger than lateral, lateral short, wide, round-
ed at caudal end; each lateris with its margin
prolonged and thickened in three places, probably
altaceratubae, plates tapering, very long, situated
caudad of lobes and prolongations of margin, ar-
ranged 0, 1, 4, those of each lateris arranged
1-1-1-1; genacerores 4-4(19-20)25-30.— South Aus-
tralia on Xanthorrhoea xanthorrhoeae Fuller.
gg. Pygidium with second pair of lobes incised, mesal
lobelet larger than lateral, often approximating a
median lobe in size; median pair of lobes well de-
veloped, distant, separted by distance about equal
to width of median lobe; plates prominent, arranged
0, 1, 4-5, those of each lateris arranged l-lor2-2;
latadentes located in median and second incisurae
and associated with plates of lateres; lateres cleft at
intervals, margin denticulate; genacerores 5-7(9-14)
7-14. — District of Columbia on East Indian bamboo.—
howardi Cooley.
dd. Pygidium with distance between median pair of lobes more
than twice width of a median lobe.
e. Pygidium with median pair of lobes subquadrangular,
mesal and lateral margins parallel, caudal margin trans-
verse, with three teeth; second pair of lobes distinct, in-
cised, lobelets long, slender, caudal ends rounded, mesal
larger than lateral; plates only slightly longer than lobes,
arranged 0, 1, 2, those of lateris placed one near second
lobe and other distant and very small, other plates may
be present; dorsal oraceratubae numerous on all parts of
body, arranged in regular series on pygidium; genacerores
10-10(26-32)30-42.— Niger, Africa on Xymenia
nigerensis Vayss.
ee. Pygidium with median pair of lobes triangular, pointed,
lateral and mesal margins oblique and concave, shouldered
on proximal portion, about as broad as long; second pair
of lobes incised, mesal lobelet similar in form to median,
as large or larger than median, lateral lobelet triangular,
small, one-third size of mesal lobelet; plates twice as long
as lobes, proximal portion stout, arranged 0, 1, 2, those
of each lateris distant; lateres irregular; genacerores 7-8
(18-23)34-40. — South Africa on aloe margaritae Brain.
TRIBE DIASPIDINI 337
SPECIES OF UNACHIONASPIS
a. Pygidium with median pair of lobes large and distinct.
b. Pygidium always with less than ten plates on each lateris.
c. Pygidium with genacerores 2-3(2-3)2-3; median pair of lobes
large, projecting, constricted at proximal end, separate; plates
arranged 0, 5, those of each lateris arranged 1-1-1-1-1; anus
located near middle of length of pygidium; scale of adult fe-
male pyriform. — South Australia on Eucalyptus
assimilis Mask,
cc. Pygidium with genacerores 7-11(10-14)6-9; median pair of
lobes large, proximal half of mesal margins parallel, subadja-
cent, distal portion diverging, distant, lateral margins oblique,
long, crenulate; plates, except one adjacent median lobe, long
and dagger-like, arranged 0, 5, those of each lateris arranged
1-2-2; lateres with latadentes; setae slender, about as long as
median lobes; spiracerores wanting. — South Africa on lilac.—
amhigua Brain,
bb. Pygidium with eleven to fourteen plates on each lateris, arranged
3-3-5-8, wanting in median incisura; median pair of lobes large,
elongate, crenate on both margins, adjacent at proximal end, di-
vergent caudad; each lateris with two shallow notches, one be-
tween each two groups of plates; segments of preabdomen and
thorax, beginning at caudal end, bear plates on each side, as fol-
lows: 9-5-8-4 or more — 4 or more, only rudiments; genacerores in
five groups. — Antigua on Heliotrope major Ckll.
aa. Pygidium with median pair of lobes small and inconspicuous.
b. Pygidium with six or more large plates; median pair of lobes
pointed, triangular, almost invisible, other lobes represented by
angular projections, distant, twice width of one of them apart;
plates several times length of lobes, large, prominent, arranged
0, 7, those of each lateris arranged 1-1-2- (1-1-1), cephalic plate
forked; dorsal oraceratubae numerous near caudal margin; anus
located between pregenacerores ; genacerores 7-10(12-16)9-15. —
Japan on bamboo colemani Knw.
bb. Pygidium with two short, stout plates on each lateris, distant
from each other and from cephalic plate, smaller than caudal;
median pair of lobes small, identified as plates by Brain, short,
notched at distal end, broad as long; lateres irregular, with
indentations; mesospiracerores 4-5; genacerores 3-5(4-6)4-7. — South
Africa on Euphorbia globosus Brain.
SPECIES OF FUNDASPIS
a. Pygidium with distinct mesal indentation, marking division between
two median lobes.
b. Pygidium with median pair of lobes with distinct lateral notch,
otherwise entire, distal end bluntly rounded, each lobe longer than
broad; second pair of lobes incised, mesal lobelet with its lateral
margin oblique with two notches, more than twice as long as
lateral lobelet, lateral lobelet small and inconspicuous, entire or
338 THE COCCIDAE
with single notch; third pair of lobes incised, small, inconspicu-
ous; plates and pectinae long, slender, those of each side of each
third lobe often furcapectinae, arranged 0, 1, 1-2, 9-13, those of
each lateris arranged 2-2to4-5to7; dorsal oraceratubae few in
number, arranged in rows; anus located about its own width
cephalad of vulva; genacerores 20-30(18-42)20-30. — Massachusetts,
Connecticut, New York, Ohio, Illinois, Minnesota, Missouri, Kan-
sas, Oklahoma, Texas on Ulmus americana Johns.
bb. Pygidium with median pair of lobes not with distinct lateral
notch.
c. Pygidium with median pair of lobes entire, not notched, large,
each broader than long, broadly rounded; second pair of lobes
incised, mesal lobelet oblique, much longer than broad and
larger than lateral lobelet, their lateral margins serrate; third
pair of lobes not well marked, incised, mesal lobelet larger
than lateral, both oblique and longer than broad, lateral mar-
gins serrate; plates arranged 0, 1, 1, 7-10, those of each lateris
arranged l-lto2-5to7; dorsal oraceratubae present, arranged
in rows; genacerores 12-19(21-29)15-22. — District of Columbia on
hickory. caryae Cooley.
cc. Pygidium with median pair of lobes serrate, fused on proxi-
mal two-thirds, diverging at distal ends, forming deep notch,
lateral margins oblique; second pair of lobes deeply notched,
mesal lobelet triangular, larger than lateral, lateral margin
oblique, serrate, lateral lobelet triangular, entire; third pair
of lobes broader than long, oblique, serrate; plates not long,
those of second incisurae not as long as median lobe, arranged
0, 1, 1, 1, 5-8, those of each lateris arranged l-lto2-4to6;
dorsal oraceratubae arranged in rows; anus located slightly
cephalad of vulva; genacerores 7-10(15-26)14-18. — Ohio on
Nyssa. sylvatica Sand.
aa. Pygidium with median pair of lobes completely fused, mesal inden-
tation wanting, entire, lobes broader than long, truncate with round-
ed angles; second and third pairs of lobes not incised, each caudad
of reniform thickening, not prominent, entire, bluntly pointed;
plates subequal in length to median pair of lobes, arranged 0, 1, 1,
13-14, those of each lateris arranged 2to3-5-6; anus located thirteen
times its width from caudal margin and twice its width cephalad
of vulva; genacerores 10-10(17-20)10-18. — California, New Mexico,
Mauritius on Quercus, Pasania quercus Comst.
SPECIES OF AFRICASPIS
a. Pygidium without plates adjacent to each median lobe; median
pair of lobes fused, three deep indentations forming four subequal
rounded lobules, mesal indentation marking division between lobes;
other lobes wanting; plates about as long as median pair of lobes,
arranged 0, 3-4, those of each lateris arranged 0-l-2to3-0-0 ; three
adjacent tooth-like projections between each median lobe and caudal
plate, single projection between caudal plate and next group; dorsal
oraceratubae numerous, especially on thorax; anus located near
TRIBE DIASPIDINI 339
middle of groups of genacerores; mesospiracerores 3-4; altacera-
tubae present; genacerores 6-8(23-28)28-36. — Uganda, Africa _.__
chionaspiformis Newst.
aa. Pygidium with two plates adjacent to median pair of lobes; median
pair of lobes fused for half their length, deep lateral notch and
mesal indentation, forming two prominent rounded lobules; other
lobes wanting; plates longer than lobes, arranged 0, 7-8, those of
each lateris arranged 2-l-2-l-lto2 ; each lateris with minute tooth-
like projections between groups of plates; group of 5-7 setae near
spiracles; dorsal oraceratubae in rows; altaceratubae few in number;
genacerores 6-11(17-26)20-32. — Uganda, Africa on Cassia
cassiae Newst.
SPECIES OF GREENASPIS
Body of adult female elongate, six or more times as long as wide, head
and thorax constituting over one-fourth of length of body, sides of
caudal segments of preabdomen strongly produced and bearing two
or three plates; pygidium with median pair of lobes prominent, di-
vergent, mesal margins emarginate; second pair of lobes incised,
mesal lobelet truncate, much larger than lateral; plates stout, 0, 1, 5-6,
those of each lateris arranged l-l-l-2to3; genacerores 4-6(8-10)10-10;
mesospiracerores 1-2; anus located slightly cephalad of vulva. — Ceylon
on Arundinaria. elongata Green.
SPECIES OF TENUIASPIS
Body of adult female small, averaging less than one millimeter; seg-
ments of preabdomen without plates; pygidium with median pair of
lobes projecting, minute, narrow, bluntly rounded; second pair of
lobes incised, minute, inconspicuous; plates comparatively large, twice
as long as the lobes, arranged 0, 1, 7-8, those of each lateris arranged
l-l-l-4to5; altaceratubae four on each side; brevaceratubae located
only near altaceratubae; genacerores 4-6(9-12)8-8; mesospiracerores
represented by small group of pores; anus located twice its width
cephalad of vulva. — Ceylon on Tetranthera minuta Green.
SPECIES OF UNASPIS
Body of adult female long and narrow, about three times as long as
wide, caudal portion widest; pygidium with median pair of lobes pro-
jecting, wedge-shaped, lateral m.argins free, distal ends truncately
rounded, entire; second and third pairs of lobes incised, lobelets
wedge-shaped, mesal lobelet larger than lateral, margins entire; plates
short and stout, arranged 0, 0, 0, 4, those of each lateris arranged
1-1-2; altaceratubae six on each side; brevaceratubae nuraer(rus, prom-
iscuously arranged; genacerores 4-4(6-7)4-5; mesospiracerores repre-
sented by small group; anus and vulva superimposed. — Ceylon
acuminata Green.
SPECIE'S OF LINEASPIS
Pygidium with median pair of lobes plate-like or spine-like, short, dis-
tance between them four times their length ; second pair of lobes normal
in form, broader than long, broadly rounded; other lobes wanting;
340 THE COCCIDAE
plates short, spine-like, subequal in length to median pair of lobes,
arranged 0, 2, 3-4, those of each lateris arranged, two near a second
lobe, and group of one or two near middle; lobes and plates all some-
times wanting; genacerores 3-4(9-13)8-17. — Algeria on Cypress
striata Newst.
SPECIES OF HEMICHIONASPIS
a. Pygidium of adult female with three pairs of lobes.
b. Pygidium with median pair of lobes small, scarcely projecting
beyond distal end of second, each with single notch; second pair of
lobes incised; third pair of lobes not incised; plates 0, 1, 1, 6, those
of incisurae short, those of each lateris arranged 1-1-4; altacera-
tubae seven on each side; dorsal oraceratubae few; anus located
cephalad of vulva; genacerores 1-12(20-23)20-23. — India on Thea.
separata Green,
bb. Pygidium with median pair of lobes large, extending slightly
caudad of second pair; second pair of lobes incised, feebly chiti-
nized, distinct, distal portion of lobelets expanded; third pair
of lobes sometimes developed, incised, mesal lobelet longer than
broad, lateral lobelet broader than long, margins of lobelets
serrate; latadentes of second incisurae very prominent, longer
than broad; plates arranged 0, 1, 1, 5-6, those of each lateris
arranged l-lto2-3; brevaceratubae few in number, large and oval;
genacerores 8-10(20-23)17-23. — Ceylon on Carsia alatae Ruth.
aa. Pygidium with less than three pairs of lobes.
b. Pygidium with two pairs of lobes.
c. Pygidium without plates in each second incisura; distance
between median lobe and second lobe greater than width of a
median lobe; median pair of lobes large and prominent, mi-
nutely serrate or crenulate, hardly projecting; second pair
of lobes small, incised, narrow, inconspicuous, scarcely pro-
jecting; plates moderately long, arranged 0, 0, 1, each lateris
probably with more; dorsal oraceratubae few, large and con-
spicuous; mesospiracerores represented by small group; gena-
cerores 1-10(13-23)17-25. — Bengal, India on Ficus fici Green.
cc. Pygidium with plates in second incisurae, if wanting, median
and second pair of lobes not the width of a median lobe apart,
d. Pygidium with two median lobes together not as broad as
long or at most subequal.
e. Plygidium with second pair of lobes incised, mesal lobe-
let projecting beyond smaller lateral lobelet; median pair
of lobes long, projecting distinctly beyond distal ends
of second pair, lateral margins with two notches or irreg-
ularly crenulate; plates arranged 0, 1, 6, those of each
lateris arranged 1-1-1-3; prominent latadentis in each sec-
ond incisura between plate and third lobe; dorsal oracera-
tubae few in number; genacerores 6-7(11-12)9-11. — Socotia
Island on Dracaena. dracaenae Cooley.
ee. Pygidium with second pair of lobes not incised, longer
than broad, hatchet-shaped, distal margin oblique; median
TRIBE DIASPIDINI 341
pair of lobes with mesal margins straight or slightly di-
vergent, not adjacent, hatchet-shaped, longer than second"
lobe, lateral margin with two notches; plates small,
arranged 0, 1, 3, those of each lateris arranged 1-1-1;
margin with six altaceratubae on each side; brevacera-
tubae few in number, not in crescentic transverse rows;
genacerores 2-4(8-13)8-15. — This species is given this place
only because of its original reference. — North Australia on
Pandanus. pseudaspidistrae Green.
dd. Pygidium with two median lobes together broader than long,
e. Pygidium with anus and vulva superimposed; median pair
of lobes moderately produced, lateral margins oblique, two
notches, forming three subequal crenulations; plates longer
than lobes, arranged 0, 7, those of each lateris arranged
1-2-2-2; genacerores 12-18(15-19)13-17.— Natal, Africa on
Altermanthera. cyanogena Ckll.
ee. Pygidium with anus and vulva not superimposed,
f. Pygidium with anus located cephalad of vulva.
g. Pygidium with median pair of lobes broader than
long, margin strongly crenulate, with three depres-
sions; second pair of lobes incised, mesal lobelet
enlarged and subtruncate at distal end; third pair
of lobes wanting; plates large; vulva located between
postgenacerores and caudad of anus; genacerores 8-8
(23-23)12-15.— French Guinea, Africa on Eleis
marchali Ckll.
gg- Pygidium with median pair of lobes projecting dis-
tinctly beyond distal end of second pair,
h. Pygidium with three rows of brevaceratubae,
about four oraceratubae in each half of each row;
median pair of lobes large, crenulate, broader
than long; second pair of lobes minute, incised;
plates long, arranged 0, 2, 7, those of each lateris
arranged 2-1-1-3; altaceratubae five on each side;
genacerores 11-20(20-27)16-22; oblong chitinous
thickening laterad of genacerores; mesospira-
cerores represented by conspicuous groups; in-
sects living in small pits at base of veins on under-
side of leaves. — Ceylon on Elaeocarpus
scroMcularum Green,
hh. Pygidium never with three rows of brevacera-
tubae, rarely with two partial rows,
i. Pygidium with genacerores 6-8(11-15)10-15;
median pair of lobes prominent, crenulate; sec-
ond pair of lobes small, incised; plates stout,
longer than lobes, arranged 0, 1, 6, those of
each lateris arranged 1-1-1-3; altaceratubae six
on each side; brevaceratubae one on each side
of fourth and three on each side of fifth row:
342 THE COCCIDAE
mesospiracerores 3-4, metaspiracerores wanting;
anus and vulva rarely superimposed. — Ceylon
on Rhododendron rhododendri Green.
ii. Pygidium with genacerores 4-4(8-10)8-10; me-
dian pair of lobes large, dark colored, semicir-
cular, margins dentate or crenulate with six
small teeth, outer two very minute; second pair
of lobes incised, lobelets longer than broad,
distal end of mesal lobelet rounded and of lat-
eral pointed; large plate cephalad of each sec-
ond lobe and prominent latadentis bearing ora-
ceratuba on each side of plate; two obliquely
truncated lobe-like projections with rounded
angles adjacent to and cephalad of cephalic
latadentis; plate adjacent to cephalic lobe-like
projection and another at cephalic end of each
lateris; dorsal brevaceratubae not described. —
Philippine Islands on Uvaria
uvariae Ckll. & Robs,
ff. Pygidium with anus located caudad of vulva.
g. Pygidium with genacerores not 16-16(19-20)25-25.
h. Pygidium without plates in each second incisura;
scale of adult female long and narrow with sides
subparallel, four to six times as long as wide;
median pair of lobes indistinctly separated, broad-
er than long, crenulate; second pair of lobes
small, incised; plates short on caudal portion and
long on cephalic, arranged 0, 0, 5, those of each
lateris arranged 1-1-1-2; altaceratubae six on each
side; brevaceratubae one on each fourth and
three on each fifth row; genacerores 8-8(18-25)
16-18; pregenacerores generally with largest num-
ber; mesospiracerores distinct and one metaspira-
ceroris. — India, Ceylon on Thea, Psychotria,
Eurya. theae Mask.
hh. Pygidium with one plate in each second incisura.
i. Pygidium with 12-28 postgenacerores.
j. Pygidium with cephalic altaceratubae of each
side always distant from lateral brevacera-
tubae of fifth row; median pair of lobes in-
distinctly separated, broader than long,
crenulate; second pair of lobes incised, mesal
lobelet prominent, truncate; third pair of
lobes obsolete; plates short on caudal and
long on cephalic portion, arranged 0, 1, 5-6,
those of each lateris arranged l-l-l-2or3;
altaceratubae six on each side, brevacera-
tubae one to two on each side of fourth and
three to four on each side of fifth row; gena-
cerores 4-16(11-22)12-28, postgenacerores al-
TRIBE DIASPIDINI 343
ways with largest number; mesospiracerqres^
forming distinct group, sometimes with ln~
conspicuous group of metaspiracerores. — ^Asia,
Australia, France, England, eastern Canada
and United States, California, West Indies,
Brazil on Aspidistra, Acacia, Davallia, Cocos,
Cyanotus, Areaca, orchids, ferns, orange, fig,
mango, pepper-tree, etc aspidistrae Sign.
jj. Pygidium with cephalic altaceratubae of each
side adjacent to lateral brevaceratubae of
fifth row.
k. Pygidium with median pair of lobes promi-
nent, longer than broad, crenulate; second
pair of lobes minute, prominent; third pair
of lobes obsolete; plates comparatively
short, arranged 0, 1, 5-6, those of each lat-
eris arranged l-l-l-2or3; altaceratubae six
in number; brevaceratubae one on each
fourth and one on each fifth row near an
altaceratuba and double row of five or six
cephalad of fifth row; genacerores 6-11(14-
17)12-17; mesospiracerores represented by
small group. — Chionaspis albizziae Green. —
New Zealand, Japan, Ceylon, Florida, West
Indies, Panama, Brazil on Parsonia, Rhipo-
gonium, Hibiscus, Capsicum, Melia, Azed-
arch, Cycas, Pelargonium, Albizzia, Agave,
fig, cocoa-palm, cotton, pepper, orange, etc.
minor Mask,
kk. Pygidium with median pair of lobes broad-
er than long; scale with exuviae darker
than in type. — West Africa
minor strachani Cooley.
ii. Pygidium with genacerores 7-12(10-13)10-12;
median pair of lobes scarcely projecting, two
lobes together broader than long, strongly den-
ticulate; second pair of lobes incised, lobelets
narrow, equal in width, mesal longest; plates
longer than lobes, arranged 0, 1, 1-1, others
probably present on each lateris; anus approxi-
mately central in location; dorsal oraceratubae
few; scale whitish, almost colorless, more or
less translucent. — Bengal, India on Ficus
minima Green,
gg. Pygidium with genacerores 16-16(19-20)25-25; anus
about nine times its width from caudal margin;
median pair of lobes together broader than long, with
four crenulations, first two very deep and strong;
second pair of lobes quite rudimentary, not or hardly
extending beyond general margin; plates arranged
344 THE COCCID.VE
0, 1, 3, those of each second incisura small, those of
each'lateris very large and long, placed at long inter-
vals; caudal segment of preabdomen with three
plates on each side; dorsal oraceratubae not numer-
ous; scale pyriform, broad, varying to nearly circu-
lar, light grayish or yellowish. — This species may be
wrongly associated, placed here because compared
with theae and aspidistrae. — Philippine Islands on
Gossypium. townsendi Ckll.
bb. Pygidium with one pair of lobes.
c. Pygidium with plate adjacent to each median lobe,
d. Pygidium with genacerores 19-24(30-38)27-35; median pair
of lobes large, prominent, broader than long, crenulate; other
pairs of lobes obsolete; plates long and slender on cephalic
part of each lateris, short on caudal part, arranged 0, 10-12,
those of each lateris arranged l-l-l-2-5or7 ; caudal preabdomi-
nal segment with 5-7 plates on each side; altaceratubae five
on each side; brevaceratubae two on each fourth and five on
each fifth row; mesospiracerores and metaspiracerores pres-
ent.— Ceylon on Mussaenda, Loranthus, Debregeasia
mussaendae Green,
dd. Pygidium with genacerores 3-3(13-13)11-13; median pair of
lobes together broader than long, not fused, each with about
three notches or crenulations ; plates all long, much longer
than median lobes, arranged 0, 4, those of each lateris ar-
ranged, one adjacent to a median lobe, one in what would
be a third incisura, one near middle of each lateris, and
one at its cephalic end; caudal preabdominal segment with
two plates on each side; spiracerores wanting; prominent
latadentes bearing oraceratubae located on each side between
two caudal plates; anus located slightly cephalad of vulva.
Brazil. orlando Leon.
SPECIES OF PHENACASPIS
a. Pygidium with three or more pairs of lobes.
b. Pygidium with second pair of lobes not incised, short, broadly
rounded, entire; median pair of lobes prominent, oblique, diverg-
ing, not confluent, crenulate; third pair of lobes small, slight
extension of margin, rounded, entire; plates large, dagger-shaped,
arranged 0, 1, 2, 2, those of each lateris distant, altaceratubae
large, five on each side; spiracerores wanting; genacerores 12-12
(8-9)14-16. — Turkey on Arthrocnemum arthrocnemi Lindgr.
bb. Pygidium with second pair of lobes always incised.
c. Pygidium always with plates in each second incisura.
d. Pygidium always with plates of lateres arranged in three
groups.
e. Pygidium with one plate in median group of each lateris.
f. Pygidium with (25-34) pregenacerores present.
TRIBE DIASPIDINI 345
g. Pygidium with cephalic group of each lateris with
two plates, plates slender, arranged 0, 1, 1, 4, those
of each lateris arranged 1-1-2; median pair of lobes
moderately large, divergent, not confluent, entire;
second and third pairs of lobes incised, mesal lobe-
let truncate and larger than lateral, lateral lobelets
pointed, all entire; mesospiracerores present; breva-
ceratubae few in number; anus located less than its
own width caudad of vulva, nine times its width
from caudal margin; genacerores 16-16(25-33)16-23. —
Ceylon on Litzea. litzeae Green.
SS- Pygidium with cephalic group of each lateris with
three plates, plates stout, dagger-shaped, arranged 0, 1,
1, 5, those of each lateris arranged 1-1-3; median pair
of lobes large, divergent, confluent, distal ends scarcely
projecting, minutely serrate; second and third pairs
of lobes incised, mesal lobelets larger than lateral, all
lobelets bluntly rounded, entire; mesospiracerores and
metaspiracerores present; brevaceratubae well devel-
oped; anus located seven times its width from caudal
margin and about its own width cephalad of vulva;
genacerores 12-16(28-34)18-24.— Ceylon on Psidium,
Actinodaphne. megaloba Green.
ff. Pygidium with (10-23) pregenacei-ores present.
g. Pygidium with third pair of lobes small or wanting,
incised, broad and short; median pair of lobes con-
fluent at proximal end, serrate, large; second pair
of lobes incised, mesal lobelet larger than lateral,
extends caudad about as far as median pair; plates
arranged 0, 1, 1, 5, those of each lateris arranged
1-1-3; altaceratubae three on each side, opening in
latadentes; genacerores 7-9(17-23)23-34. — China, Cal-
ifornia at quarantine on palm cockerelU Cooley.
gg. Pygidium with third pair of lobes large and well
developed.
h. Pygidium with second pair of lobes incised, mesal
lobelet distinctly larger than lateral, lobelets
well developed, mesal lobelet broadly rounded
and lateral bluntly pointed, entire or obscurely
serrate; third pair of lobes large and well devel-
oped, incised, mesal lobelet distinct, lateral lobe-
let obscure or wanting; median pair of lobes
large, well developed, divergent, confluent, scarce-
ly projecting, serrate; plates dagger-shaped, those
of second incisurae very short, arranged 0, 1, 1,
4-6, those of each lateris arranged l-l-2or4; ora-
ceratubae of brevaceratubae with 2-3 in mesal
group of second row, third row 4-6 in mesal group
and 2-4 in lateral, fourth row 4-8 mesal and 4-6
lateral; genacerores 6-13(10-17)10-16. — Kansas,
Japan on Rhus, sycamore platani Cooley.
346 THE COCCIDAE
hh. Pygidium with second pair of lobes incised, mesa!
and lateral lobelets subequal in size, constricted
at proximal end, distal end broadly rounded, en-
tire; third pair of lobes incised, mesal lobelet
distinctly larger than lateral, constricted at prox-
imal end, distal end broadly rounded, entire;
median pair of lobes long, strongly divergent, not
confluent, serrate; plates longer than lobes, spine-
like, arranged 0, 1, 1, 3, those of each lateris ar-
ranged 1-1-1; altaceratubae apparently seven on
each side; latadentes with oraceratubae in sec-
ond and third incisurae; lateres irregularly den-
tate with four marginal oraceratubae; genacerores
8-9(16-17)16-17, pregenacerores and postgenacerores
of each side almost confluent. — Philippine Islands
on Mischocarpus. mischocarpi Ckll. & Robs.
ee. Pygidium always with more than one plate in median
group of each lateris,
f. Pygidium with three plates in median group of each
lateris.
g. Pygidium with four plates in cephalic group of each
lateris, plates long, slender, arranged 0, 1, 1, 8, those
of each lateris arranged 1-3-4; median pair of lobes
large, divergent, confluent, serrate; second and third
pairs of lobes incised, lobelets prominent, mesal
larger than lateral, constricted at proximal end, dis-
tal end bluntly rounded, entire; brevaceratuba« nu-
merous; mesospiracerores present; anus ten times its
width from caudal margin and about its own width
cephalad of vulva; genacerores 10-16(23-27)27-60. —
Ceylon on Gelonium varicosa Green.
gg. Pygidium with five plates in cephalic group of each
lateris, plates arranged 0, 1, 1-2, 9-11, those of each
lateris arranged lto3-3-5; median pair of lobes mod-
erate in size, divergent, confluent, serrate; second
pair of lobes incised, mesal lobelet larger than lat-
eral, projects farther caudad, sometimes exceeding
median lobe; third pair of lobes not incised, some-
times rudimentary; altaceratubae four on each side,
opening in latadentes; genacerores 8-14(19-28)19-33.
— Japan on Aucuba. aucu'bae Cooley.
ft. Pygidium with two plates in median group of each lat-
eris, plates long, dagger-shaped, arranged 0, 1, 1, 7, those
of each lateris arranged 1-2-4; median pair of lobes
large, confluent, divergent, crenulate; second and third
pairs of lobes incised, lobelets subequal, rounded, en-
tire; brevaceratubae numerous; anus about seven times
its width from caudal margin and about twice cephalad
of vulva; genacerores 4-6(15-15)30-30. — Samoa on palm.
samoana Doane & Ferris.
TRIBE DIASPIDINI 347
dd. Pygidium with plates of each lateris arranged in less than
three groups.
e. Pygidium with plates of each lateris arranged in two
groups.
f. Pygidium with plates of incisurae and lateres at least
twice as long as lobes.
g. Pygidium with genacerores 4-8(12-18)14-16; median
pair of lobes rather small, divergent, distal ends
broadly rounded, hardly projecting, confluent; sec-
ond and third pairs of lobes greatly reduced in size,
incised, lobelets distinctly separated, broadly rounded,
entire, mesal lobelet larger than lateral; plates long,
stout, three or more times as long as lobes, arranged
0, 1, 1, 2, those of each lateris arranged 1-1; altacera-
tubae four on each side. — Chionaspis pinifoliae
heterophyllae Cooley. — Florida, Rhode Island on
Pinus. heterophyllae Cooley.
gg. Pygidium with genacerores 5-8(20-25)21-21; median
pair of lobes large, prominent, divergent, confluent at
proximal end, distal portions broad, broadly rounded,
serrate; second and third pairs of lobes incised,
mesal lobelet much larger than lateral, truncate, en-
tire; plates long and slender, longer than lobes, ar-
ranged 0, 1, 1, 3, those of each lateris arranged 1-2;
altaceratubae at least four times its width from
caudal margin, probably cephalad of vulva. — Japan on
Quercus. kinshinensis Knw.
ff. Pygidium with plates of incisurae and lateres distinctly
shorter than lobes, lobe-like, arranged 0, 1, 1, 2, those
of each lateris arranged 1-1; cephalic one very broad
with margin serrate; median pair of lobes long, deeply
recessed, divergent, not confluent, coarsely serrate;
second and third pairs of lobes incised, constricted at
proximal end, distal portion dilated, broadly bluntly
rounded, subequal in size, entire; latadentes in second
and third incisurae and between plates of lateres, bear-
ing large oraceratubae ; vulva located between postgena-
cerores; altaceratubae apparently three on each side;
brevaceratubae small and few in number; genacerores
7-8(10-14)8-11. — German East Africa on forest tree under
fungus. lutea Newst.
ee. Pygidium with plates of each lateris arranged in a single
group.
f. Pygidium with genacerores 10-10(22-26)19-26; median
pair of lobes large, widely divergent, broader than long,
confluent, serrate; second and third pairs of lobes in-
cised, distal ends broadly rounded, subequal in size, en-
tire; latadentes in second and third incisurae, bearing
348 THE COCCIDAE
oraceratubae ; plates distinctly longer than lobes, spine-
like, arranged 0, 1, 1, 1. — Natal on mango, palm
j natalensis Ckll.
ff. Pygidium with genacerores 5-5(20-21)38-39; median
pair of lobes divergent, not confluent, hardly projecting;
second and third pairs of lobes prominent, incised,
mesal lobelets larger than lateral, projecting caudad of
median lobes, broadly bluntly rounded, entire; plates
dagger-shaped, twice or more as long as lobes, arranged
0, 1, 1, 1; altaceratubae apparently three on each side. —
Dutch East Africa on Dracaena tang ana Lindgr.
cc. Pygidium always without plates in second incisurae, plates
spine-like, much longer than lobes, arranged 0, 0, 1, 1; median
pair of lobes large, divergent, not confluent, broadly rounded,
serrate; second and third pairs of lobes incised, lobelets broadly
truncately rounded, entire, mesal lobelets larger than lateral,
mesal lobelets of second pair projecting farther caudad than
median lobes; setae of median incisura prominent, nearly as
long as median lobes; anus located just cephalad of mesogena-
cerores, evidently cephalad of vulva; spiracerores apparently
wanting; altaceratubae prominent, five on each side; gena-
cerores 1-2(6-7)3-4. — French Guinea on Macrolobium
l)ussi Newst.
aa. Pygidium always with less than three pairs of lobes,
b. Pygidium with two pairs of lobes.
0. Pygidium with two plates in each second incisura, plates longer
than lobes, arranged 0, 2, 4, those of each lateris arranged
2-1-1; median pair of lobes prominent, narrow, divergent, lat-
eral margin convex, mesal margin concave on distal portion,
distal end rounded, entire; second pair of lobes incised, promi-
nent, mesal lobelet longer than lateral, slightly broader, proxi-
mal portion constricted, distal portion expanded, faintly
notched, lateral lobelet not constricted or expanded or notched;
altaceratubae large, five on each side; brevaceratubae con-
spicuous; anus and vulva superimposed, anus four times its
width from caudal margin; genacerores 7-8(11-14)10-12. — India
on Oryza. decurvata Green.
cc. Pygidium always with less than two plates in each second
incisura.
d. Pygidium with one plate in each second incisura.
e. Pygidium with postgenacerores always more than twenty,
f. Pygidium with pregenacerores always sixty or more,
genacerores arranged 30-30(63-63)60-70; median pair of
lobes large, prominent, oblique, confluent at proximal
end, serrate, distal end broadly rounded, entire; sec-
ond pair of lobes incised, prominent, narrow, mesal
lobelet largest, distal ends bluntly pointed, entire; pro-
thorax with ventral transverse row of conspicuous oval
scars; plates small, hardly longer than median pair of
TRIBE DIASPIDINI 349
lobes, arranged 0, 1, 6, those of each lateris arranged
1-1-4; mesogenacerores present; anus located near pre-
genacerores, apparently cephalad of vulva; dorsal ora-
ceratubae very numerous. — India on Ficus, Solanum,
Thea. manni Green.
. Pygidium with pregenacerores always less than thirty,
g. Pygidium never with more than three plates on each
lateris.
h. Pygidium with genacerores 4-7(11-15)20-26; me-
dian pair of lobes low, broad, divergent, distant at
proximal end, broadly rounded, serrate; second
pair of lobes incised, mesal lobelet twice or more
size of lateral, bluntly rounded, entire, lateral
lobelet triangular, pointed, entire; plates long,
stout, arranged 0, 1, 3, those of each lateris ar-
ranged 1-1-1; mesospiracerores 3-5. — South Africa
on Rhus. lounsburyi Cooley.
hh. Pygidium with genacerores 6-9(11-20)29-46; meso-
spiracerores 5-6; plates slender, much longer than
lobes. — South Africa on Ekebergia
lounsburyi ekehergiae Brain,
gg. Pygidium always with more than three plates on
each lateris.
h. Pygidium with second row of brevaceratubae al-
ways with oraceratubae present,
i. Pygidium with second row of brevaceratubae
with 2-3 oraceratubae present, third row with
3-4 mesal and 4-5 lateral, fourth row with 3-4
mesal and 4-6 lateral; median pair of lobes
large, oblique, confluent at proximal end, dis-
tal end broadly rounded and entire, mesal mar-
gin faintly serrate; second pair of lobes large,
prominent, entire, incised, mesal lobelet larger
than lateral, bluntly pointed; plates stout, par-
ticularly those of lateres, arranged 0, 1, 5-9,
those of each lateris arranged l-l-lor2-2or4 ;
genacerores 8-15 ( 19-31 ) 13-23. — California, Japan
on Wistaria, Alnus, Salix wistariae Cooley.
ii. Pygidium with second row of brevaceratubae
always with more than three oraceratubae.
j. Pygidium with second row of brevaceratubae
with nine oraceratubae present, other rows
with numerous irregularly arranged oracera-
tubae; median pair of lobes distinct, dis-
tant, oblique, divergent, margins undulate
or toothed; second pair of lobes not incised,
mesal lobelet three times size of lateral,
margins broadly rounded, entire; plates not
prominent, arranged 0, 1, 5, those of each
350 THE COCCIDAE
lateris arranged 1-1-1-2; spiracerores well
developed, 4-5 cerores in each group; anus
about ten times its width from caudal mar-
gin and about twice cephalad of vulva; alta-
ceratubae numerous, thirteen or more on
each side, apparently not large; genacerores
11-14(16-20)23-31.— Algeria on Ceratonia
ceratoniae Marchal.
jj- Pygidium with second row of brevaceratubae
with four or five oraceratubae present, other
rows distinct; median pair of lobes contigu-
ous or confluent, divergent; second pair of
lobes distinct, not incised; plates apparently
0, 1, 5, not well developed; spiracerores dis-
tinct, 2-3 cerores in each group; altacera-
tubae not numerous and not arranged in
groups; genacerores 8-10(15-21)15-22. — Al-
geria on Bupleurum hupleuri Marchal.
hh. Pygidium with second row of brevaceratubae with
oraceratubae wanting, third row with 5 mesal
and 4 lateral pores, fourth row with 3-4 mesal
and 5-6 lateral pores; median pair of lobes con-
fluent, oblique, large, divergent, minutely ser-
rate, not projecting strongly; second pair of lobes
minute, incised, inconspicuous; plates stout, larg-
er on cephalic part of lateres, arranged 0, 1, 6, those
of each lateris arranged 1-1-1-3; abdomen and
thorax with many conspicuous oval pores; anus
near to, probably cephalad of vulva, eight times
its width from caudal margin; mesospiracerores
present; genacerores 10-10(18-20)22-22.— Ceylon on
Strobilanthus. stroMlantM Green.
ee. Pygidium with postgenacerores always less than twenty.
f. Species found, so far as known, only in Africa.
g. Pygidium with genacerores 3-4(5-12)13-18; median
pair of lobes broader than long, distant, not oblique,
not confluent, distal margin crenulate; second pair
of lobes incised, triangular, small; plates long, slen-
der, arranged 0, 1, 2, probably three distant plates
on lateres; altaceratubae small; brevaceratubae pres-
ent.— South Africa on mistletoe visci Brain.
gg. Pygidium with genacerores 8-12(14-17)12-15; median
pair of lobes divergent, confluent at proximal end,
not large, entire; second pair of lobes prominent,
incised, mesal lobelet large, projecting farther
caudad than median lobe, bluntly rounded, much
larger than lateral lobelet; plates dagger-shaped, ar-
ranged 0, 1, 3, those of each lateris placed singly and
TRIBE DIASPIDINI 351
distant; altaceratubae not prominent; brevaceratubae
not numerous. — South Africa on aloe, grass
humilis Brain,
ff. Species found in other parts of the world.
g. Pygidium with anus located cephalad of vulva.
h. Pygidium with four plates on each lateris, plates
stout, arranged 0, 1, 5-6, caudal plates much short-
er than cephalic, those of each lateris arranged
l-l-lor2-2or3 ; median pair of lobes small, di-
vergent, serrate; second pair of lobes incised,
mesal lobelet larger than lateral, entire, rounded,
spatulate; brevaceratubae with two oraceratubae
in each second row, third row with 4 in mesal
and 3 in lateral group, fourth row with 5 mesal
and 4 lateral; anus nine times its width from
caudal margin and less than once cephalad of
vulva; mesospiracerores present; genacerores 4-8
(9-16)11-18. — Ceylon on Eurycles, Myristica,
mango. dilata Green.
hh. Pygidium with three plates on each lateris, plates
dagger-like, arranged 0, 1, 3, those of each lateris
arranged 1-1-1; median pair of lobes broad, not
prominent, confluent at proximal end, divergent,
slightly constricted at middle, distal end truncate,
entire or faintly serrate; second pair of lobes in-
cised, small, mesal lobelet largest, triangular;
anus located eleven times its width from caudal
margin and four times cephalad of vulva; breva-
ceratubae with three mesal oraceratubae and five
lateral; genacerores 4-5(11-13)15-17. — Australia on
Leptospermum. angusta Green.
gg. Pygidium with anus and vulva superimposed.
h. Pygidium with seven or more plates on each lat-
eris.
i. Pygidium with genacerores 6-8(10-12)8-12; me-
dian pair of lobes large, oblique, confluent at
proximal end, mesal margin serrate; second
pair of lobes incised, mesal lobelet three times
size of lateral; plates slender, arranged 0, 1,
7, those of each lateris arranged 1-1-1-4; anus
and vulva superimposed, anus five times its
width from caudal margin. — North Carolina on
Nyssa nyssae Comst.
ii. Pygidium with genacerores 7-8(14-19)14-16;
median pair of lobes large, oblique, confiuent
at proximal end, mesal margins serrate; sec-
ond pair of lobes incised, mesal lobelet larger
than lateral, margins entire; plates longer
than lobes, arranged 0, 1, 8, those of each lat-
352 THE COCCIDAE
eris arranged 1-1-1-6; anus and vulva probably-
superimposed, anus eight times its width from
caudal margin, located on line between pre-
genacerores and postgenacerores ; brevacera-
tubae with oraceratubae prominent, second
row with 1 oraceratuba, third row with 4
mesal and 5 lateral, fourth row with 4 mesal
and 4 lateral, and fifth row with 4 mesal and
6 lateral; altaceratubae large, prominent, six
on each side; mesospiracerores present. —
Chionaspis Candida Banks. — Philippine Islands
on Cocos. inday Banks.
hh, Pygidium with four plates on each lateris, plates
dagger-shaped, arranged 0, 1, 4, those of each lat-
eris arranged 1-1-1-1; median pair of lobes promi-
nent, oblique, confluent at proximal end; second
pair of lobes incised, mesal lobelet much larger
than lateral, distal portion bluntly rounded, en-
tire; altaceratubae prominent, six on each side;
genacerores 6-6(14-15)10-11. — Italy on asparagus—
berlesii Leon.
dd. Pygidium without plates in second incisurae, plates slender,
increasing in size toward cephalic end of lateres, arranged
0, 0, 7, those of each lateris arranged 1-1-1-4; median pair
of lobes large, prominent, projecting, bluntly rounded, broad-
er than long, minutely serrate; second pair of lobes in-
cised, small, triangular, entire, mesal lobelet slightly larger
than lateral; brevaceratubae with oraceratubae of third row
with 4 mesal and 5 lateral oraceratubae; altaceratubae five
on each side; anus located near mesogenacerores, evidently
cephalad of vulva, about twelve times its width from caudal
margin; genacerores 10-12(25-26)20-27. — Ceylon on Artocar-
pus. suhcorticalis Green.
bb. Pygidium with a single pair of lobes, divergent, forming narrow
margin to pygidial incisura, distal ends prominent and broadly
truncate; plates dagger-shaped, arranged 0, 4, those of each lat-
eris arranged 1-1-1-1, plate adjacent median lobe small; breva-
ceratubae with oraceratubae of third, fourth, and fifth rows ar-
ranged 4 mesal and 4-5 lateral; anus six times its width from
caudal margin, located near mesogenacerores, evidently cephalad
of vulva; mesospiracerores about 10; dorsum of mesothorax with
transverse row of sharply pointed, hooked spines with dilated
bases and three or four similar spines on ventral aspect caudad
of each metaspiracle ; genacerores 4-6(10-16)24-28. — New South
Wales on Eucalyptus. Formosa Green.
SPECIES OF CANASPIS "^
Body of adult female ovate, broadest through mesothorax; lateral mar-
gins of mesothorax and metathorax with one or two conical processes
provided with ceratubae; pj^gidium with median pair of lobes small,
TRIBE DIASPIDINI 353
divergent, pointed, irregularly dentate; second pair of lobes incised,
minute, inconspicuous; plates stout, arranged 0, 1, 3, those of eachr
lateris arranged 1-1-1-0-0; genacerores 3-4(6-6)8-8; spiracerores want-
ing; anus located slightly cephalad of vulva. — Ceylon on Arundinaria.—
arundinariae Green.
SPECIES OF COOLEYASPIS
Body greatly elongated, length four times its width, thorax two and
one-half times as long as wide; median pair of lobes rounded, not
oblique, serrate; second pair of lobes incised, mesal lobelet much
larger than lateral, prominent, projecting farther caudad than median
pair, margin of lobelets rounded at distal end, entire; plates spine-
like, arranged 0, 1, 4, those of each lateris arranged 1-1-1-1; genacerores
4-5(8-10)23-30; mesospiracerores 4-5, metaspiracerores wanting;
brevaceratubae prominent on pygidium and preabdomen; anus eight
times its width from caudal margin; cephalad of anus transverse
group of 14-15 genaceroris-like pores. — Uganda, Africa
praelonga Newst.
SPECIES OF SINISTRASPIS
Body with sinistral margin of preabdominal segments bearing large
spine-like setae on prominent tubercles, each with brevaceratuba ; pygi-
dium with deep pygidial incision; median pair of lobes oblique, mesal
margins oblique, finely serrate; second pair of lobes incised, mesal
lobelet longer than broad and projecting caudad of median pair of
lobes, distal end bluntly rounded, entire, lateral lobelet similar in
shape to mesal, smaller; plates few in number, arranged 0, 0, 3, those
of each lateris arranged 1-1-1; altaceratubae eight on each side; spira-
cerores wanting; anus nearly twelve times its width from caudal mar-
gin; genacerores 0-2(5-6)5-8. — Barbados on palm (?Thrinax)
unilateralis Newst.
SPECIES OF AUGULASPIS
Pygidium of adult female with deep angular pygidial incision; median
pair of lobes small, each forming the end of a projection limiting one
side of pygidial incision, longer than broad, bluntly pointedly round-
ed; other pairs of lobes wanting; plates arranged 0, 2-3, those of each
lateris located one near median lobe and two or three near cephalic
end, plates small, blunt, rudimentary; brevaceratubae forming two
distinct broad bands; anus located near mesogenacerores, vulva lo-
cated slightly caudad of anus; mesospiracerores and metaspiracerores
forming more or less crescentic group of many cerores; marginal and
submarginal altaceratubae present; genacerores 26-26(19-22)32-44. —
German East Africa. nudata Newst.
SPECIES OF GRAMENASPIS
Pygidium with median pair of lobes low and broad, broader than long,
rounded, entire; second and third pairs of lobes incised, mesal lobe-
let much larger than lateral, approximating median lobe in size, all
entire, broadly rounded ; plates wanting in all incisurae and on lateres ;
354 THE COCCIDAE
anus located between postgenacerores ; dorsal oraceratubae arranged
in two short series; mesospiracerores 4-5; setae longer than lobes;
genacerores 14-14(18-19)16-18. — South west Africa on "grass-like
plant." africana Newst.
SPECIES OF CONTIGASPIS
a. Pygidium with genacerores 8-9(12-14)13-15; median pair of lobes
short, broad, broader than long, adjacent, sometimes appearing as
if fused; other lobes wanting; plates wanting; setae adjacent each
median lobe very long, others on lateres shorter; lateres irregularly
crenulate; marginal and submarginal altaceratubae present; dorsal
oraceratubae numerous and extending to margin; anus located be-
tween postgenacerores; mesospiracerores 4, metaspiracerores want-
ing.— Southwest Africa. suhnudata Newst.
aa. Pygidium with genacerores 2-4(9-12)14-22; median pair of lobes
distinct, fused at proximal end, lateral margin oblique, longest, dis-
tinct lateral notch, mesal margin oblique; setae about as long as
lobes; plates wanting; lateres irreglarly crenulate; altaceratubae
small, seven on each side, arranged 2-2-3; anus and vulva not de-
scribed.— South Africa on Scutia scutiae Brain.
SPECIES OF NATALASPIS
Pygidium without lobes or plates, margin coarsely corrugate or undu-
late, mesal portion prolonged, rounded; setae normal in size; oracera-
tubae not abundant; genacerores 18-25(24-32)48-60. — Adult female con-
tains well developed embryos. — Chionaspis simplex Brain (not Green). —
Natal, Africa on bamboo simplex Brain.
SPECIES OF PARACHIONASPIS
Pygidium with median pair of lobes prominent, conical processes, pro-
jecting, mesal margins entire, lateral serrate; second pair of lobes
incised, lobelets serrate, subequal, each subequal to a median lobe,
mesal margin of mesal lobelets entire, lateral margin and both mar-
gins of lateral lobelets serrate; plates minute, arranged 0, 1, 3, short-
er than lobes, those of each lateris arranged 1-1-1; lateres with deep
indentations; genacerores 18-22(24-32)20-30; mesospiracerores and
metaspiracerores present; altaceratubae apparently three on each side;
brevaceratubae promiscuously arranged; anus more than its own width
cephalad of vulva; forming galls on young stems, petioles and mid-
ribes of leaves. — Ceylon on Hedyotis. galliformens Green.
SPECIES OF PARLATOREOPSIS
Antennae with single strongly curved and deeply forked spine; pygidium
with median pair of lobes large, bluntly rounded, mesal notches want-
ing, two lateral notches; second pair of lobes smaller, as long as
broad, not incised, mesal margin straight, lateral margin obliquely
rounded, notched or dentate; plates arranged 0, 0, 2, plates located
near but not adjacent to second pair of lobes, appear like incised third
lobe, not longer than lobes, bluntly pointed; setae very long, particu-
larly those of cephalic half of each lateris; each lateris with two
TRIBE DIASPIDINI 355
prominent latadentes; vulva located near middle of pygidium, anus
between vulva and caudal margin; genacerores single, transverse curved'
row of three or four cerores; insects insinuate themselves beneath
cuticle of food-plant. — Egypt on Justicia longispina Newst.
SPECIES OF POLIASPIS
a. Pygidium with nine groups of genacerores, arranged 1-1(18-21)25-35,
four other groups forming cephalic arch, lateral group of each side
8-12, two mesal groups 1-2 cerores; brevaceratubae numerous; me-
dian pair of lobes minute, scarcely perceptible; apparently without
plates. — Australia on Exocarpus, Santalum, Leptospermum, Ixylo-
bium, Dillwynia. exocarpi Mask.
aa. Pygidium with less than nine groups of genacerores.
b. Pygidium with eight groups of genacerores.
c. Pygidium with three pairs of lobes,
d. Pygidium with second and third pairs of lobes not incised,
lobes similar in form, second pair smaller than median and
larger than third pair; median pair of lobes broad, broadly
rounded, distant, not divergent or confluent, entire; plates
apparently wanting; genacerores in eight groups, arranged
10-10(10-12)12-12, three other groups in cephalic transverse
arch, mesal group of about 10 and lateral groups of about 4
cerores; brevaceratubae numerous. — Japan on Pinus, Abies.__
pini Mask,
dd. Pygidium with second and third pairs of lobes incised,
lobelets subequal, all minute, blunt, triangular, second pair
of lobes represented by three lobelets, cephalic lobelet prob-
ably latadentis; median pair of lobes long, projecting, long-
er than broad, blunt, subtriangular, mesal and lateral
notches, distant, not divergent, not confluent; plates long
and slender, arranged 0, 0, 1, 1; genacerores in eight groups,
arranged 3-5(9-14)20-25, other groups in transverse row,
mesal group of 2-4, each lateral group of 7-8. — West Australia
on leguminous plant intermedia Fuller.
cc. Pygidium with less than tLree pairs of lobes,
d. Pygidium with two pairs of lobes.
e. Pygidium with median pair of lobes confluent at proximal
end.
f. Pygidium with second pair of lobes incised, rounded,
margins entire, mesal lobelet larger than lateral; me-
dian pair of lobes confluent, at proximal end, large,
slightly divergent, finely and widely serrate or entire;
plates slender, arranged 0, 1, 1, 8-10, caudal plates
longer and more slender than cephalic, those of each
lateris arranged l-l-l-5to7, caudal group short and
broad at proximal end; genacerores arranged in eight
groups, 2-4(10-13)21-25, other three groups in trans-
verse row, each group 2-4; mesospiracerores about 12,
metaspiracerores 4-6; altaceratubae seven on each side;
356 THE COCCIDAE
anus located on line drawn through cephalic end of
postgenacerores. — District of Columbia, England on
Cycas, Dion, palms cycadis Comst.
ff. Pygidium with second pair of lobes small, not incised,
long, narrow, finely serrate; median pair of lobes nar-
row, divergent, confluent at proximal end, serrate;
plates four on each side of median lobes, simple, spine-
like, caudal three situated between median lobes and
third row of brevaceratubae, arranged 0, 0, 4; breva-
ceratubae with 2 oraceratubae in second row, third row
with 4-6 mesal and 5-7 lateral, fourth row with 4-5 mesal
and 6-7 lateral, and fifth row with 5 mesal and 6 lateral
oraceratubae; anus located near mesogenacerores, seven
times its width from caudal margin and about twice
cephalad of vulva; mesospiracerores 6-10, metaspira-
cerores 4-6; genacerores in eight groups, 2-4(14-20)21-
25, three cephalic groups of 2-5 each. — New South Wales
on Coprosoma. argentosis Brittin.
ee. Pygidium with median pair of lobes as far apart as width
of a median lobe.
f. Pygidium with plate in each second incisura, plates
twice as long as lobes, arranged 0, 1, 2; median pair of
lobes distant, distal margin rounded, serrate; second pair
of lobes incised, mesal lobelet twice as large as lateral,
projecting slightly beyond median, distal end blunt,
lateral lobelet triangular; brevaceratubae prominent;
genacerores in eight groups, 2-5(11-17)17-27, three
groups cephalad of mesogenacerores, distant, in trans-
verse row, 3-4 cerores. — Natal on Carissa
carissae Ckll.
ff. Pygidium without plates in second incisurae, plates
small, inconspicuous, arranged 0, 0, 2; median pair of
lobes rounded, broad, distant, angles rounded, margins
entire; second pair of lobes small, incised, mesal lobe-
let slightly larger than lateral, triangular, entire; each
lateris with triangular lobe-like projections, arranged in
pairs; mesospiracerores 6-10; genacerores in eight
groups, arranged 1-10(14-20)23-36, three cephalic groups
of 4-9, 1-6, and 5-8 cerores. — Natal, Africa on Kiggelaria,
willow. kiggelariae Brain.
dd. Pygidium with single pair of lobes; median pair of lobes
long, broad, divergent, not confluent, apparently entire;
plates apparently wanting; three or four large setae on
each side may be plates; brevaceratubae represented by three
distinct rows, third to fifth, on each side; anus about six
times its width from caudal margin and nearly twice
cephalad of vulva; lateres irregular; genacerores in eight
groups, arranged 4-6(20-30)20-30, three other groups in trans-
I
TRIBE DIASPIDINI 357
verse row, mesal group of row 3-5, each lateral group 8-10.
— New Zealand on Veronica, Leucopogon, Cyanthodes, ferns.^^
media Mask,
bb. Pygidium with seven groups of genacerores, arranged 3-4(15-17)
16-20, and a group cephalad and in line with pregenacerores, 7-8;
median pair of lobes prominent, divergent, confluent, short, wide,
entire; second pair of lobes minute, triangular, entire; plates
much longer than lobes, arranged 0, 1, 1; lateres serrate. — West
Australia on Daviesia nitens Fuller.
SPECIES OF POLIASPOIDES
Pygidium of adult female without lobes or plates; margin with three
indentations, limited on cephalic side by latadentes; lataceratubae ap-
parently wanting; brevaceratubae irregularly arranged: ventral sur-
face with numerous small pores adjacent to margin; dorsal and ven-
tral surfaces coarsely carinated; genacerores 19-22(30-35)50-55, two
groups caudad of postgenacerores, each consisting of 1-3 cerores; anus
located cephalad of vulva, near cephalic margin of mesogenacerores
and vulva between postgenacerores; scale of adult female mussel-
shaped. — Ceylon on bamboo simplex Green.
SPECIES OF CRYPTASPIDUS
Pygidium of adult female with median lobes short and broad, broader
than long, distal margin with notches and roundly produced between
them; second pair of lobes small, incised, mesal lobelet largest, two
notches on distal margin, lateral lobelet entire; plates large, twice as
long as lobes, arranged 2, 1, 2, those of each lateris located one near
second lobe and one near middle, remainder of each lateris not figured
or described; 8 long dorsal setae and 6 much shorter ventral setae;
latadentes with oraceratubae in each second incisura between plate
and second lobe and cephalad of caudal plate of lateres; mesospira-
cerores 4-7; lateres dentate, irregular; genacerores 6-8(14-17)9-13. —
Madagascar on Cocos. nucum Lindgr.
SPECIES OF CRYPTODIASPIS
a. Pygidium of adult female with plates, never with pectinae, ar-
ranged 0, 1, 1, 6-7, those of each second and third incisura large,
much longer than lobes; median pair of lobes narrow, bluntly point-
ed, serrate; second and third pairs of lobes incised, lateral lobelets
smaller than mesal, all lobelets notched; two altaceratubae on each
side opening in prominent latadentes in second and third incisurae;
genacerores 4-4(8-10)6-8; second nymphal female with three pairs
of lobes and deep pygidial incision; median pair of lobes oblique,
mesal margins serrate, not projecting; second and third pairs of
lobes incised, broadly truncate; plates arranged 0, 1, 1, 2-3; altacera-
tubae four on each side opening in latadentes. — Kamerun, Africa
on Euphorbiaceae. conservans Lindgr.
aa. Pygidium of adult female with furcapectinae in second and third
incisurae; median pair of lobes small, bluntly rounded, faintly
notched on mesal margin; second and third pairs of lobes deeply
358 THE COCCIDAE
incised, mesal lobelet much longer and broader tlian lateral and
truncate; furcapectinae and plates arranged 0, 1, 1, 5, furcapectinae
adjacent to lobes and two or three times as long; altaceratubae
wanting; genacerores 4-4(10-10)8-10; second nymphal female with
four pairs of lobes and deep pygidial incision; median pair of lobes
located in pygidial incision, oblique, mesal margins serrate; sec-
ond, third, and fourth pairs of lobes incised, mesal lobelet much
larger than lateral, entire, truncate; plates only present, arranged
0, 1, 1, 1, 1, 0, subequal in length to lobes; altaceratubae four on
each side, large, opening in latadentes in second, third, and fourth
incisurae and beyond fourth lobes; latadentes nearly as large as
mesal lobelets. — Kamerun on Macrolobium hamata Lindgr.
SPECIES OF FURCHADASPIS
Scale of adult female circular; pygidium of adult female with median
pair of lobes distant, divergent, subprojecting, finely and widely ser-
rate; second and third pairs of lobes subequal, incised, lobelets with
broadly rounded margins, entire or faintly notched, mesal lobelet larg-
er than lateral; pectinae and plates short and stout, arranged 2, 1, 1,
11-12, those of insurae and cephalad of third pair of lobes bifurcate
pectinae, others plates, those of each lateris arranged l-l-2-3or4-2or
3-1; genacerores wanting; mesospiracerores and metaspifacerores 3-6,
latter fewer in number; brevaceratubae numerous, irregularly ar-
ranged; anus located slightly cephalad of middle of pygidium. — Eng-
land, Portugal, Italy, New York, Massachusetts, Bermuda on Zamia.
Cycas. zamiae Morgan.
SPECIES OF MITULASPIS
Pygidium small; median pair of lobes large, distant, not divergent, not
confluent, striate, faintly serrate, mesal and lateral margins converg-
ing, bluntly pointed, constricted at proximal end; second pair of lobes
incised, mesal loblet three or more times as broad as lateral, similar
to median in form, smaller; third pair of lobes small, incised; plates
spiniform, longer than lobes, arranged 2, 2, 2, 3, those of each lateris
arranged 1-1-1, distant from third lobe; anus twenty or more times
its width from caudal margin and about once cephalad of vulva; meso-
spiracerores 5-6, metaspiracerores 2-3; altaceratubae not clearly in-
dicated, 3 on each side. — Uganda, Africa on Funtumia
funtumiae Newst.
SPECIES OF DENTACHIONASPIS
Pygidium with median pair of lobes small, incised, lobelets subequal,
pointed, lobes distant, twice width of lobes apart; second pair of
lobes lobe-like, larger than median pair, not incised, notched, bluntly
rounded; furcapectinae sometimes plates, twice as long as lobes, ar-
ranged 0, 1, 5, those of each lateris arranged 1-2-2; anus located near
cephalic end, vulva slightly cephalad of middle; dorsal oraceratubae
in groups; mesospiracerores 9-11, generally arranged in two or three
groups, metaspiracerores 2; altaceratubae three on each side in two
groups, 1-2. — Transvaal, Africa on Acacia capensis Newst.
TRIBE DIxVSPIDINI 359
SPECIES OF INCHOASPIS
Pygidium of adult female with minute median lobe located on each
side of pygidial incision, triangular, entire, with a prominent oracera-
tuba at its proximo-mesal angle; second, third, and fourth pairs of
lobes incised, mesal lobelet at least twice as large as lateral, margin
broadly rounded, lateral lobelet more angular with an irregular mar-
gin; plates longer than lobes, arranged 0, 1, 0, 2, 1, those of each
lateris adjacent to a fourth lobe, there may be others; second and
third incisurae with two minute adjacent triangular projections, prob-
ably latadentes; dorsal oraceratubae numerous, in rows; spiracerores
wanting; altaceratubae present; second exuvia of female with deep
transverse thoracic constriction. — Southwest Africa
amaniensis Lindgr.
SPECIES OF GRAPHASPIS
Pygidium of adult female with three pairs of lobes; median pair of
lobes fully projecting, curved, divergent, mesal margin finely serrate,
lateral margin entire; second and third pairs of lobes incised, lobe-
lets similar, bluntly pointed, mesal lobelets of second pair of lobes
projecting farther caudad than median pair of lobes; plates small,
not much longer than lobes, arranged 0, 1, 1, 4, those of each lateris
arranged 1-1-2; altaceratubae seven on each side; brevaceratubae
fairly numerous, irregularly arranged; genacerores wanting; anus and
vulva superimposed; scale of adult female pyriform. — Ceylon on
Antidesma. permutans Green.
SPECIES OF AMETROCHASPIS
Scale of adult female short, pyriform; pygidium with three pairs of
lobes; median pair of lobes divergent, not projecting, elongate, placed
in pygidial incision, serrate; second and third pairs of lobes incised,
lobelets constricted at proximal end, distal end broadly rounded, mar-
gins entire, mesal lobelet larger than lateral; plates distinctly longer
than lobes, arranged 0, 1, 1, 4, those of each lateris arranged 1-1-2;
altaceratubae eight on each side; brevaceratubae numerous, irregular-
ly arranged; genacerores wanting; mesospiracerores represented by
small group of pores ; anus and vulva superimposed. — Ceylon on
Antidesma. flava Green,
SPECIES OF PRONTASPIS
a. Pygidium with median pair of lobes with lateral and mesal mar-
gins serrate, divergent, lateral margins subparallel; second and third
pairs of lobes small, subequal, incised, serrate; setae short; plates ar-
ranged 0, 1, 1, 3, each lateris with a plate near a third lobe, one at
cephalic end, and one midway between these two; preabdomen with
caudal segment with 5-6 plates; anus broader than long, about fifteen
times its width from caudal margin and about twice caudad of vulva;
altaceratubae about four on each side. — Southern States, West Indies,
Mexico, New Zealand, Australia, Japan, Samoa on Euonymus, Osman-
thus, orange, palms. citri Comst.
aa. Pygidium of adult female with all lobes entire; median pair of
lobes distant, divergent, distal end round; second and third pairs of
360 THE COCCIDAE
lobes incised lobelets of second pair larger than those of third, mesal
lobelet of both pairs larger than their lateral lobelets; plates slightly-
longer than lobes, arranged 0, 1, 1, 10-11, those of each lateris ar-
ranged l-l-3-5or6; latadentes bearing oraceratubae located in median,
second, and third incisurae, one just cephalad of each third lobe and
near middle of lateres; spiracerores wanting. — Canary Islands on
Euphorbia. harrancorum Lindgr.
SPECIES OF REMOTASPIS
Scale of adult female white, highly convex, mytilaspiform ; exuviae
orange-red; cuticle of body, including pygidium, soft and thin; pygl-
dium with median pair of lobes widely separated, as broad as long,
broadly rounded, entire except for a distal notch; second pair of
lobes incised, mesal lobelet similar in form but larger than a median
lobe, lateral lobelet much smaller than mesal, one or two deep notches,
so that lobelet appears to have two or three teeth; third pair of lobes
deeply notched, mesal lobelet similar in size and form to a median
lobe, lateral lobelet smaller, dentate; triangular latadentis caudad of
each second lobe and three toothed ones cephalad of third lobe; alta-
ceratubae four on each side; plates limited to one in each second
incisura, arranged 0, 1, 0, 0; dorsal aspect with two broad bands of
oraceratubae; anus located opposite "distal ends" of dorsal oracera-
tubae.— Uganda, Africa on unknown shrub dentiloHs Newst.
SPECIES OF ASYMMETEASPIS
a. Pygidium with second pair of lobes nearly as large as median pair,
lateral margin oblique, not incised; median pair of lobes large,
broader than long, mesal margins oblique, divergent, distant, distal
margins sometimes dentate, subadjacent; dorsal oraceratubae few,
in two transverse crescentic rows; plates inconspicuous, arranged
0, 0, 2-3, frequently broken; mesospiracerores 4-5; body with ce-
phalic half strongly chitinized, caudal half including pygidium not
strongly chitinized, line between two areas sharply marked; setae
small, arranged two in median incisura, one in each second, and
one on cephalic part of each lateris; four submarginal setae cephalad
of lobes; anus eleven times its width from caudal margin and
nearly four times cephalad of vulva. — Transvaal, Africa
distorta Newst.
aa. Pygidium with second pair of lobes always smaller than median pair,
b. Pygidium with median pair of lobes serrate, broader than long,
short, very distant; second pair of lobes incised, mesal lobelet
larger than lateral, longer than a median lobe, much narrower,
bluntly rounded, entire, lateral lobelet subtriangular, generally
notched on both margins; plates long, slender, arranged 0, 1, 2,
those of each lateris located one near second lobe, other at
cephalic end; surface with several large clear vacuole-like oval
areas; mesospiracerores 1-3, mesosperitreme depressed; surface of
cuticle reticulate; altaceratubae apparently five on each side;
brevaceratubae distinct; anus over twelve times its width from
caudal margin. — Uganda, islands of Lake Victoria, Africa
tenuidisculis Newst.
TRIBE DIASPIDINI 361
bb. Pygidium with median pair of lobes entire, about as broad as_
long, mesal and lateral margins subparallel, distal end broadly
rounded; second pair of lobes minute, inconspicuous, not incised,
distal end with two notches; plates not longer than lobes, ar-
ranged 0, 0, 7, those of each lateris arranged 1-1-1- ( 2-2 ) ; alta-
ceratubae apparently not prominent; brevaceratubae distinct;
mesospiracerores 3-4, metastpi^acerores cssranting; anus; located
about eighteen times its width from caudal margin and about
three times cephalad of vulva; body elongate; caudad of and in
line with antennae, on each side a row of two or three blunt spiny
processes and pair of long setae. — Uganda, islands of Lake Vic-
toria, Africa on unknown tree dura Newst.
SPECIES OF OBLUCTASPIS
Pygidium with median pair of lobes minute, inconspicuous, distal end
concave, mesal angle prolonged, sides parallel; second pair of lobes
smaller than median, triangular, pointed, lateral margin oblique;
plates small, short, broadened at proximal end, arranged 0, 1, 2, those
of each lateris distant; brevaceratubae small, smaller than lamacera-
tubae, promiscuously arranged; anus eight times its width from
caudal margin and three times cephalad of vulva. — New Mexico on
Quercus. lohata Ferris.
SPECIES OF SITULASPIS
Pygidium with median pair of lobes large, prominent, broader than long,
distant, broadly rounded, lateral notch; second pair of lobes adjacent
to median pair, minute, bluntly rounded, as long as broad, one-fifth
width of median; plates slightly longer than median pair of lobes,
broadest at proximal end, arranged 0, 0, 1, adjacent second lobe;
altaceratubae wanting; brevaceratubae promiscuously arranged; anus
located four times its width from caudal margin and six times caudad
of vulva; setae small. — ^Arizona on Condalia condaliae Ferris.
SPECIES OF KUWANASPIS
Scale of adult female very long and slender, sides parallel and usually
straight; second exuvia distinct; pygidium with median pair of lobes
short, divergent, slightly notched on mesal margin; latadentes round-
ed, serrate; plates large, much longer than median lobes, arranged
0, 3, those of each lateris placed one near a median lobe, one near
where third incisura would be located, and one midway between caudal
and cephalic plates; anus located cephalad of middle of pygidium;
dorsal oraceratubae numerous on caudal portion; latadentes few in
number. — Japan on Phyllostachys. hikosani Knw.
SPECIES OF INCISASPIS
Pygidium with median pair of lobes distant, broadly divergent, project-
ing, oblique mesal margin with four to six notches; plates placed near
median pair of lobes, caudal three of each side shorter than others,
about 12 on each side, stout, blunt, usually curved; setae on dorsal
aspect 6, minute; mesospiracerores 2. — South Africa on Mitragyne
pugionifera Lindgr.
362 THE COCCIDAE
SPECIES OF CORONASPIS
a. Pygidium with median pair of lobes rounded, not tricuspid; two
or three pairs of minute inconspicuous bluntly lanceolate lobes,
median pair simple, second pair incised; plates prominent, ar-
ranged 0, 1, 8, those of each lateris arranged 1-1-1-1-4; dorsal ora-
ceratubae large, conspicuous, arranged in more or less definite
rows; anus broader than long, about eight times its width from
caudal margin; body after gestation reddish and densely chitin-
ous; antennae consisting of corona-like area bearing four to six
stout spines and central curved seta; caudad of each antenna and
scattered over thoracic region oval translucent spaces; margin of
preabdominal and caudal thoracic segments with numerous ora-
ceratubae, obscure in densely chitinous individuals; scale of adult
female white, sometimes tinged with ochreous, exuviae reddish,
sides of scale sloping up and forming median longitudinal rounded
ridge. — Ceylon on undetermined tree coronifera Green.
aa. Pygidium with median pair of lobes small, mesal and lateral mar-
gins subparallel, distal end tricuspid; second pair of lobes incised,
mesal lobelet large, lobe-like, much larger than a median lobe, mar-
gin rounded, entire, lateral lobelet tricuspid, similar in form to a
median lobe; plates slender, longer than lobes, arranged 0, 1, 2,
those of each lateris located one near second lobe and other at
cephalic end; altaceratubae 5-6 on each side; lateres serrate be-
tween altaceratubae ; antennae adjacent cephalad of head skeleton,
each corona-like, margin with long spiny process, two longer than
others, and two long setae; mesospiracerores 3, mesoperitreme
very large and depressed; scale of adult female elongate, strongly
convex, pure glistening white, finely laminate, exuviae orange yellow
to orange brown; male scale strongly carinate, secretionary portion
dull gold color. — Uganda, islands of Lake Victoria, Africa
auratilis Newst.
SPECIES OF DENTASPIS
Scale of adult female satiny-white, transversely striate, striae equidis-
tant and distinct, sides subparallel; body with cephalic region nar-
rowed; pygidium with median pair of lobes distant, not located in
pygidial incision, projecting, distal margin deeply serrate; second
pair of lobes distant from median pair twice its width, similar in
form, smaller, incised, deeply serrate; third pair of lobes only slight
projection on margin, serrate, not incised; plates longer than lobes,
arranged 0, 1, 1, 1-2, those of each lateris near third lobe; altacera-
tubae with tubes about as long as wide, arranged two cephalad of
plate in second incisura, one associated with each second lobe and
one near each third lobe. — Uganda, Africa on palms.. -sw&striaia Newst.
SPECIES OF HOWARDIA
Scale of adult female broadly oval, caudal end narrowed, often almost
circular, exuviae terminal; pygidium with median pair of lobes large,
prominent, mesal angle roundly protuberant, lateral margin finely ser-
rate, mesal margin substraight, not adjacent; second pair of lobes
not incised, small, bluntly pointed, entire; lateres with serrate pro-
TRIBE DIASPIDINI 363
Jections; plates stout, arranged 2, 2, 18-20, those of each lateris ar-
ranged 3-3or4-4or5-6 ; mesospiracerores and metaspiracerores pres-
ent; brevaceratubae numerous, irregularly arranged; paraphyses cluB-
shaped, two or three times as long as median pair of lobes, cephalic
end of each enlarged, knob-like, extending from median incisura; anus
located its own width cephalad of vulva. — Eastern United States, Cali-
fornia, West Indies, Mexico, England, Ireland, Hawaiian Islands,
Ceylon, Mauritius, Japan, Tahiti on orange, lime, tea, tamarind, Fla-
courtia, Cinchona, Ochrea, Ficus, Anona, Hibiscus, Microglossa
Mclavis Comst.
SPECIES OF PROCERASPIS
Scale of adult female elongate, narrow, flat, more or less distinct me-
dian carina; body of adult female elongate, thorax occupying nearly
three-fourths of length of body, metathoracic spiracles located near
middle of caudal half; pygidium with median pair of lobes contigu-
ous, two lobes together much broader than long, lateral margins
oblique, minutely serrate; plates near median lobes smaller than those
near cephalic end, arranged 0, 7, those of each lateris arranged
1-1-1-1-3; latadentis on each side between caudal first and second plates
and two between second and third; paraphyses club-shaped, short, ex-
tending to margin near lateral proximal angle of each median lobe;
dorsal oraceratubae wanting; lateres irregularly indented; anus ap-
proximately central in position. — Ceylon on Cinnamomum
cinnamomi Green.
SPECIES OF MARCHALASPIS
Pygidium with median pair of lobes distinct, entire, proximal portion
strongly constricted, distal portion broadly rounded; second and third
pairs of lobes incised, entire, lobelets subequal in size, similar in form,
proximal portion strongly constricted, distal portion truncately round-
ed; median, second, and third incisurae each with triangular projection
about as long as lobes, each identified as a plate; each lateris with
six or more lobe-like serrate projections; altaceratubae 9 on each
side; dorsal oraceratubae few, poorly developed and indistinct; anus
located midway between vulva and caudal margin. — Guinea on Copai-
fera. vuilleti Marchal.
SPECIES OF CUPIDASPIS
Pygidium with median pair of lobes small, mere prominences, broadly
rounded, entire; second pair of lobes not incised, large, not project-
ing strongly, broadly rounded, entire; pectinae or plates wanting;
lateres with angular projections; altaceratubae prominent, five or six
on each side, one in each second incisura; anus large, six times its
width from caudal margin and over twice cephalad of vulva; breva-
ceratubae distinct, apparently third row with 4 oraceratubae, fourth
row with 5 mesal and 2 lateral oraceratubae. — California on Cupressus.
cupressi Colem.
SPECIES OF PSEUDODIASPIS
a. Pygidium with two pairs of lobes; median pair of lobes broad, distal
end squared, "resembling human incisor teeth," distinctly separated;
364 THE COCCIDAE
second pair of lobes hardly half length of first, much narrower; each
lateris with angular prominence like third lobe, cephalad of this irreg-
ularly crenulate; setae large; plates cephalad of second and third
lobes quite large, broad; scale of adult female convex, exuviae lateral,
within margin, first exuvia at margin of second. — Mexico on Mimosa,
?Acacia. dentiloMa Ckll.
aa. Pygidium with one pair of lobes.
b. Pygxdium with median pair of lobes broad, low, broader than
long, close together, not adjacent, hardly projecting; plates want-
ing; lateres irregularly dentate, crenulate between dentations;
apparently with three or four altaceratubae ; scale of adult female
flat, irregular, round to suboval, exuviae lateral to sublateral;
scale of male mytiliform, exuvia at end. — Arizona on Larrea
larreae Ckll.
bb. Pygidium with median pair of lobes large and prominent, not
broader than long.
0. Pygidium with anus located near vulva, five times its width
from caudal margin and eighteen times its width caudad of
vulva; median pair of lobes large, prominent, mesal margins
parallel, near together, prominent lateral notch, distal end
rounded; setae as long as lobes; plates short, dagger-shaped,
arranged 0, 3, those of each lateris distant and located on
caudal one-fourth of pygidium; altaceratubae apparently want-
ing; dorsal oraceratubae few in number, promiscuously ar-
ranged; mesal calles present and fused, lateral calles short
and distant from mesal. — Apparently not congeneric. — California
on Atriplex. atriplicis Ferris.
cc. Pygidium with anus and vulva superimposed, about eleven
times its width from caudal margin; median pair of lobes
large, widely separated, prominent, broader than long, mar-
gins broadly rounded, crenulate; lateres broadly irregularly
crenulate, caudal crenulation in position of second pair of
lobes; plates wanting; altaceratubae apparently wanting; dor-
sal oraceratubae fairly numerous, small, irregularly arranged.
— Apparently not congeneric. — California on Phoradendron
multipora Ferris.
SPECIES OF VERSICULASPIS
Pygidium with deep angular pygidial incision; median pair of lobes
located in pygidial incision, not projecting beyond its margin, dis-
tant, suboblique, divergent, not confluent, margin serrate, subtruncate;
second pair of lobes apparently incised, triangular, bluntly pointed;
plates apparently wanting; altaceratubae small, slender, apparently
numerous; brevaceratubae apparently few in number, — South Africa
on Diosma. diosmae Brain.
SPEaES OF PROTODIASPIS
a. Pygidium with two pairs of lobes.
b. Pygidium with four very low, broad, rounded, colorless lobes,
distant, margins inclined to be crenulate; plates wanting; setae
TRIBE DIASPIDINI 365
very small; cuticle with numerous oraceratubae ; mesogenacerores
about 11; anus distinct, fair distance from caudal margin, cuticle
around anus strongly concentrically striate. — Mexico on oak
parvula Ckll.
bb. Pygidium of adult female with four lobe-like projections; median
pair of lobes near together, broader than long, broadly rounded,
entire; second pair of lobes incised, mesal lobelet largest, mar-
gins rounded, entire; dorsal oraceratubae numerous, small, irreg-
ularly arranged; anus large, about fourteen times its width
cephalad of caudal margin and about once cephalad of vulva;
setae minute; pygidium of second nymphal female with four
minute distant subequal lobes, distance between lobes about
three times width of lobe; setae few and minute; anus prominent,
about five times its width from caudal margin; plates wanting. —
Arizona on Quercus. pulchra Ferris.
aa. Pygidium without lobes or lobe-like projections.
b. Pygidium as broad as or broader than long, margin broadly
rounded, not pointed.
c. Pygidium with margin coarsely irregularly crenulate, without
lobes or plates; setae four in number, moderate in length;
anus located distant from caudal margin, around anus a num-
ber of conspicuous thin spots in cuticle, appearing as perfora-
tions; pygidium of second nymphal female with three plates
on each side; spiracerores wanting; surface of body finely
carinate. — South Africa on Euclea imhricata Brain.
CO. Pygidium with margin entire, smooth, broadly rounded, bear-
ing four minute distant setae; anus small, about eighteen times
its width from caudal margin and four times caudad of vulva. —
Resembles species of Gymnaspis or Aonidia. — Arizona on
Acacia. edentata Ferris.
bb. Pygidium longer than broad, distal portion pointed, never broad-
ly rounded; body of adult female more or less globose; pygidium
flattened, cordate, acuminate; lobes and plates wanting; margin
with two irregular teeth-like projections at caudal end; anus
small, near cephalic end, twenty or more times its width from
caudal margin; setae minute, one on each side near cephalic end
of margin, few on disk; many linear thickenings radiating in-
wards from margin; pygidium of second nymphal female broad,
rounded, hinged, semicircular plate; setae, lobes, plates, wanting;
similar to preceding species. — ^Australia on Acacia
anomala Green.
SPECIES OF NUDACHASPIS
Insects forming galls upon leaves and stem of host-plant, external open-
ing to gall typically closed by first exuvia, second exuvia thin, enclos-
ing adult puparium-like in early adult, ruptured later and insect ex-
truded; body of insect "comma-shaped," enlarged cephalic portion bent
at right angles to caudal portion; pygidium of adult female without
lobes, plates, or setae; with seven serrate lobe-like projections on each
side, probably pseudolobes, projections disappear with age; altacera-
366 THE COCCIDAE
tubae and brevaceratubae wanting; mesospiracerores and metaspira-
cerores present; anus and vulva located near cephalic end of pygidium;
pygidium of second nymphal female with pair of median lobes, project-
ing, lateral margins free, triangular in outline, confluent at proximal
end, second and third pairs of lobes much smaller, distal margin
notched at middle. — Ceylon on Loranthus fodiens Green..
SPECIES OF XANTHOPTHALMA
Insects with body nearly circular, less than one-half millimeter in
length; caudal end of abdomen minutely serrate with three pairs of
serrate triangular lobe-like projections with setae at their distal ends,
arranged like lobes; anus located over three times its width from
caudal margin; female probably transforms in a puparium. — Mexico
on "laureltree." concinnum Ckll. & Parr.
SPECIES OF CREDODIASPIS
Pygidium with adult female with median pair of lobes large, longer than
broad, distal end bluntly rounded, sides parallel; other lobes obsolete;
each lateris with three long dagger-shaped plates, largest adjacent to
each third lobe, smallest near middle of each lateris, other plate nearly
midway between cephalic and caudal plates, slightly nearer caudal;
spiracerores and genacerores wanting; two minute latadentes bearing
oraceratubae located between median pair of lobes and an oracera-
tuba on each side between two caudal plates; pygidium of second
nymphal female with four pairs of lobes, median pair oblique, mesal
margins with two or three indentations; second, third, and fourth
pairs of lobes quadrangular, subequal, entire; plates arranged 0, 1,
1, 1, 5, cephalic plate of each lateris similar in form to those of in-
cisurae, others broad, pointed, sides crenulate; oraceratubae located
in second, third, fourth incisurae and between two caudal plates of
each lateris.— South Africa on Cynometra limuloides Lindgr.
SPECIES OF ADISCODIASPIS
Pygidium of adult female without lobes, setae, plates, or genacerores;
ceratubae short, numerous, arranged in a series or in a mass on the
pygidium and on the border of the segments of preabdomen; scale
rounded, ostreiform, white with yellow first exuvia, placed near per-
iphery, ventral scale thick and flat. — France on Erica
ericicola MarchaL
CHAPTER XXVI
tRIBE FIORINIINI
The Uhlerian Scales
The body of the adult female is always distinctly longer than broad
with the cephalic end bluntly rounded and the caudal end bluntly pointed.
The lateral margins are usually parallel or subparallel with the inden-
tations marking the position of the sutures between the thoracic and
abdominal segments generally more or less distinct. The lateral m.ar-
gins of the segments of the preabdomen are sometimes provided with
bracteae in the adult female or second nymphal female. Lobes are
usually present in the adult female, usually about two pairs, more or
less than this number may be present. Plates although frequently pres-
ent, may be absent; furcapectinae are never present. The incisurae are
narrow and the spaces may be filled or not with plates. Densariae and
paraphyses are never present. Altaceratubae are rarely absent and are
not arranged in pairs, their oraceratubae are ovate in outline, always
longer than broad. Brev'aceratubae are rarely if ever present, never ar-
ranged in rows. Genacerores are frequently present, usually arranged
in five or three groups. The mesogenacerores, when present, usually
consist of about five cerores. The scale of the adult female is thin,
brownish or yellowish in color. It is much longer than broad, two or
more times, usually with the sides parallel, not enlarged at one end,
never transversely ridged, sometimes with a mesal carina. The first
exuvia projects beyond the cephalic end of the secretionary portion and
bears the porrect remains of the antennae. The wax is limited to the
periphery and is small in amount. The female always transforms in a
puparium and the pygidium of the adult female may be provided with
pseudolobes. The scales are usually exposed upon the host-plant, but
may be concealed in pits in the leaf or in galls. The scales produced in
galls are frequently opaque white, consisting in great part of wax and
the exuvia is one-fourth or less the length of the scale. The surface may
be flat, slightly convex, or slightly carinate.
The first species of this tribe was described by Targioni
Tozzetti in 1867 as Diaspis fioriniae. The following year he erected
a new genus, Fiorinia, for this species and renamed the insect
Fiorinia pellucida. This latter name was used by Signoret and by
Comstock who described a second species, Fiorinia camelliae, which
was a serious pest in the conservatories of the United States
Department of Agriculture in Washington. In 1883 Comstock
called attention to the fact that Targioni Tozzetti had no authority
367
368 THE COCCIDAE
for changing the name of his species and restored the original name
of the insect, Fiorinia fioriniae. There was at this time a rule of
the British Association code, which was generally followed in mat-
ters of nomenclature, prohibiting the use of the same word for both
generic and specific names. Comstock consequently renamed the
genus after one of the early American entomologists who had
made a special study of the Hemiptera, Philip Uhler, designating
the genus Uhleria. This rule regarding the use of the same word
for both names of an animal was afterwards rescinded and the
species of Targioni Tozzetti is now known as Fiorinia fioriniae.
The camelliae of Comstock has also been shown to be the same as
the fioriniae of Targioni Tozzetti. The selection of a vernacular
name for the insects of this tribe has been a difficult one. It was
finally decided to form a name out of the generic name of Com-
stock and call them The Uhlerian Scales.
Leonardi included the genus Fiorinia together with the genera
Mytilaspis and Ischnaspis in the tribe Mytilaspides in his original
subdivision of the subfamily Diaspidinae published in 1898. In
the same volume, where there is given the first statement of his
monograph of the Mytilaspides, he states that the following table,
one to genera and subgenera, relates only to the genus Mytilaspis,
because he combines Ischnaspis with this genus, rather than to the
Mytilaspides and that the lacking genus Fiorinia will be considered
a little later. The Mytilaspides was published in 1903 in which
he states, to translate the thought rather than the words, the
group Mytilaspides is conveniently distinguished as two sections,
one formed from the species of the genus Mytilaspis and the other
from the species of the genus Fiorinia. The essay on the classifi-
cation of the Fioriniini, '' Generic e Specie di Diaspiti. Saggio di
sistematica delle Fioriniae" was published in 1906. In this re-
vision four genera and twenty-two species are recognized, while
ten unidentified species are referred to the tribe.
The scale of the adult female is elongate, two or more times
as long as wide, and usually narrow. The two sides are usually
parallel or subparallel, but in certain species the caudal end is
widened so that the scale is distinctly pyriform. The scales are
generally some shade of brown or yellow, but in some species are
opaque white. The surface of the scale is generally flat or slightly
convex externally, but may be slightly carinate. The exuvia
of the first nymph is always placed at the cephalic end so that
it projects beyond the scale for at least one-half the length of
TRIBE FIORINIINI 369
the exuvia. This exuvia retains the antennae of this stage as
minute rigid structures at its cephalic end. The caudal part of
the first exuvia is placed over the cephalic part of the second,
which in all species of this tribe serves as a puparium for enclosing
the adult female. The skin of the second nymphal female is very-
large and in most species is almost as large as the entire scale.
The wax is represented in addition to what covers the second
exuvia by a narrow peripheral band and the puparium seems to
constitute the entire scale. The puparium is fastened to the first
exuvia and is covered by a thin sheet of wax to which it adheres.
The wax of the scale is very thin, a transparent colorless layer,
and the color of the adult scale is due to the color of the exuvia
and not to the color of the secretionary part of the scale as in
most other Diaspidinae. The ventral scale like the dorsal scale
is very thin and colorless. It forms a continuous sheet of wax
completely closing the scale in some species and is interrupted
on the meson or discontinuous in others.
The exuvia of the females of the second nymphal stage or the
puparium is much larger than the body of the adult female which
it encloses. The cuticle is tough and the pygidium and its caudal
end is well developed. The body of the adult female occupies the
caudal two-thirds or one-half of the puparium, but when the female
begins to deposit eggs, which from the necessity of the case are
placed in the caudal end of the puparium, her body is forced into
the cephalic part of the puparium. The absence of genacerores
in the species forming galls would suggest that they are probably
ovoviviparous, but nothing is known as to their development.
The scale of the male is elongate, about three times as long
as the broadest part. The sides are parallel or subparallel or
sometimes they very gradually diverge caudad and near the caudal
end after reaching their greatest divergence they converge sud-
denly and form a bluntly rounded caudal end, producing a scale
similar in general form to those of many male Diaspidini. The
dorsal surface of the scale may be flat or slightly convex externally,
sometimes with a slight mesal carina or slightly tricarinate. The
scale is frequently white or opaque white and the single exposed
exuvia is yellowish. The males, as in the males of certain species
of Diaspidini, are gregarious and several may be found collected
together in a small area. They are sometimes more or less con-
cealed by a filamentous or flocculent excretion of wax or the
portion of the leaf to which they are attached may be coated with
a mass of whitish powdery wax.
370 THE COCCIDAE
A few of the species live in depressions or pockets in the
leaves of the host-plant or produce abnormal growths or galls
upon their surface. In the case of Adiscofiorinia secreta described
by Green, the enclosing gall is minute and conical. The puparium
and the body of the adult female is peculiar in form, having the
cephalic portion greatly swollen, so that it resembles the pupa of
a syrphid-fly according to Green. The galls are confined to the
upper surface of the leaf with an opening to the exterior extending
into the gall from the under surface which is always closed by the
first exuvia. The puparium rests in the gall in an erect position
with its pygidium directed toward the opening and the first exuvia.
*'The male scales are sunk in shallow depressions on the under
surface. ' '
The species are found in great part, at least so far as described
species are concerned, in Asia and Australia. Of the twenty-five
species listed in Fernald's Catalogue, nine are reported from
Australia, five from New Zealand, three from Mauritius, five from
Ceylon, one from India, two from China, three from Japan, one
from Kew Gardens, England on Howea, a palm from the Australian
region, and one species of common occurrence on many species of
plants grown under glass in Europe and America which is also
found commonly in Brazil, Australia, Ceylon, and Japan upon
plants in the open air. This species is probably also a native of
the Indian or Australian region.
The body of the adult female is usually elongate. The seg-
mentation of the thorax is usually obscure, but that of the preab-
domen is frequently distinct. The cephalic end of the body is
truncate or broadly rounded. The antennae are rudimentary and
usually do not differ in form from those of other adult female
Diaspidinae, except that they are frequently larger. They differ
in certain species as to their location on the ventral aspect, being
very distant in some and close together in others. The ventral
surface of the head is sometimes elevated horn-like between them.
The pygidium is generally distinctly separated from the pre-
abdomen, frequently somewhat five-sided. The mesal portion of
the caudal end is more or less truncate and frequently also more or
less hollowed out forming a pygidial incision. The lobes in the
generalized species are represented by three pairs, but there is
usually less than this number of pairs, frequently reduced to a
single pair, and sometimes wanting. The lateres are without
projections of any sort or with pseudolobes. The median pair of
lobes is generally the largest, they are frequently oblique with
1
TRIBE FIORINIINI 371
their margins more or less serrate. The second pair of lobes are
small and are frequently deeply incised. The plates are usually
small and inconspicuous and greatly reduced in number. The
maximum number appears to be 2, 2, 3, 3, but the usual number is
0, 1, 1 or 2, 3 or none. Pectinae, even furcapectinae, are so far as
known, always wanting. The plates of the median incisura are
very variable in their presence or absence. The oraceratubae of
the altaceratubae are usually placed singly, about four on each
side of the meson or lateral half of the pygidium. They gen-
erally open in latadentes or lobe-like projections in the incisurae.
The altaceratubae are comparatively long broad tubes in the gen-
eralized species, but Green in his figures of the pygidium of
saprosomae and similis shows the altaceratubae as slender tubes like
the clavaceratubae or lamaceratubae of other Diaspidinae. This
observation is of interest as indicating the form of the primitive
ceratubae and the way in which they are modified. The absence
of plates and the proven accuracy of Green's work leaves slight
chance for these to be lamaceratubae. The brevaceratubae are
apparently always wanting.
The vulva is usually located near or slightly cephalad of the
middle of the pygidium. The anus is located at nearly the same
level as the vulva, usually less than its own width or diameter
cephalad of the vulva. This is different from the condition found
in most Mytilaspini where the anus is typically located near the
cephalic end of the pygidium distant from the position or level
of the vulva. Although the Fioriniini have been more generally
associated with the Mytilaspini, this arrangement is much more
like that of typical Diaspidini. Genacerores are present in all the
genera but one which contains only six species. There are usually
five groups, the mesogenacerores are only rarely wanting and the
cerores of the mesogenacerores and pregenacerores or even of all
five groups are sometimes contiguous so that the pygidium appears
to bear a single inverted U-shaped group or omnogenaceroris.
The most comprehensive published account of the species of
this tribe is that of Leonard! to which reference has already been
made. The fact that the females of all the species become mature
in a puparium, if judgment may be formed of the pygidial struc-
tures found in the females of other tribes with a similar develop-
ment, leads to the conclusion that a perfect understanding of the
pygidial structures of the adult female must be secured through a
study of the pygidium of the second and first nymphal stages.
Leonardi has unfortunately figured and described only the margin
372 THE COCCIDAE
of the pygidium of the adult female and even these figures are not
every thing to be desired. The custom of Green, Newstead, and
Lindinger of figuring the pygidial fringe of one or both of the
nymphal stages is an excellent one and will need to be done for
all of the species of this tribe before an accurate classification of
the species can be made.
Leonardi recognized three new generic names : Trullifiorinia,
for which acaciae Mask, may be taken as the type, Anamefiorinia,
for which casuarinae Mask, may be taken as type, and Adisco-
fiorinia, for which secreta Green may be taken as type. Lindinger
in 1911 proposed the name Crypthemichionaspis. While he does
not describe this genus, he states Anamefiorinia Leon, and Trulli-
fiorinia Leon, are synonyms. There are three species included
under the genus Crypthemichionaspis, acacia Mask., which may be
taken as the type, lidgetti Green, and nigra Lingr. The description
of alaeodendri Charm, was not available and could not be placed.
The following synopsis, based in the main upon that of Leonardi,
will aid in the identification of the species : —
GENERA OF FIORINIINI
a. Pygidium of adult female always with genacerores,
b. Pygidum of adult female with plates and usually with one to three
pairs of lobes, rarely without lobes Fiorinia Targ.
bb. Pygidium of adult female entirely without plates.
c. Pygidium of adult female with three or five groups of gena-
cerores, if with three groups, mesogenacerores and pregena-
cerores confluent,
d. Pygidium of adult female with at least one pair of lobes;
pygidial margin with projections. Trullifiorinia Leon.
dd. Pygidium of adult female without lobes; pygidial margin
without projections, entire or serrate. Anamefiorinia Leon.
cc. Pygidium of adult female with two groups of genacerores. —
Type, Fiorinia bambusae Maskell I schna fiorinia MacG.
aa, Pygidium of adult female always without genacerores.
b. Pygidium of adult female with single median pair of lobes. — Type,
Fiorinia rubra Mask. Parafiorinia MacG.
bb. Pygidium of adult female always without lobes.
Adiscofiorinia Leon.
SPECIES OF FIORINIA
a. Pygidium always with at least one pair of lobes.
b. Pygidium with median pair of lobes distinctly separated.
c. Pygidium with pair of plates between median pair of lobes.
TRIBE FIORINIINI 373
d. Pygidium narrow, caudal end convex, not hollowed out; me-
dian pair of lobes prominent, longer than broad, sides paral-
lel, distal end bluntly rounded; plates long and slender, much
longer than lobes, arranged 2, 2, those of each lateris adja-
cent to median lobe; lateres cephalad of cephalic plates finely
serrate and without projections; setae minute, not one-half
length of plates; genacerores 3-3(3-4)3-3; mesospiracerores
obsolete. — Ceylon. Mdens Green.
dd. Pygidium comparatively broad, caudal end deeply hollowed
out; median pair of lobes widely divergent, strongly oblique,
located in concavity, mesal margins with four to six teeth,
other lobes wanting; plates arranged 2, 4, those of median
incisura short, much shorter than median lobes, those of each
lateris arranged in two groups of two each, one group near
median lobe; each lateris with angular projections; gena-
cerores 5-6(10-10)10-13. — Philippine Islands on Machilus
phantosura Ckll. & Rob.
cc. Pygidium without plates between the median pair of lobes,
d. Pygidium with five groups of genacerores combined into a
single omnagenaceroris, total number of cerores 25-29 ; median
pair of lobes small, short and narrow, deeply angularly
notched at middle of distal margin; second pair of lobes
minute, hardly produced; plates long, broad at base, much
longer than the lobes, arranged 0, 1, 6 to 7, those of each
lateris arranged 1-1-4 or 1-3-3; genacerores 5-5(7-10)10-14;
spiracerores obsolete. — Ceylon on Odina odinae Leon.
dd. Pygidium with genacerores combined into a single large
omnaceroris, total number of cerores 86-91. — India on Taxus.
odinae multipora Lindgr.
bb. Pygidium with median pair of lobes fused at proximal end.
c. Pygidium with three pairs of lobes.
d. Pygidium with each second lobe doubly incised, three lobe-
lets subequal, entire, distal end bluntly rounded, median pair
of lobes strongly oblique, inserted in deep concavity, mesal
margin of each finely serrate; third pair of lobes about as
large and similar in shape to lateral lobelet of second lobe;
plates arranged 0, 1, 1, 0, long and slender, much longer than
lobes; genacerores 7-7(13-14)17-17; mesospiracerores 2-3. —
Ceylon on Juniperus. juniperi Leon.
dd. Pygidium with each second lobe not incised.
e. Pygidium with genacerores 10-11(22-28)16-27; median pair
of lobes small, adjacent, margins strongly dentate; second
pair of lobes elongate, sometimes spatulate, distal end
truncate; third pair of lobes similar in length to the sec-
ond pair, long and narrow, distal end rounded; plates ar-
ranged 0, 1, 1, 2, plates more than twice as long as the
lobes, those of each lateris arranged, one short distance
cephalad of third lobe and one at cephalic end; second
and third incisurae with projections as large or larger than
374 THE COCCIDAE
adjacent lobes, bearing oraceratubae ; brevaceratubae in
tbree continuous series; mesospiracerores arranged in com-
pact group; antennae close together; anus located some
distance cephalad of vulva. — Java on Piper
diaspiformis Newst.
ee. Pygidium with genacerores 4-4(4-6)5-7; median pair of
lobes small; second and third pairs of lobes small; margin
of pygidium serrate and bearing several plates arranged in
pairs along serrate edge. — New Zealand on Ribes
grossulariae Mask,
cc, Pygidium with less than three pairs of lobes,
d. Pygidium with two pairs of lobes.
e. Pygidium with second pair of lobes deeply incised.
f. Head with dorsal aspect with prominent proboscis-like
projection which extends cephalad and has distal end
slightly globularly enlarged; pygidium with median pair
of lobes fused at proximal end, strongly oblique, placed
in concavity, margins serrate; second pair of lobes in-
cised, mound-like, inconspicuous, lobelets subequal and
similar; plates arranged 0, 1, 3, those of each lateris ar-
ranged, one adjacent to third lobe and two near middle;
; genacerores 6-6(13-16)21-23; mesospiracerores 1. — Cey-
lon on Gelonium prodoscidaria Green.
ff. Head never with dorsal proboscis-like projection, anten-
nae situated in median depression and adjacent,
g. Pygidium with genacerores 5-7(11-12)12-17; median
pair of lobes fused at proximal end, strongly oblique,
placed in concavity, margins serrate; second pair of
lobes incised, mesal lobelet longest and sometimes
notched on mesal and lateral margins, lateral lobelet
bluntly rounded, entire; plates arranged 0, 1, 3, those
of each lateris arranged two, longest, near a second
lobe, and one near cephalic end; mesogenacerores and
pregenacerores confluent, forming a solagenaceroris ;
mesospiracerores 4-5. — Fiorinia fioriniae minor Mask.
— America, Europe, Australia, Ceylon, China, etc. on
Camellia, Citrus, Psidium, Areca, Cycas, Anthurium,
Phytelephas, Kentia, Hedera, Leptospermum, Ficus,
Livistona, Podocarpus, Cupressus, palms, tea, ferns,
etc. fioriniae Targ.
gg. Pygidium with genacerores 4-4(16-22)24-27. — Japan on
Podocarpus, Pinus. fioriniae japonica Kwn.
ee. Pygidium with second pair of lobes not incised, distal end
bluntly rounded, faintly notched on lateral margin; me-
dian pair of lobes fused at proximal end, strongly oblique,
placed in concavity, margins serrate; plates arranged 0, 1, 0,
single plate of each second incisura much longer than
lobes; genacerores 2-2(6-9)15-15; mesospiracerores obsolete.
— Ceylon on Grevia. tumida Leon.
TRIBE FIORINIINI 375
dd. Pygidium with single pair of lobes.
e. Head with antennae located on ventral aspect near ce^
phalic margin, adjacent, cuticle between antennae pro-
longed beyond margin of body as prominent projection
with large globular end; pygidium with median pair of
lobes fused at proximal end, oblique, placed in concavity,
distal portions serrate; second pair of lobes represented
by slight incised serrate thickening of margin; plates ar-
ranged 0, 2, those of each lateris placed one cephalad of
serrate thickening representing a second lobe and the other
caudad of it; genacerores 4-5(10-13)15-18; mesospiracerores
2-3. — India on tea plant. theae Green.
ee. Head with cuticle between articulations of antennae not
prolonged into prominent projection with globular end; pygi-
dium with median pair of lobes not fused at proximal end,
not oblique, placed in slight concavity, sides parallel, slight-
ly longer than wide, distal end bluntly rounded, entire;
plates arranged 0, 2; oraceratubae of each second incisura
located in latadentes, cephalad of which is located two
plates of each lateris; lateres indented and crenulate; gen-
acerores 3-3(4-5)8-9; mesospiracerores 4. — New Caledonia
on Baeckia. neo-caledonia Ldgr.
aa. Pygidium without lobes.
b. Pygidium with genacerores 0-3(14-18)12-16; pygidial margin semi-
circular, broken by many irregularities, regular or conspicuous
lobes wanting; margin with fourteen to eighteen setae on each
side, about one-half of them short and fine, remainder long and
strong; scale of adult female with rather strong transverse striae.
— Japan on Bambusa. signata Mask.
bb. Pygidium with genacerores 5-8(5-8)5-8; pygidial margin semi-cir-
cular, regular lobes wanting, bearing numerous rather long and
thick spine-like projections, eight to ten pairs, giving pygidial mar-
gin deeply serrated appearance; dorsal surface with brevacera-
tubae ; scale or adult female not with transverse striae. — Considered
as possible species of Leucaspis by Leonard!. — Japan on Bambusa.
tenuis Mask.
SPECIES OF TRULLIFIORINIA
a. Pygidium with the mesal portion of caudal margin not deeply in-
cised with median pair of lobes situated in incision.
b. Pygidium with margin, at least adjacent to median pair of lobes,
with two or more tooth-like projections.
c. Pygidium with genacerores arranged in five distinct groups,
d. Pygidium with genacerores arranged 3-6(6-8)6-8; mesospira-
cerores wanting; median pair of lobes short, bluntly rounded
at distal end, contiguous at proximal end, sometimes faintly
notched, mesal margins divergent, mesal margins sometimes
adjacent and two lobes forming single hemispherical area,
other individuals intermediate between these two types;
plates wanting; each lateris with about three indentations
376 THE COCCIDAE
with tooth-like projections between them; anus and vulva lo-
cated at about same distance from caudal margin. — Fiorinia
acaciae bilobis Full. — Australia on Acacia acaciae Mask.
dd. Pygidium with genacerores arranged 3-3(6-6)5-6; mesospira-
cerores 0-2; median pair of lobes adjacent, entire, bluntly
pointed, hardly projecting beyond general contour of caudal
margin; plates wanting; pygidium cephalad of each median
lobe with three latadentes, bearing large oraceratubae, out-
lets of large allaceratubae ; setae large and prominent; pygid-
ium of second nymphal female with median lobes large,
mesal margins straight, adjacent, prominent lateral notch;
second pair of lobes large, deeply incised, mesal lobelet much
larger than lateral; third pair of lobes deeply incised, lobe-
lets short and broad; each second and third incisura with
long triangular plate and prominent latadentis and plate on
each lateris near third lobe. — Crypthemichionaspis nigra
Lindgr. — South Australia on Acacia, Phyllodian. nigra Lindgr.
cc. Pygidium with five groups of genacerores, mesogenacerores
and pregenacerores confluent or almost so, forming solagena-
ceroris of 40 to 50 cerores, postgenacerores consisting of 15-20
cerores; median pair of lobes short, broad, broadly rounded,
distant, with minute tooth-like projection between them; each
lateris with lobe-like projections; plates apparently wanting;
anus located in line with cephalic portions of postgenaceroris. —
New Zealand on Brachyglottis. minima Mask.
bb. Pygidium with margin adjacent to median pair of lobes finely
serrate, not with tooth-like projections; median pair of lobes nar-
row, much longer than wide, oblique, distal end obliquely trun-
cated;— Leonardi figures them as triangular, serrate, and contigu-
ous at proximal end; — plates wanting; altaceratubae long and
slender, linaceratuba-like; each lateris with indentations; vulva
on line drawn between pregenacerores and postgenacerores, anus
little farther cephalad; articulations of antennae distant; gena-
cerores 3-5(13-18)13-18; mesospiracerores obsolete; margin of body
with blunt marginal tubercles. — Ceylon. simillis Green.
aa. Pygidium with mesal part of caudal margin deeply incised, median
pair of lobes situated in incision.
b. Pygidium as broad as long, never twice as long as wide.
c. Head with antennae articulated close together and with short
cuticular projection bearing number of small spine-like points
between them.
d. Pygidium with median pair of lobes strongly oblique, proxi-
mal ends fused, mesal margins finely serrate on distal por-
tion; plates wanting; altaceratubae long and slender, lina-
ceratubae-like ; each lateris with tooth-like projections adja-
cent to median pair of lobes and undulate cephalad of them;
anus on line drawn between pregenacerores and postgena-
cerores, vulva slightly farther cephalad; genacerores 8-8(15-
29)14-19; mesospiracerores 3; margin of body with blunt
marginal tubercles. — Ceylon on Saprosma. .saprosmae Green,
TRIBE PIORINIINI 377
dd. Pygidium with median pair of lobes more prominent and dis-
tinctly serrate; second pair of lobes sometimes present. —
Ceylon on Gelonium. saprosmae gelonii Green.
CO. Head with articulations of antennae distant, never with circular
projection between articulations,
d. Pygidium with vulva situated between pregenacerores, anus
situated on line with mesogenacerores ; median pair of lobes
strongly oblique, fused at proximal ends, mesal margins ser-
rate, caudal ends pointed; plates wanting; each lateris with
rounded projections adjacent to median lobes, one or two in-
dentations cephalad of them, margin serrate; genacerores
4-4(9-12)12-12; mesogenacerores 2. — Ceylon on Murraya
ruJ)roUneata Leon,
dd. Pygidium with vulva situated between cephalic part of post-
genacerores and anus on line drawn through mesogenacerores ;
median pair of lobes small, hardly projecting, strongly oblique,
not adjacent at proximal end, mesal margins entire, caudal
ends rounded; plates wanting; each lateris with two or three
pointed projections adjacent to each median lobe and three
equidistant indentations cephalad of them, margin entire;
genacerores 4-6(7-11)7-13; mesogenacerores apparently want-
ing; living in glandular pockets in angles of veins of under
surface of leaves; scale of male longer than that of female. —
Ceylon on Gaertnera. scroMcularum Green.
bb. Pygidium greatly elongated, lateral margins long, slightly con-
vergent, subparallel, length over three times width of narrowest
part; median pair of lobes small, inconspicuous, oblique, finely ser-
rate; plates wanting, each lateris long, without projections or in-
dentations, undulate; two setae between median lobes, two adja-
cent to lateral margin of each median lobe, and three subequidis-
tant groups of 2-2-1, one dorsal and one ventral, cephalad of this
later group; genacerores present, number of mesogenacerores and
pregenacerores unknown and of postgenacerores 8-9; mesospira-
cerores obsolete. — Java on Rhaphis macroprocta Leon.
SPECIES OF ANAMEFIORINIA
,. Thorax with mesospiracerores present.
b. Thorax with mesospiracerores 18-20; pygidium without lobes,
plates, or pseudolobes, margin crenulate, apparently without setae;
anus located between pregenacerores; caudal portion of pygidium
with two transverse rows of irregular chitinized areas, four areas
in a row; genacerores 15-15(23-34)23-26,— Green shows 3-3(17-18)
8-9; — pygidium of second nymphal female with four pairs of lobe-
like projections, caudal pair largest, others successively smaller,
triangular, notched; indentations between lobes each with a single
short blunt projection, not extending beyond indentation. — Fiorinia
maskelli Brittin. — New Zealand on Plagianthus, Veronica, Pit-
tosporum. maskelli Brittin.
bb. Thorax with mesospiracerores 4-5; pygidium without lobes, plates,
pectinae, or pseudolobes, margin entire, only slightly undulate,
378 THE COCCIDAE
several setae, some distance from margin; pygidium broad, broad-
ly rounded; anus located much farther caudad than vulva; gena-
cerores 10-10(6-9)6-7. — Australia on Casuarina casuarinae Mask.
aa. Thorax with mesopsiracerores wanting; pygidium without lobes,
plates, pectinae, or pseudolobes, margin slightly undulate and finely
serrate; dorsal and ventral row of marginal setae; pygidium with
sides converging rapidly caudad, subtriangular, not broad, broadly
rounded; anus located cephalad of vulva in line with mesogena-
cerores and vulva in line with postgenacerores; genacerores 2-2(5-5)
4-4. — Australia on Acacia lidgetti Green.
SPECIES OF ISCHNAFIORINIA
Scale of adult female elongate, twenty-five or more times as long as wide,
excessively slender, sides straight and parallel, wax thin and trans-
parent, color white, appears yellow from puparium showing through;
body elongate, several times longer than broad; pygidium with two
median trifoliate lobes, sometimes apparently wanting, figured as
slender knobbed spatulate structures; pygidial margin finely serrate
with few "spiny hairs" between and near median lobes; genacerores
arranged in two groups of three cerores each. — China on Bambusa
'bam'busae Mask.
SPECIES OF PARAFIORINIA
a. Insect living free, not producing galls.
b. Pygidium with median pair of lobes adjacent, short, broad, broad-
er than long, mesal angle prominent, lateral margin broadly
roundly oblique, minutely serrate; caudal end of pygidium not
hollowed out, lobe-like angular projection at lateral margin of each
lobe; each lateris serrate with about nine "spines" on each side;
brevaceratubae apparently present. — Fiorinia rubra propinqua
Mask. — Australia on Acacia. rul)ra Mask.
bb. Pygidium with median pair of lobes adjacent only at proximal
end, strongly oblique, placed in a deep pygidial incision, distal
ends bluntly pointed, mesal margin finely serrate; each lateris
distinctly dentate, with few short thick "spines." — China, Formosa,
Queensland on Nephelium. nephelii Mask.
aa. Insects producing galls, arranged in series around margin of leaf;
pygidium of adult female with median pair of lobes projecting, ex-
ceeding in length adjacent setae; other lobes wanting; dorsal and
ventral oraceratubae wanting. — Algeria on Nephelium
hirsuta Marchal.
SPECIES OF ADISCOFIORINIA
a. Pygidium with irregular arched group of "circumgenital glands" on
ventral aspect at cephalic end adjacent to preabdomen; margin semi-
circular, without projections or indentations, with few short setae
just within margin; anus central in position; pygidium of second
nymphal female with distinct quadrangular lobes, longer than wide,
deeply notched for at least one-half their length, with four angular
adjacent lobe-like projections cephalad of each median lobe. — Egypt
on Poplar. afrieana Newst.
TRIBE FIORINIINI
379
aa. Pygidium not with transverse band of oraceratubae or cerores at
cephalic end of ventral aspect adjacent to preabdomen.
b. Pygidium short and broad, never twice as long as wide. ~ ~
c. Pygidium with caudal margin not truncate and lateral margins
straight.
d. Pygidium with anus located little cephalad of vulva; spir-
acles with spiracerores; pygidium with sides converging
caudad and caudal end transverse and indented to form four
rounded lobules; pygidial margin with band of seven or eight
setae. — Ceylon on Atalantia. atalantiae Leon.
dd. Pygidium with anus located near middle of its length and
vulva little farther caudad; spiracles encircled by minute
spiracerores; pygidium with sides converging or broadly
rounded, distinctly broader than long, margin with about
eight long slender setae arranged in three groups, group of
three on middle of each side and group of two at caudal
end, latter longer than others, and two rows on disk of
pygidium, row of six on line drawn through anus and row
of two on line between anus and margin. — England, Kew
Gardens on Howea. kewensis Newst.
cc. Pygidium with caudal margin truncate and each side straight
and not converging caudad, caudal margin coarsely crenulate;
cephalic end of body narrow, cylindrical, and truncate; margin
of pygidium with minute median depression on each side with
two or three short setae. — Fiorinia expansa Mask. — Australia
on Melaleuca. expansa Mask.
bb. Pygidium twice as long as broad, meson of caudal end with
asymmetrical lobe-like projection, sides with asymmetrical pseudo-
lobes, surface longitudinally striate; vulva located near cephalic
end of pygidium and anus located near caudal end; spiracles ele-
vated on small round tubercles; insects living in galls formed upon
upper surface of leaf, body swollen, pupa-like. — Ceylon on Grewia.
secreta Green.
CHAPTER XXVII
TRIBE ASPIDIOTINI
The Deltoid Scales
The body of the adult female is broader than long, rarely slightly
longer than broad. The cephalic end is usually broadest, broadly
rounded, and the two lateral margins converge caudad to a bluntly
rounded point. The body has a distinct triangular or deltoid outline.
The indentations in the lateral margin of the thorax and preabdomen
marking the position of the coriae may be distinct, but are frequently
obscure. The segmentation of the preabdomen is always more distinct than
that of the other parts of the body. The lateral margins of the segments
of the preabdomen are never provided with bracteae in either the
adult or second nymphal female. Lobes are usually present in the
adult female, typically three pairs, although more or less than this
number may be present, or they may be absent, or absent and repre-
sented by pseudolobes. Pectinae and plates are usually present,
pectinae may be of any type, only rarely distapectinae. The incisurae
are usually narrow and filled with pectinae or plates, sometimes small.
Densariae and paraphyses are frequently present. Altaceratubae are
never present or at least very rarely. Brevaceratubae at most only
rarely present, are never arranged in crescentic transverse rows, and
are replaced by clavaceratubae and linaceratubae. Genacerores are
frequently present, arranged in five, four, or two groups. The meso-
genacerores, when present, usually consist of only a few pores, about
six or less, frequently wanting. Scale of the adult female coriaceous
or thin and usually brownish, reddish, blackish, or grayish in color,
rarely white. Typical in form, round with the exuviae at the center,
one exuvia superimposed over the middle of the other, scale rarely
oval in outline with the exuviae slightly eccentric, but never at or
projecting beyond the margin, never pyriform or mussel-shaped, rarely
two or three times as long as broad with subparallel sides. The
female sometimes transforms in a puparium so that the scale of the
adult female may contain only the first exuvia and the pygidium is
usually provided with pseudolobes. The scale of the male is similar
in shape and color to that of the female but differs in having a single
exuvia. It is never provided with a mesal carina and is rarely chalky
white in color.
A considerable number of the injurious species of the Diaspi-
dinae belong to the tribe Aspidiotini. In fact not only some of
the most injurious but some of the best known species of coccids,
380
TRIBE ASPIDIOTINI 381
as the San Jose Scale, belong to this tribe. The general form of
the scale and of the body makes it an easily identified group and,
since there is but slight variation in their general structure, there
is but slight reason for misplacing the species of this tribe in any
systematic scheme. The species show a wide range of variation
in many of the characters peculiar to them. This is particularly
true when they are compared as to the serial modification of their
lobes, pectinae, plates, genacerores, transformation in a puparium,
and the development of pseudolobes. Because of this range of
variation, the tribe includes several very generalized species and a
number of greatly modified and highly specialized species repre-
senting a number of distinct lines of development.
The scale of the adult female is quite constant in form. It is
circular in outline and usually varies from this shape only in those
individuals that occur on narrow leaved plants or where the space
for development of the scale is limited either through the crowding
together of the insects upon the plant or the form of the plant.
In a few other species, apparently without cause, certain individ-
uals also depart from the typical circular form of the scale. The
scales of such individuals are usually short oval in outline, very
rarely twice as long as wide. They are readily differentiated from
the round or oval scales of other tribes in that the two ends of the
scale, if oval, are similar and broadly rounded, while the two
exuviae are always placed near the center of the scale, very rarely
if ever adjacent to the periphery of the scale and never projecting
over the margin. They differ also in that the first exuvia is borne
superimposed over the middle of the second and not at one end as
in the scales of the other tribes. In individuals of this tribe the
nymphs of the first stage when ready to cast their cuticle or molt,
rupture it around the periphery of the entire body and the
antennae remain attached to the ventral portion of the exuvia and
not to the dorsal portion as in the scales of the previously described
tribes. The wax of the scales is generally of considerable extent
and while ordinarily dark in color, brownish, reddish, or blackish,
they are often grayish, and only rarely are whitish or white. The
waxy covering may be either thick and opaque or thin and trans-
parent. The first exuvia is usually covered by a very thin cover-
ing of wax and in many species frequently bears a minute central
nipple-like prominence of wax, the cicatrix, which is the remains
of the first downy ball of wax formed by the nymph of the first
stage. The cicatrix is white in color and when placed upon a dark
colored exuvia is very conspicuous. The exuviae are generally
382 THE COCCIDAE
light yellowish in color, but may be blackish or brownish. The
ventral scale varies greatly in consistency from a delicate tissue-
like sheet to a rather thick pellicle. It usually remains attached
to the host-plant and is in most cases so delicate that the ventral
portion of the exuvia is generally lost. The scales are more or
less convex or obconical, the exuviae and cicatrix being placed on
the highest part of the scale. This arrangement gives the scales
quite an armored shield-like appearance from which has been de-
rived many of their scientific names as well as the common name of
the Armored Scales, applied to the subfamily as a whole. This
structure is typical, however, only of the scales of the tribe
Aspidiotini.
The scale of the male is similar in shape, appearance, and
color to that of the female. They differ from the scales of the
adult female only in having a single exuvia and their much smaller
size. The scale of the male is so similar to that of the first stage
female that it is generally quite difficult to differentiate them and
frequently this can only be done by examining the insects under
the scales. The scales of those males having females with oval
scales are frequently also oval.
The body of the adult female is located under the central
part of the scale. It is typically triangular or deltoid in shape,
broadest across the thorax or head, the exact location varying with
the species. When widest across the head, the cephalic end of the
body is frequently very broad and sometimes truncate. The body,
even when fully expanded, is rarely twice and in most species is
about one and one-half times as long as wide, but, when the seg-
ments of the preabdomen and pygidium are retracted or telescoped
as is the usual condition in the dead and dried individuals, the
body is not any longer than wide and is frequently even wider
than long. In those species that lay eggs the retraction of the
caudal end of the body makes a place for the eggs which are usually
mixed with the fine pulverulent wax excreted by the genacerores.
The antennae are typical of the form found in the subfamily.
Each consists of a small tubercle bearing a long curved seta sur-
rounded by one or more much shorter ones, usually about four.
The antennae are often greatly reduced in size or wanting. The
segmentation of the thorax is generally obscure. There are a few
genera as Selenaspidus and its near allies in which there is a deep
constriction between the mesothorax and metathorax. In fully
expanded individuals the segmentation of the preabdomen is fairly
distinct and the lateral indentations are rather prominent. The
TRIBE xVSPIDIOTINI 383
segments are for the most part crescentic in outline which permits
of their being telescoped together and the pygidium drawn into-fehe
cavity formed at their caudal end. Their lateral margins are not
or only rarely prolonged and do not bear lateral groups of bracteae
or plates as is so characteristic of many mytilaspids, diaspidids, and
parlatorids.
The preabdomen and pygidium are usually distinctly sep-
arated. The pygidium is generally provided with a pygidial
fringe that is typical in form and arrangement. The triangular
shape of the pygidium is also characteristic. The maximum num-
ber of pairs of lobes appears to be four, there are frequently only
two or three and in a number of species there is only a single pair.
Those species transforming in a puparium, as Aonidia, not only
lack lobes, but are generally provided with pseudolobes. A few
species not transforming in this way also lack all indication of a
pygidial fringe and have the pygidial margin entire. When more
than one pair of lobes is present, the lobes located cephalad of the
median pair are usually much smaller than the median pair.
These cephalic lobes vary greatly in shape and size and often
resemble the latadentes found in other tribes. The caudal margin
of the pygidium is only rarely concave or provided with a pygidial
incision with the median pair of lobes located in the incision. The
lobes are either notched or entire, rarely crenulate or serrate.
The second pair of lobes and those cephalad of them are only
rarely incised to form two or more lobelets of each lobe. The
median lobes are usually much closer together than in Lepidosaphes
or Chionaspis, frequently subadjacent, the median incisura is very
narrow, but only rarely are the lobes fused to form a single mesal
lobe.
The margin of the pygidium typically bears pectinae and
plates or both. The pectinae vary greatly in the number and
arrangement of their teeth or pectinations. The simplest type, the
distapectinae, are sometimes found cephalad of the median pair of
lobes, but probably more often between this pair of lobes where
with specialization they become reduced to bifurcate furcapectinae.
The number of species provided with latapectinae is not great,
while there are many provided with unapectinae. The pectina-
tions are usually situated on the side away from the median pair
of lobes and are sometimes long and slender, at other times are
longer than the central axis of the pectina in the unapectinae.
These pectinae are of common occurrence in the incisurae cephalad
of the median pair of lobes and upon the lateres. Furcapectinae
384 THE COCCIDAE
are of common occurrence in a number of species but only rarely
do they have more than two projections or pectinations and these
are usually distal. Some species, however, show an intergrading
between the reduced unapectinae and the furcapectinae. A single
individual may show bifurcate furcapectinae in the median incisura,
furcapectinae with three or four distal projections in the second in-
cisura, furcapectinae with three or four distal projections or those
with &\e or six distal projections in third incisura, unapectinae in
the fourth incisura, and broad mutipectinate unapectinae upon
each lateris. The plates are usually long and slender. The occurrence
of plates and furcapectinae upon the same individual is not un-
usual, the plates being confined to the median or the median and
second incisurae while the second and cephalic incisurae may
contain comparatively simple unapectinae as well as furcapectinae.
The simpler forms of pectinae, those with a small number of pec-
tinations, are found upon the caudal portion of the pygidium and
those with a considerable number on the cephalic portion. The
pygidial fringe in certain species is so reduced that not only lobes
but pectinae and plates are also wanting. The pectinae and plates
may both be wanting even when the lobes are present. The plates
of the median incisura are often greatly reduced in diameter and
length so that it is sometimes difficult to differentiate them from
the setae, which seem in such cases to be larger, but only rarely are
the plates or pectinae wanting. The plates and pectinae are quite
typical in their arrangement, usually as follows, 2, 2, 3 or 2, 3 or
2, 0 to several. The greatest variation is found upon the lateres
which lack them, bear only a few, or bear one or more groups c(m-
sisting of several.
A characteristic feature of each lateris in the pygidia of many
lepidosaphids and diaspidids is the presence of distinct indenta-
tions which are usually limited by more or less prominent projec-
tions, latadentes. These indentations are believed to mark upon
the margin of the pygidium the divisions between the segments.
There are usually one or two plates located on all or several of
these areas of the lateres in these tribes. In the aspidiotids not
only the indentations and projections are wanting but there is
no indication of a segmental arrangement of the pectinae or plates
when they occur upon the lateres. It may be found, when they
are more carefully studied, that the location of the dorsal and
ventral setae, which are associated with the dorsal and ventral
surfaces of the lobes and latadentes in the other tribes, although
not always located upon their respective lobes, will indicate some-
TRIBE ASPIDIOTINI 385
thing as to the segmentation of the pygidium of the aspidiotids.
The lateres may be entire, serrate, or crenulate, but all deep inden^
tations seem to be secondary in origin.
The ceratubae in the Aspidiotini are very different and charac-
teristic in form and generally much more numerous in number
than in the other tribes. One of the most striking features is the
absence of altaceratubae, which are so characteristic of the species
of the other groups. There may be oraceratubae located along
the pygidial margin but their ceratubae are not of the altaceratuba
type. Another striking feature is the absence of brevaceratubae.
The records are such that a definite statement can not be made,
but the presence of typical brevaceratubae in any aspidiotid is
very doubtful. The plates and pectinae are provided with lamacer-
atubae as in the other tribes, but they are often very long, some-
times two or three times as long as the pectina or plate which they
penetrate. The linaceratubae and clavaceratubae are found in
most species although there is considerable variation in number.
They are frequently fifteen to twenty times as long as wide, often
as long as the entire pygidium. Where they occur in considerable
numbers, their oraceratubae are often arranged in characteristic
longitudinal rows. The oraceratubae are usually placed in the
bottom of more or less distinct longitudinal furrows which usually
extend from or near the margin of the pygidium obliquely or longi-
tudinally across the pygidium.
The vulva is usually located near the middle of the pygidium,
on a line drawn between the pregenacerores and postgenacerores,
but may be situated some distance cephalad of this point, on a
line drawn through the cephalic part of the pregenacerores. Where
the genacerores are wanting, it can generally be identified near
the middle of the pygidium by the radiating ridges of the cuticle
which extend from its opening.
The anus is extremely variable in position. It is generally
located about half way between the middle of the pygidium and its
caudal end, but is commonly situated some distance caudad of the
vulva and is only rarely if ever superimposed over the vulva or
located cephalad of it.
Genacerores are frequently present, while the typical number
of groups, five, is often present, there is more frequently four than
five groups present. The number of groups in a few species is
reduced to two. These are apparently formed by the fusion of
the pregenacerores and postgenacerores of the same side. The
only apparent explanation of the condition found in Selenaspidus,
386 THE COCCIDAE
where only two groups are found, is that all the mesogenacerores
have been lost and that there has been a later reduction in the
number of cerores in each lateral group. This must not be taken
to imply that this is considered as the usual line of reduction
followed where the genacerores have been lost. Where there is
only a single group of genacerores on each side of the vulva, each
group may be known as a latagenaceroris. The five groups are
never fused to form a single, large, inverted, U-shaped mass of
pores and only rarely are there more than five groups present.
The number of cerores in each group of genacerores is generally not
large, the mesogenacerores varying from one to eight, the pre-
geacerores from one to forty-two, and the postgenacerores from two
to thirty. The most usual number is about two to five mesogena-
cerores and less than twelve pregenacerores and postgenacerores.
The dorsal surface of the cephalic portion of the pygidium is
sometimes provided with a series of interlacing thickened lines
forming the so-called lattice-shaped thickenings. While the caudal
portion of the pygidium on the ventral aspect may be provided
with densariae or the dorsal aspect with a few or numerous para-
physes which may be comparatively short or long and slender, the
cephalic portion of the dorsal aspect is at times provided with
distinct calles or other thickenings.
Cockerell and Leonardi have proposed and recognized several
of the dichotomous subdivisions of the genus Aspidiotus as genera
or as subgenera. On the following pages wherever possible, these
groups and names have been recognized as of the rank of genera.
The unnamed portions of these dichotomies have been recognized
as of equivalent rank and named.
The following species have all been referred to the genus Aspidiotus,
but this generic name has been omitted in each species, and, for various
reasons, mostly because of the poorness of the description, others be-
cause the description was not obtainable, have been omitted from the
tables: — alaeodendri Charm., Fiorinia (Aonidia), from Mauritius; alatus
Frogg. from New South Wales on Eucalyptus; allaudi Charm., (?Aonidia),
from Mauritius; allaudi galliformens Charm., (Aonidia), from Mauritius;
bilobis Mask, from China on grass; bossieae Mask, from Australia on
Bossiea; caldesii Targ. from Italy, west Australia on Daphne, Acacia;
coralinus Frogg., (Targionia), from New South Wales on Eremophila;
chamaeropsis Sign, from France on Chamaerops; confusus Frogg. from
New South Wales on Eucalyptus; dysoxyli Mask, from New Zealand on
Dysoxylum; eglandulosus Lindgr. from Guatemala, Panama on Cereus;
gidgei Frogg. from New South Wales on Acacia; junctilobius Frogg.
from New South Wales on Exocarpus; kennedyae Bdv. from Australia
on Kennedya; metrosideri Mask., Mytilaspis (Anoplaspis), type of
TRIBE ASPIDIOTINI
387
Anoplaspis, from New Zealand on Metrosideros ; myopori Lidg. from
Australia on Myoporum; osmanthi Vail, from France on Olea; pandnnt
Bdv. from India on Pandanus; phormii Sign, from France, Switzerland
on Phormium; rubribullata Frogg., (Aspidiella), from west Australia,
New South Wales on Eucalyptus; santali Mask., (Diaspis), from New
Zealand on Santalum, pear, plum, other fruits; serrata Frogg. from New
South Wales on Acacia; and sophorae Mask, from New Zealand on
Sophora. The various groups recognized as genera can be separated by
means of the following table: —
GENERA OF ASPIDIOTINI
Body of adult female never enclosed in puparium; pygidium of adult
female usually with lobes and pectinae or plates, never with pseudo-
lobes, never with asymmetrically arranged projections,
b. Mesothorax and metathorax never separated by deep transverse
constriction.
c. Pygidium of adult female always with genacerores.
d. Pygidium of adult female always with pectinae or plates or
with both, sometimes few in number and small.
e. Pygidium always without densariae or paraphyses.
f. Pygidium of adult female always with four or five groups
of genacerores.
g. Pygidium with four pairs of distinct lobes. — Type,
Aspidiotus subrubescens Mask. Octaspidiotus MacG.
gg- Pygidium with less than four pairs of lobes,
h. Pygidium always with three pairs of lobes.
i. Pygidium with mesal portion of caudal margin
convex, never with deep square pygidial incis-
ion.— Evaspidiotus Leon Aspidiotus Bouche.
ii. Pygidium with mesal portion of caudal margin
with deep square pygidial incision with median
pair of lobes located in incision. — Type, Aspidi-
otus excisus Green Temnaspidiotus MacG.
hh. Pygidium with less than three pairs of lobes.
i. Pygidium with two pairs of distinct lobes
Aspidiella Leon,
ii. Pygidium with single pair of lobes; second and
third pairs replaced by short, colorless, plate-
like, one or two-toothed projections. — Type,
Aspidiotus corticis-pini Lindgr
Unaspidiotus MacG.
ff. Pygidium with two groups of genacerores.
g. Pygidium with three pairs of lobes. — Type, Aspidi-
otus corokiae Mask Aspidioides MacG.
gg. Pygidium with one pair of lobes. — Type, Aspidiotus
implicatus Mask Marlattaspis MacG.
ee. Pygidium always with densariae or paraphyses.
388 THE COCCIDAE
f. Pygidium always with densariae.
g. Pygidium of adult female with three pairs of lobes.
Furcaspis Lindgr.
gg- Pygidium of adult female with less than three pairs
of lobes,
h. Pygidium with two pairs of lobes. — Type, Aspidio-
tus ostreaeformis Curtis Quadraspidiotus MacG.
hh. Pygidium with single pair of lobes
Diaspidiotus Leon,
ff. Pygidium always with paraphyses.
g. Pygidium always with four or five groups of gena-
cerores.
h. Scale of adult female circular or nearly so, exuviae
superimposed one above the other, situated at or
near middle of scale; pygidium with anus fre-
quently nearer caudal margin than postgenacerores.
Chrysomphalus Ashm.
hh. Scale of adult female elongate and narrow, two or
three times as long as broad, sides parallel or sub-
parallel, exuviae superimposed one above the other
and situated at or near one end of scale; pygidium
with anus located between postgenacerores
Pseudischnaspis Hemp.
gg. Pygidium never with more than two groups of gena-
cerores, one on each side of vulva. — Type, Melanaspis
samoana Lindgr. Lindingaspis MacG.
dd. Pygidium of adult female without pectinae or plates.
e. Pygidium always without paraphyses.
f. Pygidium with four to six groups of genacerores.
g. Pygidium always with densariae; genacerores in
four or five groups,
h. Pygidium with two pairs of lobes. — Type, Aspidi-
otus forbesi Johns For'besaspis MacG.
hh. Pygidium with single pair of lobes. — Type, Aspidi-
otus covilleae Ferris Ferrisaspis MacG.
gg- Pygidium never with densariae; genacerores in five
or six groups; projections and mesal indentation
present, lobes lacking. Comstockiella Ckll.
ff. Pygidium with three groups of genacerores or with
three groups fused into single inverted U-shaped group.
— Type, Odonaspis janeirensis Hemp—Ligulaspis MacG.
ee. Pygidium always with distinct paraphyses.
f. Pygidium with two large distinct groups of genacerores;
lateres with lobe-like prolongations, caudal end rounded,
two pairs of indentations continued as paraphyses; anus
about midway between vulva and caudal margin
Odonaspis Leon.
TRIBE ASPIDIOTINI 389
ff. Pygidium with three groups of genacerores and two
pairs of paraphyses; anus located near vulva. — Type,
Odonaspis bambusarum Ckll Berlesaspidiotus MacG,
cc. Pygidium of adult female never with genacerores.
d. Pygidium of adult female always with pectinae or plates or
both,
e. Pygidium always without densariae or paraphyses.
f. Pygidium with distinct median incisura, median pair of
lobes never fused into single lobe.
g. Pygidium with anus located near proximal end of
median pair of lobes, distinctly nearer than midway
between vulva and median pair of lobes.
h. Body never with pair of large prominent, oval,
gland-like areas on dorsal aspect near cephalic end.
i. Pygidium with at least three pairs of lobes, sec-
ond and third pairs rudimentary, median pair
not convergent at distal end.
j. Pygidium with normal pointed plates or pec-
tinae.— Type, Aspidiotus yuccae Ckll
Neosignoretia MacG.
jj. Pygidium with cylindrical plates with sub-
parallel sides and truncate ends. — Type,
Aspidiotus mauritianus Newst
Tollaspidiotus MacG.
ii. Pygidium always with less than three pairs of
lobes.
j. Pygidium with two pairs of lobes, mesal mar-
gins of median pair distant and not converg-
ent. Gryptaspidiotus Lindgr.
Jj. Pygidium with single pair of lobes, mesal
margins of median pair strongly convergent
or lobes contiguous. M organella Ckll.
hh. Body always with pair of large, prominent, oval,
gland-like areas on dorsal aspect near cephalic
end. Pseudotargionia Lindgr.
gg- Pygidium with anus located distant from proximal
ends of median pair of lobes, about midway between
vulva and median pair of lobes or nearer to vulva than
to lobes.
h. Pygidium with three or four pairs of lobes.
i. Pygidium with four pairs of lobes.
j. Plates, at least in part, twice as long as me-
dian pair of lobes, those of median and sec-
ond incisurae a single deeply divided plate. —
Type, Aspidiotus portoricensis Lindgr
Crenulaspidiotus MacG.
390
THE COCCIDAE
jj. Plates all subequal with each other and lobes,
all plates distinctly separated. — Type, Fur-
caspis proteae Brain Separaspis MacG.
11. Pygidium with three pairs of lobes, second and
third pairs distinctly lobe-like.
j. Body normal in form, slightly longer than
broad, as broad as long, or broader than long,
never four times as long as broad.
k. Pygidium with caudal margin with deep
pygidial incision; median pair of lobes,
about one-half size of second pair, placed in
incision. — Type, Aspidiotus kellyi Brain
Brainaspis MacG.
kk. Pygidium with caudal margin never with
pygidial incision; median pair of lobes as
large as or much larger than second pair.
1. Ventral aspect of head, thorax, and pre-
abdomen not with submarginal series of
large setae or short conical setae.
m. Pygidium with pectinae between lobes.
n. Insects producing galls upon plants.
Cryptophyllaspis Ckll.
nn. Insects not producing galls upon
plants. — Type, Aspidiotus fissidens
Lindgr. Spinaspidiotus MacG.
mm. Pygidium with plates between lobes;
insects not producing galls upon
plants. — Type, Aspidiotus helianthi
Parrott. RMzaspidiotus MacG.
11. Ventral aspect of head and thorax with
a submarginal series of long stout setae
and preabdomen with a band of four to
six conical setae on each side. — Type,
Aspidiotus capensis Walk
Truncaspidiotus MacG.
jj. Body abnormal in form, length four times
greatest width, head, thorax, and preabdomen
distinctly more strongly chitinized than pygi-
dium.— Type, Aspidiotus fiorineides Newst
Varicaspis MacG.
hh. Pygidium with less than three pairs of lobes.
i. Pygidium with two pairs of lobes.
j. Pygidium with median pair of lobes large and
prominent, second pair of lobes smaller but
lobe-like. — Type, Aspidiotus minimus Leon.__
Gonaspidiotus MacG.
TRIBE ASPIDIOTINI 391
I
jj. Pygidium with median pair of lobes short
and broad, broader than long, subrudimen^
tary; second pair of lobes long, spatulate,
more than twice as long as broad, notched
near distal end on lateral margin. — Type,
Aspidiotus maculata Newst
Paranewsteadia MacG.
ii. Pygidium with single pair of lobes.
j. Pygidium with lobes large, projecting promi-
nently beyond general contour of margin;
each lateris with several plates or pectinae. —
Type, Aspidiotus guterreziae Ckll
Chorizaspidiotus MacG.
jj. Pygidium with lobes not prominent, not or
hardly projecting beyond general contour of
margin; each lateris without plates or pecti-
nae.— Type, Aspidiotus chenopodii Marl
Remotaspidiotus MacG.
ff. Pygidium with median incisura obliterated by complete
fusion of median pair of lobes Chentraspis Leon.
ee. Pygidium always with densariae or paraphyses.
f. Pygidium always with densariae.
g. Pygidium with three pairs of lobes; second and third
pairs minute and usually pointed at distal end, some-
times plate-like Hemiberlesia Ckll.
gg. Pygidium with less than three pairs of lobes.
h. Pygidium with densariae of second incisurae nor-
mal in form,
i. Pygidium with two pairs of distinct lobes. —
Type, Aspidiotus perniciosus Comst
Comstockaspis MacG.
ii. Pygidium with only single pair of lobes.
j Pygidium with unapectinae; anus located at
least twice its own width from proximal end
of median pair of lobes; median pair of lobes
with mesal margins parallel and distant; in-
sects producing galls upon plants. — Type,
Aspidiotus liquidambaris Kotins
Chemnaspidiotus MacG.
jj. Pygidium with plates, if with pectinae, not
with unapectinae. — Type, Aspidiotus ulmi
Johns. Hendaspidiotus MacG,
hh. Pygidium with mesal margin of densariae of each
second incisura greatly enlarged, paraphysis-like,
three or four times as long as lateral margin,
cephalic end circular. — Type, Aspidiotus subsimilis
anonae Houser. Clavaspis MacG.
392 THE COCCIDAE
ff. Pygidium always with paraphyses.
g. Pygidium with median incisura always present and
separating median pair of lobes.
h. Pygidium with three or four pairs of lobes and
with two or more pairs of paraphyses.
i. Pygidium with four pairs of lobes and four or
more pairs of paraphyses. — Type, Aspidiotus
lilacinus Ckll. Pelomphala MacG.
ii. Pygidium usually with three pairs of lobes,
rarely with only two.
j. Body with cephalic transverse constriction
bounding broad bluntly rounded head-like
area. Mycetaspis Ckll.
jj. Body with cephalic portion uniformly con-
vex, never transversely constricted forming
head-like area,
k. Pygidium with second pair of lobes deeply
incised; pectinae long and broad, truncate
at distal end. — Type, Aspidiotus cladii Mask.
Paraonidiella MacG.
kk. Pygidium with second pair of lobes not
deeply incised; at most only notched; pec-
tinae or plates or both present, if pectinae,
not truncate at distal end.— Aonidiella Leon,
hh. Pygidium with single pair of lobes and one or more
pairs of paraphyses. — Type, Aspidiotus ceratus
Mask. Monaonidiella MacG.
gg. Pygidium with median incisura obliterated by fusion
of median pair of lobes into single lobe, other lobes
obsolete; with two pairs of paraphyses. — Type, Aspid-
iotus extensa Mask Neoleonardia MacG.
dd. Pygidium of adult female always without pectinae or plates,
e. Pygidium always with at least one pair of lobes,
f. Pygidium with three or four pairs of lobes.
g. Pygidium with four pairs of lobes and paraphyses. —
Type, Aspidiotus phyllanthi Green.— Greenoidea MacG.
gg. Pygidium with three pairs of lobes.
h. Pygidium without densariae or paraphyses; in-
sects producing galls upon plants. Maskellia Fuller,
hh. Pygidium with densariae but without paraphyses;
Insects never producing galls upon plants. — Type,
Aspidiotus yuccarum Ckll Targaspidiotus MacG.
ff. Pygidium with one or two pairs of lobes,
g. Pygidium without densariae.
h. Pygidium with two pairs of lobes. — Type, Aspidi-
otus cedri Green. Pygidiaspis MacG.
TRIBE ASPIDIOTINI 393
hh. Pygidium with single pair of lobes
Targionia Sign.
SS' Pygidium with densariae.
h. Pygidium with two pairs of lobes. — Type, Aspidi-
otus socotranus Lindgr Affirmdspis MacG.
hh. Pygidium with single pair of lobes. — Type, Aspidi-
otus campylanthi Lindgr Targionidea MacG.
ee. Pygidium without lobes, margin sometimes with angular
projections or rounded protuberances but never with even
inconspicuous lobes,
f. Pygidium without parphyses or densariae.
g. Pygidium with caudal margin entire, coarsely crenu-
late, or without projections of any sort, lateral mar-
gins strongly converging caudad, forming a right
angle.
h. Pygidium with deep mesal and lateral indenta-
tions.— Type, Aspidiotus canaliculata Green.
Circulaspis MacG.
hh. Pygidium with indentations of normal depth, not
with deep mesal and lateral indentations. — Type,
Diaspis arizonica Ckll Rugaspidiotus MacG.
gg. Pygidium with caudal margin with thirty or more
subquadrate, irregular, blunt, closely adjacent projec-
tions, lateral margins only slightly convergent
caudad, forming very wide obtuse angle. — Type,
Aspidiotus rhizophilus Newst. Ohtusaspis MacG.
ff. Pygidium with two or more pairs of paraphyses.
g. Anus and vulva located near middle of pygidium and
superimposed. — Type, Aspidiotus pimentae Newst
Leonardianna MacG.
gg. Anus located near middle of cephalic half of pygidium
and cephalad of vulva. Froggattiella Leon.
bb. Mesothorax and metathorax always separated by deep transverse
constriction.
c. Pygidium of adult female always with genacerores.
d. Pygidium with dorsal aspect always without lattice-shaped
thickenings.
e. Pygidium always without paraphyses.
f. Pygidium with four or five groups of genacerores. — Type,
Aspidiotus culculiginis Green 8tring aspidiotus MacG.
ff. Pygidium always with less than four groups of gena-
cerores.
g. Pygidium with three groups of genacerores; scale of
adult female capsulate, posterior margin elevated by
thickened ventral scale. — Type, Aspidiotus cistuloides
Green. Semelaspidus MacG.
394 THE COCCIDAE
gg. Pygidium with two groups of genacerores; scale of
adult female never capsulate Selenaspidus Ckll.
ee. Pygidium always with paraphyses. — Type, Pseudaonidea
clavigera Ckll. Duplaspidiotus MacG.
dd. Pygidium with dorsal aspect always with lattice-shaped
thickenings.
e. Pygidium with four or five groups of genacerores
Pseudaonidia Ckll.
ee. Pygidium with two groups of genacerores.
f. Pygidium without paraphyses. — Type, Pseudaonidia pae-
onia Ckll. ^ Pseudaonidiella MacG.
ff. Pygidium with paraphyses. — Type, Chrysomphalus mal'
leolus Green. Paraonidia MacG.
CO. Pygidium of adult female never with genacerores.
d. Pygidium always with pectinae or plates or both.
e. Pygidium always without paraphyses.
f. Pygidium always without lattice-shaped thickenings.
g. Head and prothorax forming area much broader than
remainder of body, head never with deep mesal incis-
ion.— Type, Pseudaonidia magna Lindgr
Entaspidiotus MacG.
gg. Head and prothorax forming area much narrower than
area formed by combination of preabdomen and re-
maining thoracic segments; head with deep mesal in-
cision. Schizaspis Ckll. & Robs.
ff. Pygidium with lattice-shaped thickenings. — Type, Pseud-
aonidia lycii Brain. Amhigaspis MacG.
ee. Pygidium always with paraphyses and lattice-shaped thick-
enings.— Type, Pseudaonidia tesserata Charm
Lattaspidiotus MacG.
dd. Pygidium always without pectinae or plates.
e. Pygidium with four pairs of lobes and without paraphyses;
segments of preabdomen distinctly constricted. — Type,
Aspidiotus artocarpi Green. Partargionia MacG.
ee. Pygidium with less than four pairs of lobes.
f. Pygidium always with three pairs of lobes and single pair
of paraphyses; preabdomen with transverse constriction
between first and second segments. — Type, Aspidiotus
moorei Green. Parrottia MacG.
ff. Pygidium with single pair of lobes and without paraphy-
ses; preabdomen not with transverse constriction be-
tween first and second segments. --Neomorgania MacG.
aa. Body of adult female always enclosed in puparium; pygidium of
adult female frequently with pseudolobes, often with asymmetrically
arranged projections, rarely with lobes and plates or pectinae.
TRIBE ASPIDIOTINI 395
b. Pygidium of adult female with two groups of genacerores and with
distinct lobes. — Type, Odonaspis schizostachyi Ckll. & Rob ^^i__r
Bakeraspis MacG.
bb. Pygidium of adult female never with genacerores.
c. Pygidium of second nymphal female with pectinae.
d. Pygidium of second nymphal female with two or three pairs
of lobes and prominent lacinate pectinae on each lateris
Greeniella Ckll.
dd. Pygidium of second nymphal female with three pairs of lobes
and without pectinae on each lateris Aonidia Targ.
cc. Pygidium of second nymphal female without pectinae.
d. Pygidium of adult female never with two deep parallel sec-
ondary incisions; scale always consisting of wax and a first
exuvia, frequently shed from old scales; adult female en-
closed in a puparium.
6. Pygidium of adult female with two pairs of small lobes;
pygidium of second nymphal female with three pairs of
lobes. Xerophilaspis Ckll.
ee. Pygidium of adult female with single, subquadrate, lobe-
like, mesal area; pygidium of second nymphal female with
single pair of lobe-like projections. — Cryptoaonidia Leon
Phaulaspis Leon,
dd. Pygidium of adult female always with two deep, parallel,
secondary incisions; scale of male and female never contain-
ing any wax; adult female and male enclosed in puparia
Ancepaspis Ferris.
SPECIES OF OCTASPIDIOTUS
a. Pygidium of adult female with median pair of lobes entire.
b. Pygidium with median pair of lobes broadly rounded, not trun-
cate, entire; second pair of lobes narrower, as long as median
pair, entire; third and fourth pairs of lobes minute, pointed, much
smaller than second pair; pectinae long and slender, much longer
than lobes, unapectinae or latapectinae, arranged 1, 1, 1 or 2, 1
or 2, 2; genacerores (15-15)9-10. — New Zealand on Atherosperma...
atherospermae Mask,
bb. Pygidium with median pair of lobes truncate or slightly emargi-
nate, not notched at distal end.
c. Pygidium with pectinae longer than lobes; median pair of lobes
truncate or slightly emarginate at distal end, not notched; sec-
ond and third pairs of lobes narrower, rounded at distal end,
each with lateral notch; fourth pair of lobes small, distal end
acute; pectinae between lobes broad, deeply lacinate, those of
each lateris long, slender, few lacinations, arranged 2, 2, 3, 3,
4 to 5; genacerores (16-18)10-12. — Australia on Banksia, Eu-
calyptus, Pittosporum subrubescens Mask.
cc. Pygidium with pectinae subequal or shorter than the lobes,
deeply lacinate; adult female larger, more circular, pygidium
396 THE COCCIDAE
proportionately broader than in typical form; genacerores (17-
27)6-10. — ^Australia on Eucalyptus
suhruhescens corticoides Green,
aa. Pygidium with median pair of lobes with prominent mesal and
lateral notches, distal end rounded and prolonged between notches;
second, third, and fourth pairs of lobes minute, bluntly pointed,
spine-like, second pair largest; furcapectinae slightly longer than
lobes, arranged 0, 1-2, 2, 2-3, 3; spiracerores wanting; genacerores
(2-4)1-3. — South Africa on Acacia. furcillae Brain.
SPECIES OF ASPIDIOTUS
a. Pygidium with median pair of lobes shorter than either of other
pairs, not projecting nearly as far caudad, short and narrow, each
with lateral notch; second and third pairs of lobes narrow, bluntly
rounded, each with lateral notch; pectinae longer than lobes, una-
pectinae, those of each lateris broad, arranged 2, 2, 3, 6 to 7; gena-
cerores (6-16)3-9; anus distant from caudal margin three times its
own diameter; linaceratubae variable in length, with exception of
few mesal ones, eight or more times as long as broad. — West Indies,
Mexico, C^ina, Formosa, India, Laccadive Islands, Ceylon, Bourbon
Islands, Mauritius on Celtis, Terminalia, Alcocasia, Psychotria,
Manihot, Loranthus, Moesa, tea, pepper, palms, mango, banana, nut-
meg, etc. destructor Sign.
aa. Pygidium with median pair of lobes at least as long as second pair,
usually distinctly longer, at least projecting farther caudad.
b. Pygidium with lobes spatulate, constricted at proximal end,
widened at middle, narrowed at distal end, subequal in size, entire;
pectinae located between lobes and along each lateris for almost its
entire length, arranged 1, 1, 2, 7 or more; genacerores (17-21)8-13. —
Australia on Eugenia. virescens Mask.
bb. Pygidium with lobes not spatulate, wider at proximal than at distal
end.
c. Pygidium with third pair of lobes with margins entire,
d. Pygidium with third pair of lobes not deeply incised.
e. Pygidium with prominent, truncate, serrate, lobe-like pro-
jection near middle of each lateris and with seta near its
lateral margin; median pair of lobes rounded, prominent,
slight lateral notch; second and third pairs of lobes
smaller, lateral margins oblique, lateral margin of third
pair serrate; pectinae subequal in length to lobes, arranged
1, 2, 3, 3, caudal pectina of each lateris broader than
others; genacerores 3-5(8-8)2-5. — California on Pinus
californicus Colem.
ee. Pygidium not with prominent lobe-like projection near
middle of each lateris.
f. Pygidium with pectinae on each lateris.
g. Pygidium with median pair of lobes projecting dis-
tinctly farther caudad than second pair.
ft
TRIBE ASPIDIOTINl 397
Pygidium with pectinae or plates in median in-
cisura. — -
i. Pygidium with two pectinae in median incisura.
j. Pygidium with genacerores (6-14)5-11.
k. Pygidium with median pair of lobes dis-
tinctly longer than broad.
I. Pygidium with each lateris with three
pectinae; median pair of lobes long and
broad, broadly rounded, mesal and lateral
notches, latter largest; second pair of
lobes large, blunt, rounded, lateral notch;
third pair of lobes small, triangular;
pectinae of third incisurae longer than
lobes, unapectinae, others subequal, ar-
ranged 2, 2, 3, 3, those of each lateris
narrower ; genacerores ( 6-8 ) 5-5. — New
Mexico on Chilopsis coloratus Ckll.
II. Pygidium with each lateris with seven
or eight pectinae; median pair of lobes
elongate, distinctly longer than broad,
distinct mesal and lateral notches, distal
end bluntly pointed; second pair of lobes
smaller, longer than broad, distinct lat-
eral notch, distal end bluntly pointed;
third pair of lobes subequal to second;
triangular, entire; pectinae deeply laci-
nate, arranged 2, 2, 3, 7-8, those of third
incisurae and lateres unapectinae; anus
distant from caudal margin four times
its width; genacerores (12-14)9-11. —
Dahomey, Africa on Elaeis
elaeidis Marchal.
kk. Pygidium of adult female with median pair
of lobes as broad as or broader than long.
1. Pygidium with only a single pectina
cephalad of each third lobe; median pair
of lobes broader than long, quadrangu-
lar, deep lateral and faint mesal notches,
distal margin rounded; second pair of
lobes broadly rounded, broader than long,
distinct lateral notch; third pair of lobes
minute, bluntly rounded, entire; setae
large, longer than lobes; pectinae sub-
equal in length or slightly longer than
median lobes, arranged 2, 2, 2, 1; lateres
distinctly crenulate; genacerores (6-6)5-6.
— Specimens from Madagascar have lobes
longer, entire, longer pectinae, and gena-
cerores (2-4)4-8. — German East Africa,
Madagascar on Cocos varians Lindgr.
398 THE COCCIDAE
11. Pygidium with four to six pectinae
cephalad of each third lobe; median pair
of lobes large, prominent, obscure mesal
and lateral notches, distal margin blunt-
ly produced between them; second and
third pairs of lobes small, triangular,
pointed; pectinae slightly longer than
the lobes, arranged 2, 2, 3, 4-6; anus dis-
tant from caudal margin four times its
width; oraceratubae conspicuous, ar-
ranged in longitudinal rows; genacerores
1-3(10-14)5-7.— Australia on Ribes, Ampe-
lopsis, Eucalyptus, Acacia, Cytisus
tasmaniae Green,
jj. Pygidium with genacerores (3-4)2-3, said to
be in five groups; median pair of lobes broad
and stout; second pair of lobes dentiform,
lateral margin strongly oblique; third pair of
lobes reduced to tooth of margin; setae long
and stout, longer than plates and pectinae. —
Corsica on Stachys, Teucrium
laMatarum Marchal.
ii. Pygidium with two plates in median incisura;
median pair of lobes long, broad, quadrangular,
broadly rounded, mesal and lateral notches
prominent, second pair of lobes longer than
broad, bluntly rounded, lateral and possibly me-
dian notch; third pair of lobes elongate, half as
long as second pair, triangular; plates and pec-
tinae arranged 2, 2, 2, 5-7, plates of median in-
cisura as long as median lobes, others broadly
deeply lacinate distapectinae; setae very long
and stout, particularly those of second and third
pairs of lobes; genacerores (3-6)3-6. — District of
Columbia, England on Camellia, Arenga
spinosus Comst.
hh. Pygidium without pectinae or plates in median
incisura; median pair of lobes large, stout, promi-
nent, irregularly and obscurely excised; second and
third pairs of lobes minute, proximal portion
broad and distal portion aciculate; pectinae stout,
varying from deeply furcate furcapectinae to
plates, arranged 0, 2, 2, 6-9; setae long, stout,
longer than pectinae; spiracerores wanting; gena-
cerores (8-11)3-6; anus located slightly nearer
caudal margin than a line drawn through postgen-
acerores. — Java on Erythrina pustulans Green.
gg. Pygidium with median pair of lobes projecting but
little if any farther caudad than second pair, broad,
distal ends truncately rounded, faint mesal and lat-
TRIBE ASPIDIOTINI 399
eral notches; second pair of lobes large, broad,
smaller than median, distal margin undulate; third"
pair of lobes similar to second in form, smaller, mar-
gin undulate; pectinae short, those of each third in-
cisura longer than lobes, other pectinae distapectinae,
arranged 2, 2, 3, 3; genacerores 0-3(8-12)6-9. — Europe,
Maine, Massachusetts, New York, New Jersey, Georgia
on hemlock, pine, fir, maple abietis Schr.
fC. Pygidium without pectinae on each lateris.
g. Pygidium with median pair of lobes entire, large,
rounded; second pair of lobes small, similar in shape
to median; third pair of lobes minute, rounded, en-
tire; pectinae subequal in length with median lobes,
arranged 2, 2, 1, 0, those of third incisurae broadest;
each third incisura with latadentls-like projection;
genacerores (3-7)3-7. — California on Pinus
florenciae Colem.
gg. Pygidium with median pair of lobes with prominent
lateral notch, sometimes faint mesal notch, subquad-
rangular, wide, distal end truncately rounded; second
and third pairs of lobes small, projecting, angular,
long; setae stout, projecting beyond median lobes;
pectinae narrow, subequal in length with median
lobes, arranged 2, 2, 2, 1-2, each lateris irregular in
outline with one or two minute plates placed distant
from each other and third lobe; genacerores (2-6)2-4;
anus three times its own width from caudal margin.
— Italy, Greece on Hypericum, Thymelaea
privignus Lindgr.
dd. Pygidium with third pair of lobes deeply incised.
e. Pygidium with only third pair of lobes incised; third pair
of lobes consisting of two adjacent conical lobelets; median
pair of lobes large, triangular, margins serrate, distant;
second pair of lobes small, triangular, adjacent to median
lobe, appearing as deeply incised lateral lobelet; median
and second incisurae without pectinae or plates, each third
incisura with long furcapectina, each lateris adjacent to
third lobe with another similar furcapectina and few
plates on cephalic portion; genacerores (6-10)15-15. — Fiji
Islands on "various forest trees." vitiensis Mask.
ee. Pygidium with second and third pairs of lobes deeply in-
cised.
f. Pygidium with three pairs of lobes; scale of adult fe-
male oval, larger near one end, exuviae placed on smaller
end; median pair of lobes moderately large, distant;
lateres thick and serrate; two plates between median
pair of lobes and on margin between other lobes; setae
short; dorsal oraceratubae numerous; genacerores (25-
400 THE COCCIDAE
39)31-35; ordinarily with one to five cerores between
anterior groups. — Brazil on Drymus. — See next species.
pisai Hemp,
ff. Pygidium with four pairs of lobes; scale of adult female
more or less circular, flattened, exuviae placed little to
one side of center; median pair of lobes smallest; sec-
ond, third, and fourth pairs deeply incised, mesal lobelet
of second and third pairs largest, lobelets of third pair
subequal; plates and pectinae arranged 2, 1, 1, 1, 12,
median incisura with plates, other incisurae with furca-
pectinae, and lateres with plates grouped near fourth
lobe; genacerores 16-18(16-30)16-21; also two to three
glands at sides of pregenacerores. — Brazil on Drymus. —
This species and the preceeding one do not appear like
species of this tribe moreirai Hemp.
CO. Pygidium with third pair of lobes with margins notched.
d. Pygidium with four or five groups of genacerores, if with
four groups, number of cerores in groups very dissimilar,
e. Pygidium with six or more pectinae on each lateris.
f. Pygidium with three pectinae in each third incisura.
£:. Pygidium with second and third pairs of lobes dis-
tinctly broader than long; median pair of lobes pro-
jecting, bluntly rounded, faint lateral notch; second
and third pairs of lobes much shorter than median,
lateral margin strongly oblique, finely serrate; pec-
tinae subequal in length to lobes, arranged 2, 2, 3, 7,
those of each lateris unapectinae; genacerores 4-5(7-
8)4-5. — Japan on Cry ptomeria,.-. cry ptomeriae Kuwn.
SS- Pygidium with second and third pairs of lobes dis-
tinctly longer than broad.
h. Pygidium with anus distant from caudal margin
about four times its width.
i. Pygidium with two pectinae in median incisura.
J. Pygidium with longest linaceratubae not
more than four times as long as wide; me-
dian pair of lobes round, large lateral and
smaller mesal notches; second and third pairs
of lobes variable in form, second pair round-
ed and notched, third pair bluntly pointed,
faintly notched; pectinae longer than lobes,
arranged 2. 2, 3, 6, those of median and sec-
ond incisurae and caudal pectina of third in-
cisurae distapectinae, others unapectinae ; gen-
acerores ( 8-15 ) 4-10. — Aspididiotus hederae
simplex Charm. — Canada, United States, Mex-
ico, West Indies, Southern Europe, Africa,
Australia, etc. on orange, lemon, olive, palms,
orchids, cherry, plum, currant, maple, and
many other plants hederae Vail.
TRIBE ASPIDIOTINI 401
jj. Pygidium with linaceratubae eight to ten
times as long as wide.
k. Pygidium with genacerores (7-9)8-15; ine^
dian pair of lobes rounded with large later-
al and feeble mesal notches; second pair of
lobes narrower and almost entire; third
pair of lobes smaller and notched on outer
side; median lobes extend farther caudad
than second pair; pectinae arranged 2, 2,
3, 6, those of each lateris about twice as
long as a third lobe; anus slightly nearer
postgenacerores than caudal margin. — ^Aus-
tralia on palm simillimus Ckll.
kk. Pygidium with genacerores (6-11)4-6; me-
dian pair of lobes largest, mesal and lateral
notches not prominent, distal end rounded
between notches, subequal in length to
second pair; second and third pairs slender,
longer than broad, constricted on proximal
portion, distinct lateral notch; pectinae
subequal in length or slightly longer than
lobes, arranged 2, 2, 3, 6-7, those of each
lateris unapectinae; spiracerores wanting;
caudal portion of lateres toothed, cephalic
portion entire; linaceratubae long, distinct,
opening at margin. — Aspidiotus simillimus
translucens Fern. — Ceylon, Hawaiian Is-
lands on Loranthus, Dahlbergia
transparens Green.
11. Pygidium with single pectina in median incis-
ura; median pair of lobes long and projecting,
subquadrangular, longer than broad, deep lateral
notch, distal portions rounded, converging
caudad, almost contiguous; second pair of lobes
small, longer than broad, deep lateral notch;
third pair of lobes smaller than second, similar
in form, deeply notched ; pectinae of median and
second incisurae narrow, distapectinae, others
unapectinae, arranged 1, 2, 3, 6; anus over four
times its width from caudal margin; gena-
cerores (7-9)6-8 — Philippine Islands on Corpyha.
coryphae Ckll. & Rob.
hh. Pygidium with anus distant from caudal margin
six or seven times its width.
i. Pygidium with genacerores (0-2)0-2; median
pair of lobes elongate, longer than broad, distal
end rounded, distinct lateral and faint mesal
notches; second pair of lobes similar in shape,
notches smaller; third pair of lobes longer than
broad, one or two lateral notches, bluntl-
402
THE COCCIDAE
rounded; pectinae subequal in length to mediati
pair of lobes, deeply lacinate, arranged 2, 2, 3,
6-8, those of each lateris near third lobe, una-
pectinae; setae not as long as lobes; dorsal
oraceratubae numerous on each side. — Canary-
Islands on Dracaena, Gymnosporia, Ilex, Oreo-
daphne, Picconia, Smilax, Hedera, Apollonias,
Heberdenia, Laurus lauretorum Lindgr.
ii. Pygidium with genacerores 0-3(4-14)4-13.
j. Pygidium with genacerores (11-14)11-13; me-
dian pair of lobes longer than broad, broadly
rounded, faint mesal and lateral notches;
second and third pairs of lobes subequal,
slightly smaller than median, similar in
form, distinct mesal and lateral notches;
pectinae broad, distapectinae, longer than
lobes, arranged 2, 2, 3, 8-9; each lateris finely
serrate cephalad of pectinae; dorsal oracera-
tubae numerous, irregularly arranged. — Aus-
tralia on Eugenia fimhriatus Mask.
jj. Pygidium with genacerores 0-3(5-10)4-10; pec-
tinae deeply lacinate, longer than lobes, ar-
ranged 2, 2, 3, 8-12, those of incisurae dista-
pectinae, others unapectinae. — South Africa
on Cycads, Encephalartos
ftmhriatus capensis Newst.
ff. Pygidium with two pectinae in each third incisura; me-
dian pair of lobes broad, broadly rounded, mesal and
lateral notches; second pair of lobes similar, smaller,
mesal and lateral notches; third pair of lobes sometimes
irregular, smaller than second, usually with lateral mar-
gin only or with distal margin notched; pectinae short,
arranged 2, 2, 2, 6, those of incisurae bifurcate or tri-
furcate, those of each lateris slightly lacinate unapecti-
nae; genacerores 0-3(7-10)7-8. — England, Massachusetts,
Oregon on Ruscus, holly lyritannicus Newst.
Pygidium with four pectinae on each lateris.
f. Pygidium with median pair of lobes quadrangular, broad-
er than long.
g. Pygidium with median pair of lobes entire, broad,
quadrangular, broadly truncated; second and third
pairs of lobes quite small, much shorter than pectinae,
bluntly rounded, entire; pectinae deeply lacinate, ar-
ranged 2, 2, 3, 4, those of each lateris longest; dorsal
oraceratubae not numerous; anus distant from caudal
margin six to seven times its width; genacerores (6-8)
5-7. — Mexico on ash. vagabundus Ckll.
SS- Pygidium with median pair of lobes with distinct
mesal and lateral notches, distal end rounded; second
I
TRIBE ASPIDIOTINI 403
pair of lobes smaller, similar in shape, distinct mesal
and lateral notches; third pair of lobes about one-
half size of second, similar in shape; pectinae broad,-
arranged 2, 2, 3, 3-4, those of incisurae subequal to
lobes, those of lateres much longer than lobes; gena-
cerores (2-5)3-4. — South Africa on aloe. -.regriws Brain,
ff. Pygidium with median pair of lobes elongate, much long-
er than broad, mesal margin entire or with one faint
notch and lateral margin with two to four notches; sec-
ond and third pairs of lobes large, nearly as large as
median, elongate, similar in form, similar notches; pec-
tinae deeply lacinate, much longer than lobes, arranged
2, 2, 3, 4, those of each lateris subequal in length, more
irregular than others; anus distant from caudal margin
seven or eight times its own width; genacerores (2-4)
2-4, — Canary Islands on Gymnosporia
gymnosporiae Lindgr.
dd. Pygidium of adult female with four groups of genacerores,
(6-6)6-6; median pair of lobes close together, parallel, not
quite touching, considerably longer than broad, rounded, ends
minutely serrate; second pair of lobes similar, smaller, ends
serrate, their ends extending to line drawn through caudal
margin of median lobes; third pair of lobes small, lateral
notch; pectinae serrate and branched, seven on each lateris;
, setae large, not longer than pectinae; anus small, close to
proximal end of median pair of lobes. — Hawaiian Islands,
Japan on Persea, Trachycarpus, Magnolia perseai'um Ckll.
SPECIES OF TEMNASPIDIOTUS
Pygidium with median pair of lobes large, distal margins oblique, slightly
emarginate; second pair of lobes long and slender, projecting beyond
median pair, less than one-half width of median pair, shorter, narrow,
pointed; third pair of lobes similar to second pair, shorter and blunter;
pectinae deeply lacinate, unapectinae, arranged 2, 2, 3, 4; spiracerores
wanting; anus three times its own width from vulva and five times from
caudal margin; genacerores (8-15)7-9. — Ceylon on Cyanotis
excisus Green.
SPECIES OF ASPIDIELLA
a. Pygidium with either pectinae or plates in median incisura.
b. Pygidium with pectinae in median incisura.
c. Pygidium with pectinae in each second incisura.
d. Pygidium with median pair of lobes with mesal and lateral
notches.
e. Pygidium with second pair of lobes entire.
f. Pygidium with anus not more than twice its own width
from caudal margin; median pair of lobes quadrangular,
distinct mesal and lateral notches; second pair of lobes
small, longer than wide, entire, bluntly rounded; pecti-
nae long and slender, subequal in length or slightly
404 THE COCCIDAE
longer than lobes, two to five teeth, arranged 2, 2, 6-10,
caudal three or four of each lateris unapectinae, cephalic
two to four plates; dorsal oraceratubae numerous; gena-
cerores (5-7)3-6; setae large and prominent, projecting
beyond lobes. — Florida on saw palmetto
pseudospinosa Woglum.
ff. Pygidium with anus five or more times its own width
from caudal margin; median pair of lobes quadrangular,
distinct mesal and lateral notches; second pair of lobes
long and narrow, longer than broad, entire, bluntly
rounded; pectinae long, subequal in length to lobes,
those of median incisura narrow, arranged 2, 2, 6-8, three
caudal pectinae of each lateris longest and broadest,
those cephalad of these much shorter, varying from
furcapectinae to plates; dorsal oraceratubae numerous;
genacerores (9-12)6-8; setae small, not projecting beyond
lobes. — New Jersey on Oxycoccus oxycoccus Woglum.
ee. Pygidium with second pair of lobes with notch.
f. Pygidium with seven or eight pectinae on each lateris;
median pair of lobes large, broadly rounded, proximal
portion constricted, distinct mesal and lateral notches;
second pair of lobes strongly constricted on proximal
portion, distal portion broadly rounded, lateral shoulder
and lateral notch; with shoulder-like thickening, may
represent third lobe; pectinae subequal in length to
lobes, deeply toothed, arranged 2, 2, 7-8, those of each
lateris adjacent to second lobe; dorsal oraceratubae
prominent; genacerores 2-4(3-3)3-4; mesal calles oblique,
not so distinct as oblique lateral calles; anus about
twice its own width from caudal margin. — China on
Abies. meyeri Marl.
ft. Pygidium with three pectinae on each lateris; median
pair of lobes short and broad, quadrangular, prominent
lateral and mesal notches, distal margin bluntly round-
ed ; second pair of lobes broader than long, oblique, feebly
crenulate or notched; pectinae subequal in length to
median lobes, narrow, deeply lacinate, arranged 2, 3, 3,
those of each lateris adjacent to third lobe; anus oval,
near median lobes; genacerores 4-4(7-7)3-4. — California
on Arctostaphylos arctostaphyli Ckll. & Rob.
dd. Pygidium with median pair of lobes entire or with lateral
notch only.
e. Pygidium with median pair of lobes about as broad as long,
broadly rounded, mesal margins divergent, entire or with
indistinct lateral notch; second pair of lobes much smaller,
subminute, triangular; pectinae with few lacinations, ar-
ranged 2, 2, 7, caudal pectinae distinctly lacinate, cephalic
two or three modified into plates; lateres serrate cephalad
of plates; genacerores 3-4(8-9)5-7. — Aspidiotus hartii luntii
Ckll. — West Indies on yam. hartii Ckll.
TRIBE ASPIDIOTINI 405
ee. Pygidium with median pair of lobes longer than broad,
bluntly rounded, distinct lateral notch; second pair of lobes_
similar in form, one-half size first pair, lateral notch broad;
pectinae subequal in length to lobes, arranged 2, 2, 3, those
of each lateris largest; dorsal oraceratubae prominent;
anus distant four or five times its width from caudal mar-
gin; genacerores 0-3(5-6)3-5. — California on Quercus,
Pasania. densiflorae Bremner.
cc. Pygidium with plates in each second incisura; median pair of
lobes large, projecting, rounded, mesal margins parallel, promi-
nent lateral notch; second pair of lobes much smaller, about as
broad as long, distal margin sinuate; two minute furcapectinae
in median incisura, shorter than or subequal to lobes, two
plates in second incisurae, and three pectinae on each lateris;
lateres cephalad of pectinae denticulate; genacerores (7-9)7-9. —
West Indies on Saccharum. sacchari Ckll.
bb. Pygidium with plates in median incisura.
c. Genacerores (5-5)5-5; median pair of lobes large, prominent,
distant, rounded, lateral notch; second pair of lobes small,
rounded; pectinae small, poorly developed, similar to plates, ar-
ranged as in hederae, thus 2, 2, 3, 6; genacerores (5-5)5-5. —
Chili. latastei Ckll.
cc. Genacerores 0-1(8-12)6-9; median pair of lobes large; second
pair of lobes much smaller; plates present between the lobes;
dorsal oraceratubae numerous, arranged in four rows on each
side, extending from margin, those of same row varying greatly
in length. — Spain on walnut. juglandis Colvee.
aa. Pygidium without pectinae or plates in median incisura.
b. Pygidium with second pair of lobes longer than median, project-
ing farther caudad, rounded at distal end, entire; median pair of
lobes longer than broad, faint lateral and mesal notches, bluntly
rounded; pectinae broad, deeply lacinate, arranged 0, 1, 4, caudal
pectina of each lateris very broad, others narrow; anal opening
about twice length of median lobes from caudal margin; gena-
cerores (5-6)4-4. — New York, Ohio, Illinois, Iowa on maple
comstocki Johns,
bb. Pygidium with median pair of lobes extending much farther
caudad than second pair, quadrangular, mesal and lateral notches,
adjacent; second pair of lobes broad, distal end oblique, sub-
truncate, slightly emarginate; pectinae longer than lobes, ar-
ranged 0, 2, 2; genacerores (3-4)2-3. — Jamaica, Dominica, New
York, Iowa on cocoanut, pomegranate punicae Ckll.
SPECIES OF UNASPIDIOTUS
Pygidium with median pair of lobes dark yellowish brown, subadjacent,
feebly convergent, projecting far, no mesal notch, single lateral notch;
other lobes replaced by short, colorless, plate-like, one or two pointed
processes; plates and pectinae numerous, colorless, undivided, blunt,
last three frequently somewhat thickened, those near median pair of
406 THE COCCIDAE
lobes more or less deeply forked; genacerores 0-4(7-10)7-8. — Japan on
Finus. corticis-pini Lindgr.
SPECIES OF ASPIDOIDES
a. Pygidium with median pair of lobes entire.
b. Pygidium with median pair of lobes large, long and broad, quad-
rangular, distal margin bisinuate; second pair of lobes much
smaller, elongate, bluntly pointed, slight mesal notch; third pair
of lobes triangular, as broad as long, entire; pectinae longer than
lobes, deeply lacinate, mostly unapectinae, arranged 2, 2, 3, 4,
those of each lateris in two groups; genacerores in two groups of
two pores each; anus located near margin of pygidium. — New Zea-
land on Corokia. corokiae Mask.
bb. Pygidium with median pair of lobes rectangular, entire, distal end
oblique; second and third pairs of lobes bluntly rounded, contrict-
ed at proximal end, with an outer shoulder, entire; pectinae nor-
mal in form, subequal in length to lobes, arranged 2, 2, 3, 2-4, those
of each lateris adjacent to third lobe; anus large, located near
middle of pygidium; genacerores represented only by two groups
of 1-4 cerores, probably represent pregenacerores ; dorsal oracera-
tubae numerous. — Australia on Hake comperei Marl.
aa. Pygidium with median pair of lobes with mesal and lateral notches,
quadrangular, distal end broadly rounded; second pair of lobes
smaller, similar, unsymmetrical with short oblique margin; third
pair of lobes smaller than second, somewhat pointed; pectinae of
median and second incisurae as long as median lobes, furcapectinae,
others deeply lacinate, those of each lateris longest, cephalic four
often short, arranged 2, 2, 3, 4-5; genacerores arranged in two groups
of 4-6 cerores. — Canary Islands on Olea tafiranus Lindgr.
SPECIES OF MARLATTASPIS
Pygidium with median pair of lobes sloping inwards, outer margin of
each with conspicuous notch and smaller notch on rounded lateral
margin at distal end; each lateris with two deep indentations cephalad
of lobes, and several pectinae, some are simple, plate-like and others are
broad and conspicuously forked; genacerores one or two on each side
and sometimes with a single mesogenaceroris. — China on Campanula
implicata Mask.
SPECIES OF FURCASPIS
a. Pygidium with median and second pairs of lobes subequal.
b. Pygidium with each lateris produced near middle of its length
into prominent tooth-like projection with single furcapectina
cephalad of it; median pair of lobes quadrangular, broadly round-
ed, subtruncate, entire; second pair of lobes similar in size and
shape to median; third pair of lobes similar in shape to second,
smaller; pectinae subequal in length to lobes, furcapectinae, ar-
ranged 2, 2, 3, 1, those of each lateris distant from third lobe;
second and third incisurae with densariae; genacerores (4-5)4-4.
— Chrysomphalus biformis Ckll., Aspidiotus biformis odontoglossi
TRIBE ASPIDIOTINI 40T
Ckll., Chrysomphalus odontoglossi Ckll., Chrysomphalus biformis
cattleyae Ckll. — Central America, Jamaica, Trinidad, Grenada,"
Antigua on orchids. Mformis Ckll.
bb. Pygidium with each lateris not produced into a tooth-like pro-
jection near middle of its length.
c. Pygidium with five to six pectinae on each lateris near a third
lobe, unapectinae, subequal in length to lobes, arranged 2, 2, 3,
5-6; median pair of lobes subquadrangular, usually with mesal
notch, lateral margin oblique, minutely serrate; second pair
of lobes subequal in size, lateral margin oblique, finely serru-
late; third pair of lobes slightly smaller than second pair, lat-
eral margin oblique, finely serrulate; anus located midway be-
tween vulva and caudal margin; genacerores (11-14)7-9. — Japan
on Quercus. jordani Kuwn.
cc. Pygidium without pectinae on each lateris, pectinae broad, sub-
equal to lobes in length, arranged 2, 2, 3, 0; median pair of
lobes subquadrangular, truncately rounded, entire; second pair
of lobes similar in form and subequal in size; third pair of
lobes bluntly pointed, smaller than second, lateral margin
oblique, serrate; anus distant from caudal margin over twice
its own width; densariae in median, second, and third incisurae,
those of second longest; genacerores 2-4(5-7)4-5; dorsal ora-
ceratubae numerous. — Japan on Tsuga tsugae Marl.
L. Pygidium with median pair of lobes much larger than second pair,
b. Pygidium with five or more pectinae on each lateris.
c. Pygidium with pectinae of median incisura distinctly longer
than median pair of lobes; median pair of lobes quadrangular,
subtruncate, deep mesal and lateral notches; second pair of
lobes narrow, pointed, nearly as long as median pair, not one-
half as wide; third pair of lobes much shorter than second
pair, narrow, bluntly pointed; pectinae deeply lacinate, ar-
ranged 2, 2, 3, 5, apparently all latapectinae ; genacerores (4-6)
5-6. — West Indies on cocoanut palm, banana palmae Morg.
cc. Pygidium with pectinae of median incisura but little if any
longer than median lobes; median pair of lobes quadrangular,
rounded, distinct mesal and lateral notches; second pair of
lobes much shorter and about one-third width of median, blunt-
ly pointed, sometimes faintly notched; third pair of lobes
minute, pointed; pectinae deeply lacinate, arranged 2, 2, 3, 3,
those of each lateris deeply divided, strongly furcate, those of
median and second incisura f urcapectinae ; genacerores (3-5)
3-5. — Massachusetts, District of Columbia, Ohio, Mexico, Brazil,
Venezuela, England, France, Ceylon, Mauritius on Cyanophyl-
lum, Ficus, Laurus, Cycas, Ipomoea, Pritchardia, Cinchona,
palms, orchids, tea. cyanophylli Sign.
bb. Pygidium never with more than three pectinae on each lateris.
c. Pygidium with pectinae or plates on each lateris.
d. Pygidium with two or three pectinae on each lateris.
408 THE COCCIDAE
e. Pygidium without densariae between median pair of lobes;
two or three pectinae in each third incisura; median pair
of lobes quadrangular, rounded, distinct mesal and lateral
notches; second pair of lobes smaller, rounded, narrower,
mesal and lateral notches; third pair of lobes smaller,
bluntly rounded, lateral notch; pectinae deeply lacinate,
arranged 2, 2, 2-3, 2-4, those of each lateris latapectinae,
those of median and second incisurae as long as lobes,
distapectinae ; genacerores 0-2(4-9)3-5. — Aspidiotus os-
becklae Green. — India, Ceylon on Atylosia
orientalis Newst.
ee. Pygidium with densariae between median pair of lobes.
f. Pygidium with second pair of lobes smaller than median,
subspatulate, longer than broad, bluntly rounded, faint
mesal and lateral notches; median pair of lobes quad-
rangular, distal end bluntly rounded, distinct mesal and
lateral notches; third pair of lobes smaller than sec-
ond, subspatulate, bluntly pointed, sometimes faint lat-
eral notch; pectinae deeply lacinate, slightly shorter to
slightly longer than lobes, arranged 2, 2, 3, 3, those of
each lateris deeply lacinate unapectinae, located adjacent
to third lobe; anus four or five times its own width
from caudal margin; genacerores 0-1(3-8)3-5; dorsal ora-
ceratubae in two rows on each side; each lateris with
three small serrated projections. — Cuba on Cocos,
Citrus. cocotiphaga Marl.
ff. Pygidium with second pair of lobes short, much broader
than long, oblique, mesal angle pointed or rounded;
third pair of lobes smaller than second, similar in form,
as broad as or broader than long; median pair of lobes
large, quadrangular, distal end bluntly rounded, dis-
tinct mesal and lateral notches; pectinae subequal in
length to median pair of lobes, narrow, deeply lacinate,
arranged 2, 2, 1-3, 3, those of each lateris not adjacent,
not adjacent to third lobe; anus about three times its
own width from caudal margin; genacerores (0-4)0-2,
usually (1-2)1-2. — South Africa on Acacia, Acer, Ber-
beris, Ceratonia, Cotoneaster, Crataegus, Gleditschia,
Rhus, pear, privet, alder, almond, apple, apricot, ash,
persimmon, plum, poplar, quince, pepper, willow
pectinata Lindgr.
dd. Pygidium with single pectina on each lateris; median pair
of lobes quadrangular, faint mesal and lateral notches, dis-
tant, broader than long; second pair of lobes short and broad,
twice as broad as long, smaller than median, mesal and lat-
eral notches; third pair of lobes rudimentary, hardly pro-
jecting, crenulate; pectinae slender, arranged 2, 2, 3, 1, those
of median incisura narrow furcapectinae, all others unapecti-
nae, those of each lateris distant from third lobe; gena-
cerores 0-2(4-9)7-10. — Italy on cherry patavina Berl.
TRIBE ASPIDIOTINI 409
cc. Pygidium without pectinae or plates on lateres.
d. Pygidium with an oblique row of oraceratubae extending
cephalad from each third incisura, laterad of this another
row of irregularly alternate pores and none from second in-
cisurae; median pair of lobes subquadrangular, with small
mesal and large lateral notch, rounded; second pair of lobes
small, broader than long, lateral margin oblique, notched;
third pair of lobes similar to second, wider; median incisura
with spine-like plates, each second incisura with two plates
or two furcapectinae or finely lacinate unapectinae or one
plate and one furcapectina; setae only slightly longer than
lobes; genacerores 0-4(8-14)5-11. — Europe on oak
zonata Frauenf.
dd. Pygidium with oblique row of oraceratubae extending cephal-
ad from each third incisura, cephalad of this another row of
irregularly arranged pores and a row of about five oracera-
tubae from each second incisura.
e. Pygidium with genacerores arranged 0-8(10-14)8-11; me-
dian pair of lobes subquadrangular, large lateral notch or
sinuate, rounded; second pair of lobes small, longer than
broad, lateral margin oblique, two lateral notches; third
pair of lobes minute, bluntly pointed; pectinae short, those
of median and second incisura not longer than lobes, furca-
pectinae, arranged 2, 2, 3, 0, those of each third incisura
unapectinae; setae very long, much longer than lobes;
genacerores 0-8(10-14)8-11. — Aspidiotus juglandis Colvee,
Aspidiotus juglans-regiae albus Ckll., Aspidiotus juglans-
regiae kafkae Ckll. — United States east of Rockies, Canada,
New Mexico, California, Switzerland, Spain on walnut,
apricot, apple, pear, peach, cherry, plum, locust, maple,
etc. juglans-regiae Comst.
ee. Pygidium with genacerores arranged 0-1(6-7)5-7; four
series or rows of oraceratubae. — New Mexico on plum,
Liquidambar. juglans^egiae pruni Ckll.
SPECIES OF QUADRASPIDIOTUS
a. Pygidium with six or more pectinae on each lateris.
b. Pygidium with two caudal pectinae of each lateris longer than
cephalic pectinae.
c Pygidium with six pectinae on each lateris, three caudal much
larger than three cephalic, pectinae forming a scaly fringe in
region of lobes; median pair of lobes largest, not contiguous;
anus large, about twice its own length from caudal margin;
genacerores (4-8)5-5. — Arizona, Mexico on ash. townsendi Ckll.
cc. Pygidium with pectinae long, scarcely lacinate or dentate, four
caudal pectinae of each lateris much larger than three cephalic;
median pair of lobes large, prominent, adjacent, not touching,
obliquely truncate, distinct lateral and faint mesal notches;
second pair of lobes small, broad, mesal angle longest; mesal
410 THE COCCIDAE
densaria of second incisura largest; pectinae small, about five,
arranged 2, 0, 3; dorsal oraceratubae conspicuous; genacerores
(6-7)2-4. — Colorado, New Mexico on plum howardi Ckll.
bb. Pygidium with pectinae and plates of each lateris subequal in
size; median pair of lobes large, quadrangular, distinct lateral
and mesal notches; second pair of lobes one-third or less the
width of median lobes, shorter, small lateral and mesal notches,
hyaline; small third pair of lobes sometimes present; pectinae
subequal in length to lobes, arranged 2, 2, 5-6, those of incisurae
and three cephalad of each second iobe furcapectinae, others
plates; genacerores 0-1(4-7)4-6; densariae not prominent, subequal.
— Virginia, Ohio on Epigaea. epigaeae Marl.
aa. Pygidium never with more than three pectinae or plates on each
lateris.
b. Pygidium with plates in median incisura.
c. Pygidium with second pair of lobes small, usually narrow,
longer than broad, if broader than long, never twice as broad
as long,
d. Pygidium with mesal and lateral margins of second densariae
subequal in length.
e. Pygidium with median pair of lobes quadrangular, broader
than long, rounded, prominent lateral notches, mesal mar-
gins suboblique; second pair of lobes usually as broad as,
sometimes broader, than median, but much shorter, lateral
margin oblique and emarginate or irregularly notched; me-
dian incisura with two short plates, each second incisura
with two plates or finely serrate pectinae, each lateris with
two plates or two furcapectinae in third incisura and a
single unapectina; setae long, particularly those of second
pair of lobes; genacerores 5-8(10-12)10-16; anus five times
its width from caudal margin. — Aspidiotus ostreaeformis
oblongus Goethe, Aspidiotus ostreaeformis magnus Goethe.
— British Columbia, Idaho, California, Eastern Canada,
New York, Michigan, Europe on apple, pear, plum, peach,
cherry, birch, poplar, horse-chestnut, linden, alder, maple,
oak, date-palm, Crataegus, Calluna.__05treae/ormis Curtis.
ee. Pygidium with the median pair of lobes projecting, more
or less parallel, longer than broad, distal end uniformly re-
duced, mesal and lateral notches; second pair of lobes very
small, broad with irregular margin; genacerores always in
four groups, (4-11)3-9; other structures assumed to be same
as in ostreaeformis. — Bavaria, Prussia, Austria, Norway,
England, Portugal on Calluna, Erica davaricus Lindgr.
dd. Pygidium with mesal margins of second densariae longer
than lateral; median pair of lobes wide, sometimes with
lateral notch; second pair of lobes with prominent mesal
angle, distal margin oblique, one to four notches; plates
small, arranged 2, 2, 3; genacerores 2-6(5-12)5-7. — Aspidiotus
fernaldi albiventer Hunter, Aspidiotus fernaldi hesperiua
TRIBE ASPIDIOTINI 411
Ckll. — Massachusetts, Kansas, Arizona on Juneberry, maple,
Gleditschia. fernaldi Ckll.
cc. Pygidium with second pair of lobes twice or more as broad-tj,8-
long, projecting only slightly, entire or more or less serrate;
median pair of lobes broad, short, entire, broader than long,
hardly projecting; incisurae not prominent; densariae in me-
dian incisurae prominent, mesal margin of second and third
densariae broader than lateral, second densariae longer than
median; pectinae not as long as lobes, furcapectinae, arranged
2, 2, 0; setae of second pair of lobes and lateres large, promi-
nent; oraceratubae in longitudinal rows; genacerores 4-5 (IS-
IS) 7-10; anus small, over five times its width from caudal mar-
gin.— Ohio on Pinus. glanduliferus Ckll.
bb. Pygidium with median pair of lobes contiguous and without plates
in median incisura; median pair of lobes with lateral margin cren-
ulate; plates short and inconspicuous; genacerores 4-5(10-10)7-8. —
Japan on Quercus. cryptoxanthus Ckll.
SPECIES OF DIASPIDIOTUS
a. Pygidium always with pectinae or plates in median incisura.
b. Pygidium with pectinae in median incisura, often minute and
lacinations difficult to identify.
c. Pygidium with median pair of lobes with distinct mesal and
lateral notches,
d. Pygidium with median pair of lobes entire, quadrangular,
mesal margins oblique, convergent, mesal and lateral notches,
latter deepest; two pairs of densariae; pectinae not deeply
lacinate, arranged 2, 8-9, those of each lateris arranged in
three groups, two associated with each second densaria, three
with each third, and three or four cephalad of third densaria,
two cephalic groups unapectinae; genacerores 0-8(6-14)5-8. —
Eastern United States, Colorado, New Mexico, Washington,
Germany on apple, pear, quince, cherry, peach, linden, oak,
maple, beech, ash, osage orange, hackberry, currant, etc
ancylus Putn.
dd. Pygidium with median pair of lobes with distal ends finely
serrate; densariae of second incisura about equal in length.
— Iowa on willow ancylus serratus Newl. & Ckll.
cc. Pygidium with median pair of lobes with mesal notch wanting,
lateral notch frequently obtuse or nearly wanting; mesal
densaria of each second incisura slightly larger than lateral;
densariae of median incisura larger and more marked. — Iowa
on Mountain ash. ancylus latilobis Newl.
bb. Pygidium with plates in median incisura.
c. Pygidium with two plates, often minute and process-like, in
median incisura and with more than three pectinae cephalad of
each third incisura.
d. Pygidium with mesal margins of densariae longer than lat-
eral; median pair of lobes quadrangular, distal end bluntly
412
THE COCCIDAE
rounded, prominent lateral notch; pectinae with slight laci-
nations, arranged 2, 10 or more, those of each lateris near a
lobe; dorsal oraceratubae few in number, arranged in three
rows; setae as long as or longer than pectinae; mesal calles
fused and distant from lateral calles, latter L-shaped; anus
three times its width from caudal margin, about five times
from vulva, and about eleven times from mesal calles; gena-
cerores (4-4)4-7. — Arizona on Prosopis candidula Ckll.
dd. Pygidium with mesal margins of densariae not longer than
lateral.
e. Pygidium with genacerores( 8-10) 5-6; median pair of lobes
Quadrangular, rounded, lateral and mesal notches, mesal
margins adjacent, divergent; margin of pygidium cephnlad
of each incisura produced spine-like; pectinae not deeply
lacinate, plates and pectinae, arranged 2, 7, plates of me-
dian incisura short, about as long as lobes, those of each
lateris in three groups, 2-3-2, two cephalic ones plates,
others unapectinae. — Aspidiotus cydoniae Comst. — Southern
States, Kansas, New Mexico, Mexico, West Indies, Ceylon,
Samoa on quince, fig, palms, orange, tea-plant, cactus,
Latania, Jasminum. lataniae Sign.
ee. Pygidium with genacerores 3-7(12-15)7-11; median pair of
lobes broad, prominent lateral notch, sometimes with mesal
notch; densariae fused, forming semicircular thickening,
those of second incisurae larger than those of third; plates
of median incisura short, inconspicuous, chitinous proces-
es, each lateris with 8-10 plates and f urcapectinae ; dorsal
oraceratubae numerous and prominent. — Ohio on Aesculus.
ohioensis York.
CO. Pygidium with a single plate in median incisura, three pectinae
on each lateris in second incisura and two in third; median
pair of lobes quadrangular, faint lateral notches, angles round-
ed; densariae of second incisura with mesal margins larger
than lateral, third incisura with lateral margin nearly obsolete;
genacerores 0-5(6-10)11-11. — Iowa, Texas, on currant
hunteri Newl.
aa. Pygidium without pectinae or plates in median incisura.
b. Pygidium with pectinae on each lateris.
c. Pygidium with median pair of lobes with mesal and lateral notch-
es, not adjacent, quadrangular, mesal margins oblique; two
pairs of densariae, two unapectinae associated with each second
incisura and three with third and 3-5 plates cephalad of third;
lateres finely serrate; genacerores 0-4(4-9)3-7. — Ohio, Illinois,
Indiana, Tennessee, Kansas, Florida, Jamaica, Europe on Vitis,
hickory. uviae Comst.
cc. Pygidium with median pair of lobes with lateral notch only,
d. Pygidium with three pairs of densariae, mesal thickening of
densariae of second and third incisurae only slightly longer
than lateral; median pair of lobes large, quadrangular, dis-
TRIBE ASPIDIOTINI 413
tinct lateral notch, distal margin bluntly rounded; anus two
to two and one-half times its width from caudal margin;
pectinae small, furcapectinae, arranged 0, 4, those of each,
lateris arranged 2-2-0; dorsal oraceratubae arranged in lon-
gitudinal rows; genacerores 0-3(15-23)6-14. — Ohio on Lirio-
dendron. piceus Sands.
dd. Pygidium with three pairs of densariae, mesal thickening of
densariae of second and third incisurae much broader and
longer than the lateral, those of second incisura much larger
than those of third; median pair of lobes large, quadrangu-
lar, convergent, distinct lateral notch, distal margin rounded;
anus twice its width from caudal margin; each second in-
cisura with two long furcapectinae and each third with three
long plates; genacerores (11-15)7-10. — Texas on Morus
mori Herrick.
bb. Pygidium without pectinae on each lateris, but with plates or at
most with a single pectina.
c. Pygidium with two or three pairs of densariae.
d. Pygidium with densariae in median incisura; median pair of
lobes subquadrangular, longer than broad, distal end rounded,
lateral notch only; densariae of median incisura ovate in out-
line, those of second and third incisurae subequal, those of
third incisura smaller than those of second; genacerores
0-2(5-7)3-4. — Kansas, Iowa, Georgia on Quercus, Ostrya
oshorni Newl. & Ckll.
dd. Pygidium never v/ith densariae in median incisura.
e. Pygidium with two pairs of densariae, those of mesal side
of second incisurae terminating in distinct oval knob,
longer than those of lateral side, those of second incisura
smaller, mesal side longer than lateral; median pair of
lobes quadrangular, two lateral notches, minute mesal
notch, sometimes wanting; anus minute, in line with mesal
densariae of second incisurae; plates inconspicuous; mesal
calles fused, sometimes separate, distant from oblique lat-
eral calles; genacerores (1-3)0-1. — Mexico on Coursetia
coursetiae Marl,
ee. Pygidium with two pairs of densariae, those of second and
third incisurae subequal in size, none of them swollen.
f. Pygidium with median pair of lobes twice as long as
width of anus, anus five times its width from caudal
margin and about ten times from vulva; median pair of
lobes large, prominent, bluntly rounded, distinct later-
al notch, mesal margins divergent; plates short, not as
long as median pair of lobes, arranged 0, 3-4, two in
each second incisura and one or two in third, these
sometimes lacinate; mesal calles fused, lateral calles
distinct, distant from mesal; genacerores 2-2(5-9)3-7. —
California on Abies, Libocedrus, Pseudotsuga
ehrhorni Colm.
414 THE COCCIDAE
ft. Pygidium with median pair of lobes not or only slightly^
longer than width of anus.
g. Pygidium with median pair of lobes quadrangular^
close together, two distinct lateral notches, mesal
wanting; plates short and inconspicuous; anus large,
less than twice its own width from caudal margin;
dorsal oraceratubae numerous, confined chiefly to two
irregular, double, lateral rows; calles not especially
prominent, mesal calles distant from each other and
from lateral calles; genacerores (3-9)3-8. — New Mex-
ico on Cottonwood popularum Marl.
gg. Pygidium with median pair of lobes quadrangular,
single large lateral notch; plates small, arranged 0,
4-6, those of lateres arranged 2-1 to 3-1 or more, one
of second group furcapectina; anus small, two or
three times its own width from caudal margin; dorsal
oraceratubae fairly numerous; mesal calles fused;
genacerores 0-3(9-14)6-9. — Aspidiotus aesculi solus
Hunter. — Kansas on Juglans solus Hunter.
cc. Pygidium with single pair of densariae, located in second in-
cisurae, subequal in length, if third incisurae with densariae,
these very small; lobes subquadrangular, nearly as broad as
long, not strongly projecting, distinct lateral notch, mesal want-
ing; plates small, inconspicuous, arranged 0, 3, those of each
lateris arranged 1-2 in incisurae; setae about as long as or
longer than lobes; lateres entire; anus five times its width from
caudal margin; calles not fused, not prominent, mesal calles
fused, distant from oblique lateral calles; genacerores (5-17)
4-11. — California on Aesculus. aesculi Johns.
SPECIES OF CHRYSOMPHALUS
a. Pygidium never with more than five pairs of paraphyses.
b. Pygidium with three pairs of paraphyses.
c. Pygidium with first pair of paraphyses longest, cephalic end
enlarged, knob-like, second pair similar in form, shorter, third
pair about one-half length of first pair, not enlarged at cephalic
end; median pair of lobes longest, faint mesal and lateral
notches, bluntly pointed; second, third, and fourth pairs blunt-
ly pointed, second pair smaller than others, much nearer me-
dian lobe than third; pectinae small, obscure, arranged 2, 2,
3, 3, 0; lateres practically entire; genacerores forming almost
continuous arc, 75-100 cerores. — Ceylon on Murraya
quadriclavatus Green.
cc. Pygidium with first pair of paraphyses about one-half length
of subequal second and third pairs, these latter enlarged at
cephalic end, club-shaped; median pair of lobes subtriangular,
lateral margin oblique, emarginate, serrate; second md third
pairs of lobes similar to median and subequal, serrate; pectinae
short, arranged 2, 2, 2, 0; each lateris serrate, bearing many
TRIBE ASPIDIOTINI 415
closely placed short oraceratubae ; head and thorax with several
long setae with tubular bases; genacerores (10-14)10-12. — Japan
on Quercus. setiger Mask.
bb. Pygidium with more than three pairs of paraphyses.
c. Pygidium with four pairs of paraphyses, about as long as me-
dian pair of lobes.
d. Pygidium with median pair of lobes entire, well separated,
about as broad as long, rounded; second pair of lobes distant
from median, distance equal to width of median lobe, broader
than long, entire; third pair of lobes pointed, tooth-like, less
conspicuous; each lateris with lobe-like projection; anus near
cephalic ends of paraphyses; genacerores (4-4)3-3. — Mexico
on orange. koebelei Ckll. & Towns.
dd. Pygidium with median pair of lobes with mesal and lateral
notches, bluntly rounded between notches; second pair of
lobes similar in form, slightly smaller; third pair of lobes
more irregular, conspicuous lateral notch; pectinae slightly
longer than lobes, arranged 2, 2, 3, 3; each lateris adjacent to
cephalic pectina with stout marginal prominence; paraphyses
slender, sides subparallel; anus three times its width from
caudal margin; dorsal oraceratubae in two rows on each side;
genacerores (3-4)2-3; vulva between pregenacerores. — Sey-
chelles on Cocos. ansei Green.
cc. Pygidium with five pairs of paraphyses.
d. Pygidium with fourth pair of paraphyses not more than one-
half length of third, second pair longest, first, third, and
fifth long, slightly shorter than second; median pair of lobes
longer than broad, bluntly rounded, prominent lateral notch;
second pair of lobes smaller than median, bluntly pointed,
entire; third pair of lobes smaller than second, elongate,
lateral margin notched; pectinae short, arranged 2, 2, 3, 4,
those of median and second incisurae distapectinae, others
unapectinae; genacerores (3-4)2-3; thorax not produced on
each side into a tooth. — Jamaica on mango. — mangiferae Ckll.
dd. Pygidium with fourth pair of paraphyses as long or much
longer than fifth.
e. Pygidium with all paraphyses of each side very different in
length.
f. Pygidium with first pair of paraphyses shorter than
others, second and fourth pairs longest, third and fifth
intermediate in length; median pair of lobes rounded,
deep lateral notch, mesal margins convex; second pair of
lobes nearly as large as median, similar in shape, lateral
notch; third pair of lobes elongate; bluntly rounded, lat-
eral notch; pectinae large, arranged 2, 2, 3, 6-7,
those of each lateris unapectinae, others distapectinae;
genacerores (4-8)2-4; thorax produced on each side into
a short prominent tooth-like projection. — Chrysomphalus
proposimus Banks. — India, Japan, Australia, Brazil,
416 THE COCCIDAE
Mexico, California, eastern United States, Europe,
Egypt, Ceylon, Mauritius, Natal, south Africa on orange,
lemon, cocoanut, banana, palms, rose. Ilex, Camellia,
Dictyosperma, Phyllodendron, guava, grape-fruit, olean-
der, fiig. Begonia, etc. aonidum Linn.
ff. Pygidium with first pair of paraphyses nearly as long as
fourth and more than twice as long as second, third,
and fifth, fourth pair very slightly longer than first;
median pair of lobes slightly wider than long, distal
margin transverse, subtruncated, slightly indented; sec-
ond and third pairs of lobes slightly smaller, similar in
form, broader than long; pectinae and plates subequal
in length to lobes, arranged 2, 2, 3, 3, caudal one of each
of two last groups a plate; each lateris thickened, four
or five sharp tooth-like projections with serrate edges;
genacerores (6-10)3-7; caudal portion of each lateral
margin of thorax with tubercle, not bearing a horn. —
Brazil on Laurus. pauUstus Hemp.
ee. Pygidium with all paraphyses of each side subequal in
length, first pair as long as others; median pair of lobes
bluntly pointed, elongate, prominent lateral notches; sec-
ond pair of lobes similar to first, smaller, notched; third
pair of lobes similar to second, smaller, serrate on lateral
margin; pectinae elongate, arranged 2, 2, 3, 5, those of
third incisurae and lateres unapectinae; genacerores (3-4)
1-3; thorax produced on each side into prominent tooth-
like projection. — Italy, Jamaica, Demerara on Ooton,
Cycas, Pandanus, rose, mango pinnuliferus Mask.
aa. Pygidium always with more than five pairs of paraphyses.
b. Pygidium with six or seven pairs of paraphyses.
c. Pygidium with six distinct pairs of paraphyses.
d. Pygidium with first and second pairs of paraphyses never
subequal.
e. Pygidium with second, third, fourth, and fifth pairs of
paraphyses subequal, longer than first and sixth, second
pair slightly longer than third, fourth, and fifth, first
pair shortest, and sixth pair intermediate in length; me-
dian pair of lobes large, broad, distal margin oblique, sinu-
ate; second pair of lobes much shorter and narrower than
median pair, bluntly rounded; third pair smaller than sec-
ond, more pointed; pectinae short, arranged 2, 2, 3, 3, those
of median and second incisurae distapectinae, others
unapectinae; genacerores (4-4)2-3. — Chrysomphalus dictyo-
spermi arecae Newst. — United States, West Indies, Mexico,
Brazil, Demerara, China on Dictyospermum, Erythrina,
Cycas, Latania, palms, rose, mango, etc.- -dictyospermi Morg.
ee. Pygidium never with four pairs of paraphyses subequal in
length.
TRIBE ASPIDIOTINI 417
f. Pygidium with second and fourth pairs of paraphyses
subequal in length. - .
g. Pygidium with fourth and sixth pairs of paraphyses
subequal, second, fourth, and sixth pairs longest,
others shorter and smaller; lobes broad, low, incon-
spicuous, median pair twice as broad as long; second
pair of lobes about or nearly as broad as first; third
" pair of lobes obsolete; plates scarcely visible, third
incisurae with two; each lateris with depression with
pair of spine-like plates, cephalad of this minutely
serrate with three distant pointed prominences; gena-
cerores (8-8)4-7. — Mexico reniformis Ckll.
gg. Pygidium with fourth pair of paraphyses longer than
sixth, fourth and fifth pairs subequal, second pair as
long as fourth, longer than first and third, fourth
pair very slightly longer than subequal second and
fifth, first and third pairs subequal and slightly shorter
than sixth; median pair of lobes elongate, longer than
broad, distinct lateral and mesal notches; second pair
of lobes similar in size and shape to median, distinct
lateral notch; third pair of lobes broad, bluntly
pointed, lateral margin with two or three notches;
pectinae deeply lacinate, arranged 2, 2, 3, 3, those of
incisurae other than median and of each lateris
unapectinae and longer than lobes; lateres with large
tooth-like projection adjacent to pectinae; genacerores
(4-5)1-2. — Australia on Acacia. fodiens Mask.
ff. Pygidium with second and fourth pairs of paraphyses
not subequal in length.
g. Pygidium with fourth pair of paraphyses longer than
sixth, fourth pair longest, second and sixth pairs
subequal and two-thirds length of fourth, first and
third pairs distinctly shorter than second; median
pair of lobes quadrangular, broader than long, entire,
truncate with rounded angles; second and third pairs
of lobes shorter than median and more than twice as
broad, entire, distal margins oblique; pectinae about
as long as median lobes or shorter, arranged 2, 2, 3,
3; lateres with two slight indentations; metathorax
with large blunt marginal tubercle, tubercles and
margins of abdomen strongly chitinized; anus near
middle of pygidium and vulva near middle of cephalic
half; genacerores (6-8)5-6. — British Guiana on
Lecythis. umdoniferus Newst.
gg. Pygidium with fourth pair of paraphyses distinctly
shorter than sixth.
h. Pygidium with fifth pair of paraphyses longer than
sixth, second and fifth pairs longer than others,
first and third pairs subequal, slightly more than
one-half length of second, longer than sixth, sixth
418 THE COCCIDAE
pair nearly twice as long as fourth, shortest; three
pairs of subequal lobes, lateral margins sloping
and serrulate; pectinae shorter than lobes, ar-
ranged 2, 2, 3, 1; genacerores (11-17)12-14. —
Japan. kelloggi Kuwn.
hh. Pygidium with fifth and sixth pairs of paraphyses
subequal, second, fifth, and sixth pairs longest,
first, third, and fourth pairs subequal, much short-
er than others, all enlarged at cephalic end and
club-shaped; median pair of lobes broad, short,
broadly rounded; second pair of lobes subequal in
length, broader than median pair; third pair of
lobes similar to second, broader; pectinae Y-
shaped furcapectinae, arranged 1, 1, 1, 3, caudal
one of each lateris not Y-shaped; genacerores
(8-9)7-9. — England on Cattleya alienus Newst.
dd. Pygidium with first and second pairs of paraphyses subequal,
shorter than third, fourth, fifth, and sixth pairs; median pair
of lobes small, distant, nearly outline of half -circle; second
pair of lobes twice as broad as median; third pair of lobes
fully three times as broad as median, margin minutely ser-
rulate; each lateris minutely serrulate with three distant
notches limiting four broad projections; genacerores (8-9)
6-7. — Mexico on orange. albopictus Ckll.
CO. Pygidium with seven pairs of paraphyses, fifth rarely not as
broad as others.
d. Pygidium with fifth pair of paraphyses distinctly longer than
fourth.
e. Pygidium with fifth pair of paraphyses four times as long
as fourth.
f. Pygidium with second pair of paraphyses shorter than
fifth and three times as long as first, second, fifth, and
seventh pairs longest, first, third, and fourth pairs
shortest, sixth intermediate; median pair of lobes short,
narrow, lateral margin obliquely rounded, entire; sec-
ond pair of lobes short and broad, broader than median
pair, lateral margin broadly obliquely rounded, serrate;
third pair of lobes short and broad, broader than sec-
ond pair, lateral margins obliquely rounded, serrate;
pectinae short, narrow, distapectinae, arranged 2, 2, 2, 6,
those of each lateris arranged 2-1-1-1-1; genacerores (IS-
IS) 7-9. — Mexico, Central America on Avocardo pear.
Citrus, Laurus. scutiformis Ckll.
ff. Pygidium with second pair of paraphyses as long as
fifth, about twice as long as first, seventh pair distinctly
shorter than fifth and longer than first, third, fourth,
and sixth pairs subequal, about one-half length of first;
median pair lobes short, distal end oblique on each side,
bluntly rounded; second and third pairs of lobes smaller
but similar; pectinae very inconspicuous; lateres serru-
TRIBE ASPIDIOTINI 419
late, three or four small projections; genacerores (7-7)
5-5. — Mexico on mangrove. rhizophorae Ckll^
ee. Pygidium with fifth pair of paraphyses not four times as
long as fourth.
f. Pygidium with fifth pair of paraphyses not more than
twice as long as fourth, first, third, and fifth pairs sub-
equal and longest, sixth pair slightly shorter than fifth
and longer than subequal second, fourth, and seventh;
median pair of lobes subquadrangular, as broad as long,
distinct mesal and lateral notches, distal end bluntly
rounded; second pair of lobes similar in form and size
to median pair; third pair of lobes subequal in size to
second pair, distal end bluntly rounded, prominent lat-
eral notch, no mesal notch; pectinae arranged 2, 2, 3, 3,
those of each lateris unapectinae; genacerores 1-2(5-6)
6-6. — Ceylon on unidentified tree pedronis Green.
ff, Pygidium with fifth pair of paraphyses longer than
fourth, not twice as long, fifth pair longest, second,
fourth, and seventh pairs subequal, a little shorter than
fifth, first pair two-thirds length of second and one-
fourth longer than third, third and sixth pairs sub-
equal; median pair of lobes quadrangular, distinct lat-
eral notch, as broad as long; second and third pairs of
lobes similar, smaller than median, lateral margin
oblique, notched; pectinae short, not as long as lobes,
furcapectinae, arranged 1, 1, 1, 2; lateres toothed and
serrate; genacerores 6-9(17-24)9-16. — South Africa on
Virgilia, Erythrina, Celastrus, Schinus, Robinia, olive,
apple, hawthorn, kei-apple, lilac, pear, poplar, plane-tree,
privet, rose, peach, plum, walnut corticosus Brain.
dd. Pygidium with fifth pair of paraphyses not one-half length
of fourth, if subequal in length, narrow and inconspicuous.
e. Pygidium with each lateris entire, serrate, or slightly in-
dented, but never saw-toothed.
f. Pygidium with five or six pectinae on each lateris;
paraphyses short, longest but little longer than adjacent
lobes, fifth pair slender inconspicuous thickening; me-
dian pair of lobes distinctly broader and longer than
second pair, quadrangular, mesal and lateral notches,
broadly rounded; second pair of lobes similar but
smaller; third pair of lobes bluntly triangular, short;
pectinae deeply lacinate, arranged 2, 2, 3, 5-6, those
of median incisura short, furcapectinae, those of other
incisurae unapectinae, lacinations of those of each
lateris reduced in number, particularly cephalic ones;
genacerores (1-3)3-4. — Italy on Camellia
degeneratus Leon.
ff. Pygidium with two or three pectinae on each lateris;
paraphyses in three lengths, second, fourth, and seventh
420 THE COCCIDAE
pairs longest, first and third pairs shortest, fifth and
sixth pairs slightly longer than latter, seventh pair very-
small; median pair of lobes short and narrow, rounded,
entire; second pair of lobes short and broad, broader
than median pair, lateral margin broadly oblique and
crenulate; third pair of lobes short and broad, broader
than second pair, lateral margin broadly oblique,
serrate; pectinae short and narrow, distapectinae, ar-
ranged 2, 2, 2, 2, those of each lateris adjacent to third
lobe; genacerores (10-11)6-7. — Florida, Mexico, England
on Persea, Magnolia, Anthurium, Ilex, cocoanut-palm
perseae Comst.
ee. Pygidium with each lateris with deep indentations, appear-
ing distinctly saw-toothed.
f. Pygidium with fourth pair of paraphyses distinctly
longer than third or fifth, never twice length of either,
first pair shortest, third and fifth pairs next in length,
sixth and seventh pairs next; median pair of lobes
broad, broadly rounded, subquadrangular, slight mesal
and lateral notches; second and third pairs of lobes sim-
ilar, broader than long, lateral margin longest and
strongly oblique, crenulate, subtriangular, bluntly round-
ed; pectinae short, narrow, distapectinae or furcapecti-
nae, arranged 1, 1, 2, 2; each lateris with about four
teeth; genacerores 3-3(8-11)5-7. — District of Columbia,
Ohio, Illinois, Kansas, Florida on oak, hickory, pecan,
grape. obscvrus Comst.
ff. Pygidium with fourth pair of paraphyse? three times
length of third and twice length of fifth, fourth and
seventh pairs subequal and longest, first, third, and sixth
pairs subequal and shortest, fifth pair distinctly longer
than third and slightly shorter than fourth; paraphyses
knob-like at cephalic end; median pair of lobes
broad, broader than long, caudal margin longest; sec-
ond and third pairs of lobes similar in form, wide, third
pair widest; each lateris with about six tooth-like pro-
jections; pectinae present, exceedingly minute; spira-
cerores wanting; genacerores (6-9)3-5. — British Guiana
on Erythraspis. erythraspidis Newst.
bb. Pygidium with nine pairs of paraphyses.
c. Pygidium with third pair of paraphyses four times as long as
fourth, second pair shortest, fourth, fifth, and seventh pairs
slightly longer, third and sixth pairs longest, first, eighth, and
ninth pairs intermediate; median pair of lobes broadly round-
ed, slight lateral notch; second and third pairs of lobes similar,
not notched ; pectinae truncate, arranged 2, 2, 2, 4, those of each
lateris f urcapectinae ; genacerores (4-4)3-4, — New Zealand,
Louisiana on Linum. sphaerioides Ckll.
cc. Pygidium with third and fourth pairs of parphyses subequal in
length.
TRIBE ASPIDIOTINI 421
d. Pygidium with second, third, and fourth pairs of paraphyses
subequal in length.
e. Pygidium with each lateris not deeply notched; paraphyses^
long, subequal, all much longer than lobes; median pair of
lobes quadrangular, bluntly rounded, distinct mesal and
lateral notches; second and third pairs of lobes similar
but smaller; pectinae not deeply lacinate, arranged 2, 2, 3,
1-2, those of median incisura narrow, distapectinae, all
others unapectinae; each lateris with seven tooth-like pro-
jections; genacerores (9-12)8-9, according to Brain (7-9)
3-5. — Chrysomphalus rossi victoriae Ckll. — Australia, New
Zealand, Philippine Islands, Japan, China, Ceylon, South
Africa on Xanthorrhoea, Eucalyptus, Nerium, Caffaris,
Rhinocarpus, Acacia, Araucaria, Abutilon, Artemesia, olive,
plum, etc. rossi Mask.
ee. Pygidium with each lateris deeply notched or "bayed in;"
genacerores (9-14)7-11. — Ceylon, South Africa on Capparis,
Chaetachme. rossi greeni Brain & Kelly.
dd. Pygidium with second pair of paraphyses twice as long as
either third or fourth pairs.
e. Pygidium with second and third pairs of lobes subequal to
or narrower than median pair; median pair of lobes short
and broad, lateral margin long, oblique, rounded, entire;
second and third pairs of lobes similar, faintly notched;
pectinae short, distapectinae, arranged 2, 2, 2, 5, those of
each lateris arranged 2-1-1-1; each lateris with about six
tooth-like projections; paraphyses variable in length, first,
third, fourth, sixth, seventh, and ninth subequal in length,
shorter than second, fifth, and eighth, fifth longest, second
dilated at cephalic end; genacerores 4-8(15-16)8-11. — Mex-
ico on "Trueno." nigropunctatus Ckll.
ee. Pygidium with second and third pairs of lobes wider than
median; median pair of lobes contiguous, widened; third
pair of lobes subequal to second ; second incisurae less than
one-half width of median lobe; third incisurae equal to
width of median lobe; each incisura with angular pro-
jection, cephalad of this serrate; this species is allied to
nigropunctatus; paraphyses are not described; genacerores
(13-16)8-8. — Mexico on Agave agavis Towns. & Ckll.
SPECIES OF PSEUDISCHNASPIS
Pygidium with six pairs of paraphyses, first and third pairs sub-
equal, second and fifth pairs subequal and three times as long as first;
fourth and sixth pairs subequal and about twice as long as first,
median pair of lobes subquadrangular, broadly rounded, distal mar-
gin with two notches; second and third pairs of lobes broader than
median, third broader than second, lateral margin oblique, serrate;
furcapectinae arranged 1, 2, 2, 1-3; lateres thickened and toothed;
genacerores (6-8)4-7. — Brazil on Myrcia. linearis Hemp.
422 THE COCCIDAE
aa. Pygidium with six pairs of paraphyses, first pair slightly longer than
third, second and fifth pairs subequal and longest, as long as third,
fourth and sixth pairs subequal, slightly longer than third.
b. Pygidium with three pairs of lobes, successively larger from first
to third, latter with two small notches on lateral margin; each
lateris with three serrations; genacerores 2-2(5-5)3-3, pregenacer-
ores and postgenacerores of each side forming nearly continuous
row. — Mexico on mango longissima Ckll.
bb. Pygidium with three pairs of lobes; median pair subquadrangu-
lar with rounded caudal ends, close together, entire; second and
third pairs of lobes broader and larger than median, lateral mar-
gin oblique, faintly serrate; pectinae short, narrow, distapectinae,
subequal in length to lobes, arranged 2, 2, 2, 5, each lateris with
two pectinae adjacent third lobe and three truncate plates distrib-
uted along margin cephalad of pectinae; each lateris with five
distinct serrations; genacerores (7-7)8-8. — Jamaica, Mexico on
Agave. towreyi Ckll.
SPECIES OF LINDINGASPIS
Pygidium with three pairs of more or less unsymmetrical subequal lobes,
broader than long, broadly rounded, margin finely crenulate; pectinae
broad, slightly longer than lobes, lacinations deep, arranged 1, 1, 1-2, 1,
one of each lateris adjacent third lobe; each lateris with six or more
large, equidistant, tooth-like projections, smaller teeth between two
caudal projections; spiracerores wanting; genacerores in two groups,
8-10; dorsal oraceratubae prominent, longitudinal row extending
cephalad from each third incisura; paraphyses in five pairs, first and
second pairs associated with median lobes, third pair with second pair
of lobes, fourth pair longest and opposite third incisura, fifth pair op-
posite caudal margin of third pair of lobes. — Melanaspis samoana
Lindgr. — Samoa on Myristia. samoana Lindgr.
SPECIES OF FORBESASPIS
Pygidium with median pair of lobes subtriangular, converging, sub-
adjacent, mesal portion longest, distal end bluntly rounded, lateral mar-
gin oblique, deep lateral notch; second pair of lobes small, not strongly
protuberant, distinct lateral notch; densariae present between median
pair of lobes, mesal side of densariae of second incisurae much larger
than minute lateral, those of third incisurae subequal, fused; gena-
cerores 0-4(3-7)3-5. — Canada, eastern United States, New Mexico,
Mexico, Porto Rico, Germany, south Africa on plum, peach, pear, apple,
quince, cherry, currant, gooseberry, apricot, beech, hawthorn, honey-
locust, walnut, Jasmine, Acer forhesi Johns.
SPECIES OF FERRISASPIS
Pygidium with median pair of lobes prominent, margins convex, distal
end rounded, prominent lateral notch; lateres entire, second and third
incisurae with densariae, mesal and lateral margins subequal, those of
second much larger; dorsal oraceratubae numerous, arranged in oblique
band nearly parallel with margin; setae as long as median lobes; anus
TRIBE ASPIDIOTINI 423
small, about five times its width from caudal margin and fifteen from
vulva; genacerores 0-2(2-5)2-5; mesocalles fused and distant_from
latacalles. — Arizona on Covillea. covilleae Ferns.
SPECIES OF CONSTOCKIELLA
a. Pygidium with six groups of genacerores.
b. Pygidium with genacerores 4-6(4-7)7-15; caudal margin deeply and
roundly notched at meson, pair of short setae in notch; each
lateris with five long setae and three short ventral setae located
just within margin; four dorsal oraceratubae in front of .notch
in two rows, two or three rows extending length of pygidium on
each side of anus; anus located five times its width from caudal
margin and about one-half its width cephalad of vulva. — Florida,
Guadaloupe Island on Erythea, palmetto sabalis Comst.
bb. Pygidium with genacerores 11-15(7-10)14-17. — Mexico on palms.—
sabalis mexicana Ckll.
aa. Pygidium with five groups of genacerores; margin with lobes little
developed, represented by three pairs of broad low rounded promi-
nences, all wide apart, incisurae and especially lateres irregular with
more or less angular prominences; sides of caudal part of insect
finely transversely striate; setae and pectinae or plates not notice-
able; paraphyses and densariae wanting; dorsal oraceratubae many,
row of transverse ones; genacerores 5-6(21-22)22-23; scale long, oval,
moderately convex, rough, grayish brown. — Chile on Chisquea. — This
like the type species has many characteristics of the Diaspidini
riverae Ckll.
SPECIES OF LIGULASPIS
Pygidium with three groups of genacerores, sometimes almost united into
single group, mesogenacerores 27, latagenacerores 106; pygidial mar-
gin with five plate-like projections, mesal longest, narrowest, three-
lobed, others irregularly notched and toothed; lateral margin of two
caudal segments of preabdomen chitinized and plate-like; oraceratubae
numerous on pygidium and segments of body; anus located just caudad
of mesogenacerores; mesospiracerores and metaspiracerores 36. —
Odonaspis janeirensis Hemp. — BraEil on grass janeirensis Hemp.
SPECIES OF ODONASPIS
Pygidium with single fused median lobe with prominent lateral notches,
other lobes obsolete; each lateris with four prominent indentations
from which longitudinal thickenings extend, indentations limited by
angular teeth on cephalic side, serrate between indentations; gena-
cerores in two converging groups, each consisting of 80-90; mesospira-
cerores and metaspiracerores less than 10; anus nearer vulva than
caudal margin. — Odonaspis secreta saccharicaulis Zehnt., Odonaspis
secreta greeni Ckll. — Japan, Hawaiian Islands, Java, Ceylon on bam-
boo, Arundinaria. secreta Ckll.
SPECIES OF BETiLESASPIDIOTUS
Pygidium without lobes, plates, or pectinae; no mesal indentation, in-
424 THE COCCIDAE
dentation at caudal end of second pair of paraphyses; lateres with two
deep notches; two pairs of equidistant paraphyses; anus located near
vulva, distant from caudal margin, near mesogenacerores ; genacerores
in three groups, mesogenacerores 50 and latagenacerores 150. — Odon-
aspis (Anoplaspis) bambusarum Ckll. — Japan, California on bamboo.
tam'busarum Ckll.
SPECIES OF NEOSIGNORETIA
a. Pygidium without pectinae or plates on lateres.
b. Pygidium with three pairs of lobes.
c. Pygidium with median pair of lobes notched,
d. Pygidium with plates in median incisura; median pair of
lobes wide, others rudimentary, all small; plates long and
slender, sharp, conspicuous, distinctly longer than lobes, ar-
ranged 2, 1, 1, 0; anus located near caudal margin. — The
reference of this species to Pseudodiaspis needs confirmation.
— Targionia yuccae neomexicana Ckll. — Mexico on Yucca
yuccae Ckll.
dd. Pygidium without plates or pectinae in median incisura;
median pair of lobes short, subquadrangular, deep lateral
notch; second and third pairs of lobes rudimentary, second
pair larger and with lateral notch; plates and pectinae about
as long as median lobes, arranged 0, 2, 2, 0, mesal projection
of second and third incisurae a plate and lateral pectina;
setae about as long as median lobe; dorsal oraceratubae
prominent, not numerous; anus small, distant four or five
times its width from caudal margin. — California on Quercus.
yulupae Bremner.
CO. Pygidium with median pair of lobes entire, short, truncate or
bluntly rounded; second pair of lobes broader than median,
short, broadly rounded, entire; third pair of lobes represented
by stout marginal prominence; median incisura about as wide
as median lobe, with two short blunt plates, not longer than
lobes, second incisura with two similar plates; dorsal oracera-
tubae small and inconspicuous; anus three times its width
from caudal margin; scale of adult female snowy white,
strongly convex. — Australia on Styphelia. --immacwZata Green,
bb. Pygidium with four pairs of lobes; median pair of lobes small,
moderately prominent, constricted at proximal end, hatchet-
shaped, lateral and distal margins forming continuous curve; sec-
ond, third, and fourth pairs of lobes simply marginal prominences,
angular; plates or pectinae of median and second incisurae not
longer than median lobes, of fourth incisurae longer, arranged
2, 2, 3, 4, 0, those of median, second, and third incisurae plates,
others pectinae; anus minute, ten times its width from caudal
margin and over twenty times from vulva; dorsal oraceratubae
in two longitudinal rows; second and third incisurae with two
pairs each of minute thickenings; calles present and separate. —
Northern Australia on Eucalyptus. miniatae Green.
aa. Pygidium always with pectinae or plates on lateres.
TRIBE ASPIDIOTINI 425
b. Pygidium with median pair of lobes with lateral notches.
c. Pygidium with median pair of lobes with mesal and lateral
notches,
d. Body with margin of cephalic end entire, not finely crenu-
late; pygidium with median pair of lobes quadrangular, deep
lateral and mesal notches; second pair of lobes spatulate,
bluntly pointed or bluntly rounded, entire or with faint lat-
eral notch; third pair of lobes spatulate, subequal to second
pair, bluntly rounded, entire or with faint lateral notch;
pectinae long and narrow, subequal in length to lobes, ar-
ranged 2, 2, 3, 8, those of each lateris unapectinae and
shorter than others, placed cephalad of third lobe; caudal
pectina of each third incisura much smaller than other two.
— South Africa on Euphorbia. tectaria Lindgr.
dd. Body with margin of cephalic end finely crenulate; pygidium
with median pair of lobes quadrangular, deep mesal and lat-
eral notches, distal margin rounded between them; second
pair of lobes broader than long, lateral margin oblique, deep
lateral notch, mesal portion projecting; third pair of lobes
narrower, deep lateral notch, projecting mesal portion more
angular; distapectinae in median incisura, others unapecti-
nae, latter longer than lobes, arranged 2, 2, 3, 3; spiracerores
wanting; anus about twice its width from caudal margin;
vulva located near cephalic half of pygidium. — Gymnaspis
africana Newst. — Uganda, Africa. africana Newst.
cc. Pygidium with median pair of lobes with distinct lateral
notches only, projecting, distinct, about as broad as long, distal
end bluntly rounded; second pair of lobes smaller, triangular,
bluntly pointed; third pair of lobes minute, pointed; pectinae
minute, short, narrow, lacinations minute, arranged 2, 2, 2, 1,
those of second and third incisurae and of each lateris
unapectinae; anus distant three or four times its width from
caudal margin. — Sardenia, Italy on Osirys cecconi Leon.
bb. Pygidium with median pair of lobes entire, large, broadly bluntly
rounded, distant; second and third pairs of lobes minute, subtri-
angular, more or less pointed; pectinae arranged 2, 2, 2, 5, those
of incisurae and three caudal ones of each lateris very broad,
deeply lacinate at distal end, cephalic two of each lateris slender,
unapectinae with short blunt branches; pectinae of each lateris
adjacent third lobe; setae limited to two on each lateris near
cephalic pectinae; center of pygidium cephalad of anus with large
tongue-shaped strongly chitinized area; anus distant two or three
times its width from caudal margin. — Uganda, Africa on Anona.
gowdeyi Newst.
SPECIES OF TOLLASPIDIOTUS
a. Pygidium with two plates in each third incisura; median pair of
lobes subquadrangular, distal end and angles broadly rounded, as
broad as long, entire; second and third pairs of lobes similar in
size and form; plates subequal in length to lobes, arranged 2, 2, 2, 0;
426 THE COCCIDAE
each lateris irregularly toothed; anus located midway between vulva
and caudal margin, vulva located at cephalic end of pygidium;
mesospiracerores 5-6; preabdomen with lateral margins of segments
produced, tuberculate; each bearing few setae; scale subcircular,
exuviae central, caudal end elevated by ventral scale. — Mauritius on
palm trees. mauritianus Newst.
aa. Pygidium with three plates in each third incisura; median pair of
lobes spatulate, proximal portion constricted, distal end bluntly
rounded, longer than broad; second and third pairs of lobes similar
in form and size; plates subequal in length to lobes, arranged 2, 2,
3, 0; each lateris with five large lobe-like unsymmetrical projec-
tions with serrate margins cephalad of third lobe; setae of lateres
very long and slender, much longer than lobes, others minute. —
Furcaspis rufa Lindgr. — Island of Reunion, St. Denis on Erythroxy-
Ion. rufus Lindgr,
SPECIES OF CRYPTASPIDIOTUS
Pygidium with median pair of lobes large, broader than long, entire,
broadly truncately rounded; second pair of lobes smaller, narrower,
entire, truncately rounded; pectinae subequal in length to lobes, ar-
ranged 2, 2, 3, distapectinae; anus apparently distant from caudal mar-
gin; spiracerores wanting; lateres crenulate. — Algeria on Juniperus,
Callitris. mediterraneus Lindgr.
SPECIES OF MORGANELLA
Pygidium with pectinae on each lateris; anus located near proximal ends
of lobes; median pair of lobes long, longer than broad, pointed, adja-
cent, deep lateral notch; pectinae broad unapectinae, small or long
adjacent to median lobes, successively broader cephalad, much longer
than lobes, sometimes arranged in three groups, arranged 0, 13; setae
adjacent to median lobes shorter than lobes, others much longer than
pectinae. — Morganella maskelli Ckll. — Brazil, Hawaiian Islands, Mauri-
tius, Ceylon, south Africa, Demerara on Michelia, Camellia, Artocarpus,
Cupania, Lagerstromia, mango, orange, papaw longispina Morg.
SPECIES OF PSEUDOTARGIONIA
Pygidium with median pair of lobes entire, distant, truncate with round'
ed angles, longer than wide; second pair of lobes adjacent to median,
smaller, distinct lateral notch on distal end, bluntly rounded; third pair
of lobes rudimentary; plates not prominent, arranged 0, 0, 2-3, 0,
irregular in length; lateres thickened, crenulate, bearing 5-6 setae,
twice as long as median lobes; dorsal oraceratubae in three rows on
each side; anus adjacent to median pair of lobes; mesal part of dorsal
surface of pygidium between dorsal oraceratubae distinctly thickened;
spiracerores 5-6, large; head and thorax with 8-10 long slender setae,
— ^Aonidia glandulosa Newst. — Upper Egypt on Acacia
glandulosa Newst.
SPECIES OF CRENULASPIDIOTUS
Pygidium with median pair of lobes subquadrangular, not strongly pro-
jecting, mesal and lateral margins parallel, distal margin crenulate;
TRIBE ASPIDIOTINI 427
second pair of lobes bluntly rounded, crenulate; third and fourth pairs
of lobes bluntly pointed, crenulate; plates arranged 2, 2, 3, 3, 3, those
of second incisurae described as single deeply divided plate, those of
median incisura similarly described and more than twice as long as
lobes, all other plates short; anus very distant from caudal margin. —
Venezuela on Bletia. portoricensis Lindgr.
SPECIES OF SEPARASPIS
Pygidium with median pair of lobes twice as long as wide, entire, distal
end bluntly rounded, sides parallel; second, third, and fourth pairs
of lobes similar in form and subequal in size to median pair, entire;
plates subequal in length to lobes, distal ends blunt, arranged 2, 2, 3, 0;
each lateris with four adjacent, entire, lobe-like projections, caudal
one largest, others successively smaller; dorsal oraceratubae few in
number; anus distant from caudal margin. — Transvaal, Africa on
Protea, Faurea. proteae Brain.
SPECIES OF BRAINASPIS
Pygidium with median pair of lobes small, quadrangular, mesal and lat-
eral notches, about as broad as long; second pair of lobes large, much
longer and broader than median pair, elongate, longer than broad,
rounded, entire; third pair of lobes smaller than median, triangular,
bluntly rounded, faint lateral notch; pectinae distinctly lacinate, ar-
ranged 2, 2, 2, 2, those of median and second incisurae longer than
median lobes, distinctly shorter than second pair of lobes, those of
third incisurae as long as second pair, those of each lateris as long
as third pair; anus distant from caudal margin; spiracerores wanting.
— South Africa on Andropogon. kellyi Brain.
SPECIES OF CRYPTOPHYLLASPIS
a. Pygidium with four or five unapectinae on each lateris.
b. Pygidium with margin of each second and third incisura pro-
duced into short prominent point; median pair of lobes larger
than others, constricted toward proximal end, mesal and lateral
notches; second and third pairs of lobes elongate, bluntly pointed,
third pair usually with a lateral notch; pectinae deeply lacinate,
arranged 2, 2, 3, 4-5; oraceratubae numerous, tendency to ar-
range themselves in transverse rows; insects forming and occupy-
ing small pits on underside of leaves. — Ceylon on Strobilanthes
putearia Green,
bb. Pygidium v/ith margin of each second and third incisura concave,
not produced.
c. Pygidium with proximal portion not constricted, lateral mar-
gins uniformly gradually converging toward caudal end; me-
dian pair of lobes much larger than others, long and promi-
nent, mesal and lateral notches; second and third pair of lobes
bluntly pointed, elongate, lateral notches; pectinae deeply lac-
inate, arranged 2, 2, 3, 4-5; oraceratubae few in number, not
arranged in transverse rows; insects forming galls upon upper
surface of leaves. — Ceylon on Grewea. occulta Green.
428 THE COCCIDAE
cc. Pygidium with cephalic portion abruptly constricted so that
subparallel lateral margins of portion caudad of constriction
meet cephalic portion at an angle; pectinae and lobes like
those of occultus; oraceratubae few in number; pectinae long
and deeply lacinate; insects forming cylindrical galls twice
as long as wide, constricted at base, ends often studded with
irregular tubercles, upon lower surface of leaf. — Aspidiotus
(Cryptophyllaspis) occultus elongatus Green. — Ceylon on
Grewea. elongata Green.
aa. Pygidium with three pectinae on each lateris.
b. Pygidium with three deeply lacinate furcapectinae on each lat-
eris; median pair of lobes with slight mesal and lateral notches;
three pairs of lobes not darkened; pectinae narrow and strongly
fringed, each lateris with three double pectinae each having ap-
pearance of two squames (plates?) joined at proximal end; in-
cisurae with densariae; dorsal oraceratubae not numerous, ar-
ranged in two rows on each side; setae small. — Bismarck Archi-
pelago on Codiaeum. ruhsaameni Ckll.
bb. Pygidium with three deeply lacinate unapectinae on each lateris;
median pair of lobes elongate, deep mesal and lateral notches,
distal margin bluntly pointed; second pair of lobes much smaller,
unsymmetrical, oblique, margin with deep lateral notch or cren-
ulate; third pair of lobes irregular, pointed, notched; pectinae
deeply lacinate, much longer than median pair of lobes, arranged
2, 2, 3, 3-5; dorsal oraceratubae distinct; insects forming small
conical galls on leaves. — Canary Islands, Maderia on Globularia...
hornmuUeri Riibs.
SPECIES OF SPINASPIDIOTUS
a. Body with cephalic margin entire, not thickened and regularly cren-
ulate.
b. Pygidium with median pair of lobes distinctly notched,
c. Pygidium with five to seven pectinae on each lateris.
d. Pygidium with second pair of lobes distinctly shorter than
median pair.
e. Pygidium with median pair of lobes quadrangular, longer
than broad, distal end truncately rounded, mesal and lat-
eral notches; second pair of lobes much smaller, elongate,
pointed, lateral notch; third pair of lobes much smaller
than second, pointed, triangular, lateral margin toothed;
pectinae long, narrow, subequal in length to median lobes,
deeply lacinate, arranged 2, 2, 3, 6-7, those of each lateris
adjacent to third lobe. — South Africa on Parinarium,
Strychnos. fissidens Lindgr.
ee. Pygidium with median pair of lobes shorter than in type,
second pair broader, third pair of lobes broader, short,
pointed, entire or with more teeth on oblique margin
toward distal end; pectinae with finer lacinations. — Ger-
man East Africa, Mozambique on Bosquiea, Sideroxylon,
Chrysophyllum. fissidens pluridentatus Lindgr.
TRIBE ASPIDIOTINI 429
dd. Pygidium with second pair of lobes as long as or longer than
median pair, subequal in width, similar in shape, lateral
notch; median pair of lobes large, conspicuous, close to-
gether, mesal and lateral margins nearly parallel, distal mar-
gin rounded, slight lateral notch; third pair of lobes about
one-half size of second pair, subsimilar in shape; pectinae
subequal in length to lobes, arranged 2, 2, 3, 5-6, three caudal
pectinae of each lateris deeply lacinate, others simple; dorsal
oraceratubae present, promiscuously arranged; usually
wanting, sometimes with a single ceroris on each side; anus
six times its width from caudal margin and twice its width
caudad of vulva. — Samoan Islands on cocoanut husks
pangoenensis Doane & Ferris,
cc. Pygidium with three pectinae on each lateris; median pair of
lobes prominent, distinct lateral notch, mesal notch wanting,
distal end bluntly rounded; second pair of lobes smaller, irreg-
ular, pointed, lateral notch; third pair of lobes small, triangu-
lar, pointed; pectinae subequal in length to lobes, arranged
2, 2, 3, 3, all except those of median incisura broad, irregu-
larly lacinate; each lateris crenulate, bearing two minute
plates or furcapectinae. — Dutch East Africa on Furcraea
furcraeicolus Lindgr.
bb. Pygidium with median pair of lobes entire.
c. Pygidium with pectinae or plates on each lateris.
d. Pygidium with third pair of lobes distinctly smaller than
median pair.
e. Pygidium with median pair of lobes quadrangular, short,
much broader than long, entire, truncately rounded; sec-
ond pair of lobes minute, as long as broad, entire, bluntly
rounded; third pair of lobes minute, sharply pointed,
longer than broad; pectinae broad, distinctly longer than
median pair of lobes, numerous deep lacinations, arranged
2, 2, 3, 4, those of each lateris adjacent to a third lobe,
lacinations deeper and teeth much longer than those of
other pectinae. — Abyssinia on Euphorbia fissus Lindgr.
ee. Pygidium with median pair of lobes large, longer than
broad, projecting, parallel, unsymmetrical, entire, distal
end broadly rounded; second pair of lobes almost as broad
as long, oblique, rounded, margin with several notches;
third pair of lobes similar to second, smaller; pectina
or plates subequal in length to median pair, dagger-
shaped plates or furcapectinae, arranged 2, 2, 3, 3. —
Sardina, Spain on Ephedra. ephedrarum Lindgr.
dd. Pygidium with three pairs of lobes subequal, similar in
shape, entire or finely crenulate, slightly constricted on
proximal portion; pectinae subequal in length to lobes, ar-
ranged 2, 2, 3, 3, those of incisurae subequal in width, dista-
pectinae, those of each lateris toothed lobe-like projections;
lateres with several angular or tooth-like projections; setae
except cephalic one of each side, subequal in length to lobes,
430 THE COCCIDAE
cephalic setae two or three times length of others; surface
of pygidium coarsely and irregularly reticulate; anus locat-
ed considerably cephalad of middle of pygidium; scale of
adult female coarsely transversely striate, exuviae lateral. —
South-West Africa. reticulatus Newst.
cc. Pygidium without pectinae or plates on lateres.
d. Pygidium with median pair of lobes broader than long.
e. Pygidium with pectinae tridentate, arranged 2, 2, 3, 0,
those of incisurae broad, truncate, not deeply lacinate,
subequal in length to lobes; median pair of lobes quad-
rangular, broader than long, entire, truncate with round-
ed angles; second pair of lobes similar to median, sub-
equal in size; third pair of lobes similar to second, broad-
er than long, shorter than second, truncate; lateres broad-
ly indented and crenulate. — Furcaspis oceanica Lindgr. —
Jaluit Islands on fruit hulls of Cocos oceanicus Lindgr,
ee. Pectinae bidentate; median pair of lobes broader than
long, entire, roundly truncate; second pair of lobes sub-
equal to median, similar in form; third pair of lobes sim-
iliar in form to second, much smaller; pectinae subequal
in length to lobes, not deeply lacinate, bidentate, arranged
2, 2, 3, 0; lateres with two prominent teeth, irregular pro-
jections between third lobe and caudal tooth, three oblique
projections between two teeth, oblique margin serrate. —
Furcaspis haematochroa Ckll. — Batbatan Island. Philip-
pine Islands on cocoanut palms haematochrous Ckll.
dd. Pygidium with median pair of lobes longer than broad,
truncate, entire; second pair of lobes similar to median,
broader; third pair of lobes broader than long, truncate,
expanded at proximal end; pectinae slightly shorter than
lobes, truncate, not deeply lacinate, arranged 2, 2, 2, 0; lat-
eres indented, irregularly produced with irregular projec-
tions.— Aspidiotus cladii Charmoy. — Mauritius on palms
charmoyi Brain.
aa. Body with cephalic margin thickened and regularly crenulate;
pygidium with median pair of lobes large, quadrangular, distal end
rounded, deep lateral and mesal notches; second pair of lobes
longest on mesal margin, bluntly pointed, broad lateral notch; third
pair of lobes as long as second, narrower, pointed, faint notches;
pectinae long, narrow, deeply lacinate, subequal in length or slightly
longer than median, arranged 2, 2, 3, 4, projections of each lateris
with short bluntly truncated, evenly distributed plates. — South Africa
on Dichapetalum. maeandrius Lindgr.
SPECIES OF RHIZASPIDIOTUS
Pygidium with median pair of lobes almost contiguous at distal end,
proximal ends distant, subtruncate, distal end notched, concave or al-
most truncate and broadly rounded on lateral margin, proximal por-
tion of lateral margin notched; second and third pairs of lobes small,
incised, each portion tooth-like projection; lateres crenulate or notched;
TRIBE ASPIDIOTINI 431
second incisurae broad; plates shorter than second pair of lobes, ar-
ranged 0, 2, 2, 0; setae small; oraceratubae small, numerous; anus
longer than broad, six times its width from caudal margin. — Kansas-
on roots of Helianthus. helianthi Parrott.
SPECIES OF TRUNCASPIDIOTUS
Pygidium of adult female with three pairs of lobes, similar in size,
stout, prominent, truncately rounded, entire; some of lobes some-
times missing; pectinae broadly truncated, without pectinations, ar-
ranged 2, 2, 3, 0, resembling lobes; each lateris thickened and coarsely
dentate; paraphyses wanting; dorsal oraceratubae few in number;
ventral aspect of preabdomen on each side with four to six short,
stout, conical setae inserted in calices with thickened rims; head and
thorax on each side with row of long stout submarginal setae. — Algoa
Bay, Cape Colony, Transvaal, Africa on Aloe capensis Walker,
SPECIES OF VARICASPIS
Scale of adult female elongate, sides compressed, meson rather sharply
keeled, exuviae central, superimposed, secretionary portion broad, at-
tached to edge of leaf with equal portions on each side; body with
dorsum strongly convex, venter flat ; rostrum located nearly midway
between cephalic end of body and caudal end of preabdomen, far
caudad of cephalic margin; body densely chitinized except pygidium;
pygidium with median pair of lobes quadrangular, distinct mesal and
lateral notches, as broad as long, bluntly rounded between notches;
second pair of lobes smaller than median, longer than broad, distinct
lateral notch; third pair of lobes minute, distinct, lateral notch; pecti-
nae or plates subequal in length to lobes, arranged 2, 2, 3, 3-4, one next
to third lobe a pectina, others plates ; vulva located near middle of pygi-
dium and anus almost half way between vulva and caudal margin. —
Uganda, Africa on Coffea. fiorineides Newst.
SPECIES OF GONASPIDIOTUS
a. Body with cephalic margin entire and never uniformly crenulate.
b. Pygidium with two pectinae or plates in median incisura.
c. Pygidium with pectina on each lateris, median pair of lobes
with mesal and lateral notches, subquadrangular, prominent,
broadly rounded; second pair of lobes about one-half size of
median, bluntly pointed between mesal and lateral notches;
pectinae subequal in length to lobes, narrow distapectinae, ar-
ranged 2, 2, 3; each lateris with three spine-like projections
equidistant from each other and a second lobe, one adjacent to
pectinae of lateres. — Italy on Quercus minimus Leon.
cc. Pygidium without pectinae or plates on lateres.
d. Pygidium with setae small, subequal in length to median
pair of lobes; median pair of lobes broader than long, mesal
margins parallel, distal end bluntly rounded, lateral
margin oblique, lateral notch; second pair of lobes similar
to median, smaller, lateral notch; second incisurae normal
in width, plates arranged 2, 2, 0, short, much shorter than
432 THE COCCIDAE
lobes; each lateris with five rounded undulations; marginal
ceratubae prominent; anus five times its width from caudal
margin and three times from vulva. — California on Cupressus,
Sequoia. shastae Colm.
dd. Pygidium with setae large, longer than median pair of lobes,
four on each side; median pair of lobes with lateral notches
only, rarely with mesal notch, subquadrangular, parallel or
somewhat convergent, longer than broad, symmetrically
bluntly rounded; second pair of lobes two-thirds size
of median, subadjacent to median, oblique, broadly rounded,
deep lateral notches; plates arranged 2, 1-2, 0, those of me-
dian incisura small, dagger-shaped, two, those of second in-
cisura sometimes furcapectinae, one to two; setae large,
longer than median lobes, four on each side. — Madeira on
Laurus. laurinus Lindgr.
bb. Pygidium without plates in median incisura, plates in second in-
cisurae, and pectinae on lateres, arranged 0, 2, 2; median pair of
lobes large, prominent, rounded, crenulate; second pair of lobes
small, nearly as broad as median, mesal margin straight, lateral
margin oblique, crenulate, lobes inconspicuous; dorsal oracera-
tubae arranged in four indefinite rows on each side; anus five
times its width from caudal margin and same distance from vulva;
lateres entire; setae small; mesal calles separate, distance be-
tween them subequal to distance between each mesal and lateral
callis, latter oblique. — Targionia graminellus Ckll. — New Mexico,
Colorado on ?Hilaria, Bouteloua graminellus Ckll.
aa. Body with cephalic margin thickened and uniformly crenulate;
pygidium with median pair of lobes quadrangular, broad with deep
lateral and mesal notches, distal margin bluntly rounded; second
pair of lobes much smaller, pointed, large lateral notch; pectinae
subequal in length to median lobes, deeply lacinate, arranged 2, 2,
10-11, each lateris with three pectinae adjacent to second lobe and
7-8 short bluntly truncated plates distributed along margin of lateris.
— This species is similar to Spinaspidiotus maeandrius Lindgr. —
South Africa on Acioa, Strychnos. undulatus Lindgr,
SPECIES OF PARANEWSTE'ADIA
Pygidium with two pairs of lobes, median pair minute, rounded; second
pair of lobes three times as long as median pair, slightly constricted at
proximal end, spatulate, lateral notch; pectinae and plates present, ar-
ranged 0, 2, 7-8, those of each lateris consisting of two long simple spat-
ulate plates and 5-6 unapectinae, second incisura with plates. — Africa...
maculata Newst.
SPECIES OF CHORIZASPIDIOTUS
a. Pygidium with median pair of lobes entire; quadrangular, truncate;
second and third pairs of lobes represented by slight deeply emargi-
nate projections, producing a crenulate appearance; plates and
pectinae wanting in median incisura and on lateres, arranged 0, 2,
0. — New Mexico on Guterrezia guterriziae Ckll. & Parr.
aa. Pygidium with median pair of lobes notched.
TRIBE ASPIDIOTINI 433
b. Pygidium with seven or eight or more pectinae or plates on each
lateris.
c. Pygidium with plates on lateres.
d. Pygidium with median pair of lobes with prominent mesal
and lateral notches, quadrangular; plates wanting in median
incisura, arranged in two groups of six each just cephalad of
each median lobe, separated by small marginal prominence;
each lateris cephalad of median lobe thickened for short dis-
tance, irregularly coarsely crenulate; anus distant from
caudal margin; spiracerores wanting. — Ceylon from galls of
Amorphococcus mesuae. cuculus Green.
dd. Pygidium with each median lobe only with prominent lat-
eral notch.
e. Pygidium with pectinae and plates of lateres one-fourth
longer than lobes; median pair of lobes subquadrangular,
prominent, projecting, longer than broad, adjacent, lat-
eral margin straight to convex, prominent lateral notches
only, one or two; each lateris with 5-7 plates, very long,
much longer than lobes, curved, located adjacent to lobes
in positions for incisurae on lateres; setae long and slender,
as long as plates; anus large, slightly over twice its width
from caudal margin and about three times from vulva;
incisurae may have densariae. — Diaspis (Epidiaspis) con-
spicua Brain. — South Africa on Acacia, Gardenia, privet—
conspicus Brain,
ee. Pygidium with pectinae or plates of lateres subequal to or
shorter than lobes, usually with plates, at most furca-
pectinae; anus four times its width from proximal ends
of lobes; median pair of lobes quadrangular, mesal mar-
gins adjacent, prominent lateral and faint distal notches;
plates all shorter than lobes, stout, some rarely furcate,
placed adjacent to lobes; setae slightly longer than lobes,
five on each side; calles distinct. — Targionia celtis Her-
rick. cueroensis Ckll.
cc. Pygidium with pectinae on each lateris; median pair of lobes
large, subquadrangular, longer than wide, single mesal and
two or more lateral notches; each lateris with eight to ten
furcapectinae and plates intermixed, very long and slender,
some longer than lobes, located adjacent to lobe; lateres finely
serrate cephalad of pectinae; setae of two lengths, one lot long,
nearly as long as lobes, other lot minute, not projecting beyond
margin. — South Africa on Euphorbia spiniger Lindgr.
b. Pygidium with three or four pectinae or plates on each lateris.
c. Pygidium with anus ten times its width from caudal margin;
vulva more distant from anus than anus from caudal margin;
median pair of lobes subquadrangular, slight lateral notch,
distal end angular, as broad as long, prominent, projecting;
pectinae shorter than median pair of lobes, three on each lat-
eris, latapectinae, wanting in median incisura; setae as long
434 THE COCCIDAE
as lobes; dorsal oraceratubae few, not in longitudinal rows. —
Australia on Casuarina. Hdens Green.
cc. Pygidium with anus four times its width from caudal mar-
gin; median pair of lobes large, elongate, convergent, lateral
margin oblique, two deep notches; plates, rarely furcapectinae,
six on each lateris, arranged in two or three groups of two
plates each, one near each median lobe, two short distance
cephalad, third still farther cephalad, wanting in median in-
cisura; lateres crenulate; setae long and stout, as long as or
longer than lobes. — Canary Islands on Argyranthemum
canariensU Lindgr.
SPECIES OF REMOTASPIDIOTUS
«" Pygidium with median pair of lobes short, not projecting much be-
yond general contour of pygidial margin, close together, distal
ends practically contiguous, broad and obliquely truncate; pectinae
apparently wanting, two minute plates adjacent to lobe; each lateris
bluntly serrulate, small projection near lobe, four much larger prom-
inent ones at considerable distance cephalad of caudal projection;
anus oval, about six times its width from caudal margin and four
times from vulva; surface striate with rows of small round dorsal
oraceratubae. — French Bay, Bruce Peninsula, Shores of Lake Huron,
Canada on Arctostaphylos and California on Grindelia, Eriophyllum,
Corethrogyne. dearnessi Ckll.
a,a. Pygidium with median pair of lobes short, not projecting beyond
general contour of pygidial margin, contiguous, distal ends oblique
with mesal angle produced; second and third pairs of lobes want-
ing; each lateris serrate with two inconspicuous rounded projec-
tions located cephalad where a lobe would be expected; plates lim-
ited to two, short but easily distinguished just cephalad of each
median lobe; setae normal; anal opening large, four times its width
from caudal margin; dorsal surface striate with rows of oracera-
tubae, much less conspicuous than in preceeding species. — New South
Wales on Chenopodium. chenopodii Marl.
SPECIES OF CHENTRASPIS
Pygidium with median pair of lobes fused into single rectangular lobe,
twice as broad as long, margin notched; pectinae of each lateris ar-
ranged in three groups near lobe, arranged 2-2-1, separated by indenta-
tions, unapectinae; setae stout, nearly as long as pectinae; each lat-
eris cephalad of pectinae finely striate or undulate. — Australia on
Acacia. unilobus Mask.
SPECIES OF HEMIBERLESIA
u. Pygidium with pectinae or plates in median incisura.
b. Pygidium with pectinae or plates on each lateris.
c. Pygidium never with more than four, usually two or three,
pectinae or plates on each lateris.
d. Pygidium with unapectinae on each lateris.
TRIBE ASPIDIOTINI 435
e. Pygidium with densariae of second and third incisurae
subequal; median pair of lobes large, quadrangular, mesal
and lateral notches, distant, distal margin bluntly rounded;
second pair of lobes small, spear-shaped, often with lat-
eral notch; third pair of lobes narrow, spiniform, pointed,
entire; pectinae slightly larger than lobes, long slender
unapectinae, arranged 2, 2, 3, 3-4; anus about its width
from proximal ends of lobes. — South Africa
mitchelli Marl,
ee. Pygidium with densariae of second and third incisurae
not subequal; median pair of lobes large, quadrangular,
distinct mesal and lateral notches, distal margin round-
ed between notches; second and third pairs of lobes small,
triangular, attenuate at distal end, pointed; pectinae
longer than median pair of lobes, arranged 2, 2, 3, 3, those
cephalad of each median lobe longer and broader, deeply
lacinate, unapectinae; densariae large, prominent, mesal
margin of third incisurae larger than lateral; setae long;
each lateris near middle with two tooth-like projections;
anus over twice its width from caudal margin; spiracerores
wanting. — Mansa, Africa on Trichilia. elegans Lindgr.
dd. Pygidium with plates or furcapeotinae on each lateris.
e. Pygidium with median pair of lobes large and well de-
veloped.
f. Anus from one to one and one-half times its width dis-
tant from caudal margin.
g. Pygidium with median pair of lobes with mesal and
lateral notches or sinuate, quadrangular, projecting,
mesal margins parallel or slightly convergent, lateral
margins straight or curved; second and third pairs
of lobes minute, bluntly pointed, entire; pectinae ir-
regular, pectinae and plates arranged 2, 2, 3, 2-3,
those of median incisurae furcapectinae and of
other incisurae unapectinae, each lateris with
plates or furcapectinae; anus one and one-half
times its width from caudal margin and over
five times its width from vulva; lateral and
mesal margins of densariae subequal; calles all
separate. — Aspidiotus rapax Comst., Aspidiotus
euonymi Targ., Aspidiotus lucumae Ckll. — United
States, Mexico, West Indies, Brazil, Europe, Hawaiian
Islands, South Africa, Australia, New Zealand,
Ceylon on Camellia, Fuchsia, Eucalyptus, Acacia,
Myoparum, Thamnus, Cercis, Coprosoma, cotton-
wood, walnut, willow, maple, birch, holly, laurel,
olive, fig, almond, orange, lemon, apple, pear, quince,
tea, etc. camelliae Sign.
gg. Pygidium with median pair of lobes with lateral
notch only, lobes prominent, projecting, distal end
bluntly rounded, mesal margins straight, subadjacent;
436 THE COCCIDAE
second pair of lobes minute, bluntly pointed, entire;
third pair of lobes smaller than second, pointed, pec-
tinae and plates subequal in length to median pair of
lobes, arranged 2, 2, 3, 2-3, those of median incisura
and each lateris slender plates, those of second and
third incisurae longest, f urcapectinae ; densariae of
second incisurae larger than those of third, wanting
in median; anus large, about its own width from
caudal margin. — Argentina on Ophryoporus
argentina Leon,
ff. Pygidium with anus large, over three times its width
from caudal margin; median pair of lobes large, quad-
rangular, deep lateral and mesal notches, distal margin
strongly convexly rounded between notches; second pair
of lobes small, rounded, inconspicuous; third pair of
lobes rounded, smaller than second pair, inconspicuous;
pectinae not deeply lacinate, about as long as median
pair of lobes, arranged 2, 2, 3, 2-3, those of third in-
cisurae and of lateres either not deeply lacinate or
plates, one of each third incisura much larger than
others; densariae prominent, those of second incisurae
larger than those of third, mesal and lateral margins
subequal; spiracerores wanting; each lateris on caudal
portion with short plate-like projections; pygidium of
second nymphal female similar to adult. — Tanga, Africa
on Tectona. tectonae Lindgr.
ee. Pygidium with median pair of lobes not well developed,
margin rounded; second pair of lobes represented by sim-
ple lobe of border of pygidium; third pair of lobes spini-
form; pectinae furcapectinae, arranged 2, 2, 3, 2; pectinae
of median incisura but little longer than median lobes,
others successively longer, those of each lateris twice as
long as lobes, formed with mesal fork much longer than
lateral. — France on Psamma provincialis Vayss.
cc. Pygidium with five to seven pectinae on each lateris.
d. Pygidium with densariae in median incisura; median pair of
lobes subquadrangular, longer than broad, entire, truncately
rounded, distant; second and third pairs of lobes much
smaller, short, entire, bluntly rounded; pectinae elongate,
nearly twice as long as median pair of lobes, deeply lacinate,
arranged 2, 2, 3, 5-6, those of each lateris very deeply
lacinate; anus about four times its width distant from caudal
margin; setae of median and second pairs of lobes long,
longer than lobes; densariae of second and third incisurae
typical. — Canary Islands on Euphorbia. taorensis Lindgr,
dd. Pygidium not with densariae in median incisura; median
pair of lobes longer than broad, subadjacent at proximal end,
bluntly rounded, entire; second and third pairs of lobes
large and distinct, smaller than median pair, bluntly pointed,
indistinct lateral notch; pectinae elongate, distinctly longer
TRIBE ASPIDIOTINI 437
than median lobes, deeply lacinate, those of each third in-
cisura and of lateris unapectinae, arranged 2, 2, 3, 6-7, those
of each lateris longer than others; anus over four times
its width distant from caudal margin; setae slightly longer
than lobes; densariae of second and third incisurae typical.
— Canary Islands on Dracaena. .-tinerfensis Lindgr.
bb. Pygidium without pectinae or plates on each lateris.
c. Pygidium with pectinae or plates of second and third incisurae
longer than second and third pairs of lobes,
d. Pygidium with median pair of lobes with distinct mesal and
lateral notches, quadrangular, large, distal end bluntly round-
ed; second pair of lobes similar in shape, about one-half size
of median; third pair of lobes small, pointed; pectinae sub-
equal in length to median pair of lobes, those of second and
third incisurae unapectinae, arranged 2, 2, 3, 0; each lateris
crenulate with two or three distant truncated plate-like pro-
jections; dorsal oraceratubae present. — Abyssinia on Aloe
mammillaris Lindgr.
dd. Pygidium with median pair of lobes entire, large; second
and third pair of lobes much smaller, those of third pair
dentiform; pectinae small and delicate, arranged 2, 2, 2-3, 0;
dorsal oraceratubae numerous; cephalic end of pygidium with
two transverse calles, mesal and lateral evidently fused. —
Algeria on Ephedra. trabuti Marchal.
cc. Pygidium with pectinae or plates of second and third incisurae
notably much shorter than second and third pairs of lobes,
those of third incisurae larger and with some notches, ap-
parently arranged 2, 2, 3, 0; median pair of lobes projecting;
second pair of lobes triangular, mesal side parallel with meson,
lateral side oblique, continuous with margin of pygidium;
third pair of lobes similar in form to second, but smaller and
more obtuse; setae long, extending much beyond lobes; para-
physes wanting; bundle of long and straight ceratubae on each
side of pygidium. — South Tunis on Nitraria. __mfrariae Marchal.
aa. Pygidium without pectinae or plates in median incisura.
b. Pygidium with pectinae or plates on each lateris.
c. Pygidium with pectinae in incisurae.
d. Pygidium with mesal densariae of second incisurae larger
than lateral, its cephalo-lateral angle produced; median pair
of lobes subquadrangular, mesal margins strongly convergent,
lateral and distal margins rounded, prominent lateral notch;
second and third pair of lobes minute, triangular, second
pair much broader than third; pectinae deeply lacinate, ar-
ranged 0, 2, 3, 4-5, those of each lateris and of incisurae una-
pectinae; cephalic end with four calles; setae long and
slender. — Canada, District of Columbia, Mexico, Demerara
on basswood, lilac diffinis Newst.
dd. Pygidium with mesal and lateral margins of densariae of
438 THE COCCIDAE
second incisurae subequal and symmetrical. — Mexico on
"Barenjeno Chiquito." diffinis parrotti Newell.
cc. Pygidium with plates in incisurae; median pair of lobes promi-
nent, nearly as long as broad, mesal margins subadjacent, lat-
eral notch, sometimes wanting; second pair of lobes longer
than broad, sometimes with lateral notch; third pair of lobes
inconspicuous, small, pointed; second and third incisurae with
densariae, mesal margin longest and stouter than lateral;
plates in incisurae small, furcapectinae and one plate on lateres,
arranged 0, 1, 2, 6, those of each lateris arranged 3-1-1-1, plate
at cephalic end, swollen at proximal end; anus about seven
times its width from caudal margin and about twelve times
from vulva; mesal calles fused, distant from lateral calles, each
of latter only about three times as long as broad. — Cuba on
Faberna. fabernii Houser.
bb. Pygidium without pectinae or plates on lateres, arranged 0, 1, 0,
0; median pair of lobes large, broader than long, two mesal and
one lateral notch, bluntly rounded; second and third pairs of
lobes small, crenulate, rounded, inconspicuous; lateres crenulate;
setae long, projecting beyond median pair of lobes; mesal calles
fused and widely separated from lateral calles; anus near caudal
margin; densariae of second and third incisurae distinct, want-
ing between median pair of lobes. Canary Islands on Laurus,
ApoUonias. aonidioides Lindgr.
SPECIES OF COMSTOCKASPIS
a. Pygidium with pectinae on each lateris.
b. Pygidium with median pair of lobes with lateral notch, distal end
broadly rounded, mesal margins not distant; second pair of lobes
minute, only slightly projecting, distal margin broadly emarginate
or with broad lateral notch; plates small, arranged 2, 2, 6, those
of each lateris arranged in two groups, one group associated third
densariae and a group of three pectinae located cephalad of ce-
phalic group of plates. — Aspidiotus perniciosus albopunctatus
CklL, Aspidiotus perniciosus andromelas Ckll. — Canada, United
States, Chili, Hawaiian Islands, Japan, China, Australia, Germany
on apple, pear, peach, quince, apricot, plum, walnut, almond, pop-
lar, chestnut, sumac, catalpa, willow, osage-orange, linden, ash,
elm, dogwood, orange, lemon, elder, maple, rose, currant, raspberry,
pecan, milkweed, spruce, cedar, etc. pemiciosa Comst.
bb. Pygidium with median pair of lobes with mesal and lateral notch.
c. Pygidium with six pectinae on each lateris, pectinae long, well
developed, much longer than setae, arranged 2, 2, 6, those of
each lateris unapectinae, deeply lacinate; median pair of lobes
large, projecting, constricted at proximal end, triangular,
strongly emarginate on lateral margin, small notch on mesal
margin; second pair of lobes subadjacent to median, similar in
form, much smaller, single lateral notch; dorsal oraceratubae
in longitudinal rows. — South Algeria on Zilla,.. seurati Marchal.
TRIBE ASPIDIOTINI 439
cc. Pygidium with nine pectinae on each lateris, pectinae long,
longer than median lobes, deeply lacinate, arranged 2, 2, 9,
those of each lateris unapectinae, not as deeply lacinate as
others, forming group cephalad of second lobe; median pair of
lobes qaudrangular, distal end bluntly rounded, deep lateral and
mesal notches; second pair of lobes longer than broad, deep
lateral and faint mesal notches, bluntly pointed; each side of
pygidium near cephalic portion of lateres with group of 19-32
dorsal oraceratubae. — South Africa on Ehretia, Anacardiaceae,
Illigera, Mitrangune. replicata Lindgr.
aa. Pygidium with three plates on each lateris, plates arranged 0, 2, 3;
median pair of lobes adjacent, large, broad and short, bluntly round-
ed, lateral notch; second pair of lobes short, broad, pointed, subobso-
lete; two pairs of densariae, well formed, pyriform, lateral margin
of each second densaria largest; anus over three times its width
from caudal margin and five times from vulva. — New Mexico, Cali-
fornia on pinus, Libocedrus, Cupressus coniferarum Ckll.
SPECIES OF CHEMNASPIDIOTUS
Pygidium with median pair of lobes distinct, broader than long, mesal
and lateral notches; densariae in second and third incisurae, wanting
in median incisura, subequal; plates arranged 2, 5 to several, those of
each lateris arranged, two in each second incisura, three in third in-
cisura, and several on each lateris; setae longer than lobes; dorsal ora-
ceratubae few; anus twice its width from caudal margin; living in galls
on leaves. — District of Columbia, Georgia, Mississippi on Liquidambar...
Uquidamdaris Kotinsky.
SPECIES OF HENDASPIDIOTUS
a. Pygidium with densariae of second incisurae normal in length, never
greatly elongated.
b.Pygidium with pectinae or plates in median incisura.
c. Pygidium with plates in median incisura.
d. Pygidium with densariae in median incisurae; median pair
of lobes as long as or longer than broad, rectangular, mesal
and lateral margins straight, angles prominent; two plates in
median incisura and five pectinae and three short plates on
each lateris. — Mexico. tricolor Ckll.
dd. Pygidium not with densariae in median incisura.
e. Pygidium with median pair of lobes broader than long,
mesal margins parallel, not converging, quadrangular, dis-
tal end bluntly rounded, distinct lateral and faint mesal
notches; densariae of second incisurae large, subequal in
length, those of third incisurae much smaller, subequal;
plates minute, inconspicuous, arranged 2, 2-1; mesal calles
strongly thickened, near together, distant from oblique
lateral calles; anus large, about twice its own width from
caudal margin. — South Africa on Robinia.__a/ncanws Marl,
ee. Pygidium with median pair of lobes broader than long,
distant, mesal margins converging, rounded, broadest at
440 THE COCCIDAE
middle, edges obscurely crenulate; pectinae large, narrow,
longer than lobes, arranged 2, 11, those of each lateris
close together; anus circular, located near proximal ends
of lobes; second and third incisurae with densariae. — Mex-
ico on Cupressus. cupressi Ckll.
CO. Pygidium with pectinae in median incisura, pectinae distinct,
comb-like, not notched, form of rounded knife-blades, arranged
2, 10, those of median incisura and of each lateris formed and ar-
ranged so as to make perfect contour with lobes; median pair
of lobes large, others wanting; cephalic end of pygidium with
three transverse calles. — Senegal, Africa on Balanites
vuilleti Marchal.
bb. Pygidium without pectinae or plates in median incisura.
c. Pygidium with about eight plates, adjacent to median lobe, on
each lateris; median pair of lobes quadrangular, distal end
truncately rounded, distinct mesal and lateral notches; den-
sariae marking position of second and third incisurae, mesal
margin of densariae of second incisurae much longer than lat-
eral; mesal and lateral calles distinct, subequal in length, eqi-
distant from each other. — Illinois, New York, Kansas, Japan
on elm, Catalpa, Cycas. ulmi Johns.
cc. Pygidium never with eight plates on each lateris adjacent to
third lobe, usually few in number.
d. Pygidium with median pair of lobes large, two prominent
lateral notches, mesal angle of each protuberant and round-
ed, mesal notches wanting; second and third pairs of lobes
marked by densariae; mesal margin of densariae of each sec-
ond incisura but little longer than lateral; much broader, den-
sariae of third incisurae not prominent; plates inconspicuous;
mesal calles subhyaline, inconspicuous, lateral calles dis-
tinct.— Transcaspian Russia on Populus. transcaspiensis Marl,
dd. Pygidium with median pair of lobes large, each with single
prominent lateral notch, mesal margins straight, distant, dis-
tal ends bluntly rounded; plates and pectinae inconspicuous,
not as long as lobes, arranged 0, 2, those of each lateris ar-
ranged 1-1, cephalic one a pectina; densariae large, mesal and
lateral margins subequal; setae large, prominent, longer than
lobes. — not Aspidiotus alni Marchal. — Austria on Quercus
alni Lindgr.
aa. Pygidium with densariae of second incisurae with mesal side greatly
elongated, straight, enlarged at apex, club-shaped, lateral margin
shorter and only slightly enlarged, mesal margin inversely carrot-
shaped; densariae of third incisurae subequal; median pair of lobes
with two small proximal processes; median pair of lobes rather long,
rounded, sometimes notched; second pair of lobes usually slightly
doubly concave; setae fairly large; plates minute, spine-like, hardly
to be seen, not detectable according to Newell; anus near caudal
margin, transverse; dorsal oraceratubae present. — Mexico on Caesal-
pina. subsimilis Ckll.
TRIBE ASPIDIOTINI 441
SPECIES OF CLAVASPIS
a. Pygidium with median pair of lobes with mesal and lateral notches,
prominent, lateral margin oblique, adjacent, distal ends rounded;
densariae prominent, mesal margin of those of second incisurae
over three times as long as lateral margin, cephalic end greatly
enlarged, rounded, ball-like, densariae of third incisurae small, nor-
mal in form; plates small, inconspicuous, arranged 0, 8-9, those of
each lateris arranged 2-2-2-2 or 3, two in second incisurae, two in
third incisurae, two just cephalad of third incisurae, and two or
three farther cephalad; setae long, slender, about as long as lobes;
anus longer than broad, distant five times its width from caudal
margin and sixteen or more times from vulva. — Aspidiotus subsimi-
lis anonae Houser. — Florida, Peru on Mangifera, Anone
anonae Houser.
aa. Pygidium with median pair of lobes with prominent lateral notch,
large, subadjacent, distal ends bluntly rounded; second pair of lobes
small, inconspicuous; densarie of second incisurae large, prominent,
those of mesal margin much broader, three or more times as long
as lateral, rounded, club-shaped, those of third incisurae subequal
in length, small, swollen; plates small, inconspicuous, arranged
0, 2, 3, those of each lateris arranged in two groups, one in third
incisurae and two much shorter ones some distance cephalad; anus
distant about twice its width from caudal margin; setae longer than
plates. — Society Islands. herculeana Doane & Hadden.
SPECIES OF PELOMPHALA
a. Pygidium with nine pairs of paraphyses; body with cephalic portion
on each side with tooth-like projection.
b. Pygidium with median pair of lobes with lateral notch, narrow
in form, pointed, near together, not touching; second, third, and
fourth pairs of lobes broader than median, similar, except second
and third have mesal and lateral notches, fourth serrate; each
lateris crenulate, with about five small prominences; plates short
and inconspicuous; paraphyses in nine pairs, seven short and two
long pairs, short ones considerably longer than lobes. — Chrysompha-
lus calurnus Ckll. — Mexico on Crataegus. calurna Ckll.
bb. Pygidium with median pair of lobes with distinct mesal and lat-
eral notches, quadrangular, broadly rounded between notches,
mesal and lateral margins parallel, distant; second, third, and
fourth pairs of lobes with mesal margin longer than lateral, distal
margin oblique, notched, second pair broader than median,
others successively broader; each lateris coarsely, irregularly
toothed; plates shorter than lobes, inconspicuous, arranged 0, 0,
2, 2, 0; paraphyses in nine pairs, sixth pair short and third pair
long, second, fifth, and eighth pairs long, third and ninth pairs
subequal, slightly shorter than subequal first and sixth pairs and
slightly shorter than fourth; anus located near middle of pygi-
dium; calles continuous, united with lateral thickenings; body
with margin more or less strongly and finely crenulate. — British
Guiana on Avicennia. apicata Newsi.
442 THE COCCIDAE
aa. Pygidium with four pairs of paraphyses; lobes in four pano, median
pair entire, second pair with single notch, third pair minutely ser-
rate with four notches, and fourth pair broad and serrate; each lat-
eris with three small prominences; anus located considerable dis-
tance from caudal margin. — New Mexico on Quercus lilacina Ckll.
SPECIES OF MYCETASPIS
Body with broad, rotund, cephalic, constricted, head-like area; median
pair of lobes triangular, entire or faintly notched; second pair of lobes
broad, shorter than median, one or more notches; third pair of lobes
as broad as second, distal margin dentate; plates short, rudimentary,
arranged 1, 2, 2, 3, those of each lateris not longer than third lobe,
one of median incisura deeply bifurcate; each lateris with three broad,
serrate, lobe-like projections on caudal half; paraphyses arranged in
four pairs, first pair longest, associated with second incisurae, second
pair with third incisurae, slightly shorter than first, third and fourth
pairs near together, associated with cephalic end of third lobe. — West
Indies, Mexico, Guiana, England on orange, Areca, Sabol, Jasminum,
plantain, banana, mango, fig, Tillandosia, Anacardium
personata Comst.
SPECIES OF PARAONIDIELLA
Pygidium with median pair of lobes narrow, bluntly rounded, mesal and
lateral margins subparallel; second pair of lobes much broader than
long, deeply incised, mesal lobelet larger than lateral, lobelets truncate-
ly rounded; third pair of lobes short, wider than mesal lobelet, sub-
truncate; pectinae short, broad distapectinae, not projecting beyond
lobes, arranged 2, 2, 2, 0; each lateris irregularly indented with very
large prominent seta near middle of its length; paraphyses arranged
in eight pairs, eighth pair longest, third pair next in length, others
much shorter. — Australia, Natal, south Africa on Cladium, Lepido-
sperma, Xerotes, sedges, aloe cladii Mask.
SPECIES OF AONIDIELLA
a. Pygidium with seven or eight pairs of paraphyses.
b. Pygidium with nine pairs of paraphyses, second, fifth, and ninth
pairs subequal and longest, seventh pair one-fourth shorter than
ninth and next in length, first, third, sixth, and eighth pairs sub-
equal and slightly shorter than seventh, fourth pair one-third short-
er than third and shortest; median pair of lobes broader than long,
rounded, slightly oblique; second and third pairs of lobes similar,
subequal, lateral margins oblique, crenulate; lateres with lobe-
like projections; pectinae subequal in length to lobes, furcapec-
tinae, arranged 1, 2, 2, 2, those of each lateris adjacent to third
lobe. — South Africa on Acacia (not Mimosa) phenax Ckll.
bb. Pygidium with less than nine pairs of paraphyses.
c. Pygidium with eight pairs of paraphyses, second pair one-half
length of first, slightly longer or subequal to second, third,
fifth, sixth, and eighth pairs longest and slightly longer than
subequal first and seventh; median pair of lobes rounded, one
or more lateral notches; second pair of lobes longer than
TRIBE ASPIDIOTINI 443
broad, lateral margin strongly oblique, dentate, mesal angle
bluntly rounded; third pair of lobes similar to second pair;
pectinae short, inconspicuous furcapectinae or plates, arranged
0 2, 2, 2; each lateris with indentations. — Mexico on Mimosa
mimosae Comst.
cc. Pygidium with seven pairs of paraphyses, second pair longer
than first or third, fourth and sixth pairs longest, slightly
longer than second, fifth pair slightly shorter than second and
longer than first and seventh; median pair of lobes subquad-
rangular, mesal and lateral notches; second pair of lobes
smaller, subquadrangular, mesal and lateral notches; third pair
of lobes broader than long, bluntly pointed, lateral margin
crenulate; lateres irregularly toothed and crenulate; pectinae
comparatively broad distapectinae, arranged 2, 2, 2, 2, those of
each lateris furcapectinae. — District of Columbia, Georgia, Cali-
fornia on maple, apple tenedricoa Comst.
. Pygidium with six or less pairs of paraphyses.
b. Pygidium with five or six pairs of paraphyses.
c. Pygidium with second and third pairs of paraphyses subequal
and longer than fourth, firs,t, fourth, and sixth pairs subequal,
all short,
d. Scale of adult female reddish or brownish; median pair of
lobes rounded, mesal and lateral notches; second pair of lobes
similar in shape, mesal and lateral notches or lateral mar-
gin oblique and crenulate; third pair of lobes similar, lateral
and sometimes mesal notches; pectinae subequal in length
to lobes, median and second incisurae with narrow dista-
pectinae, others unapectinae, arranged 2, 2, 3, 3; calles four
in number. — New York, Florida, Ohio, California, West
Indies, Hawaiian Islands, Fiji, Samoa, New Caledonia,
New Zealand, Australia, Japan, China, Ceylon, Mauritius,
south Africa, Syria, southern Europe on orange, lemon,
cocoanut, fig, olive, agave, plum, rose, pear, quince, apple,
willow, oak, grape, etc. aurantii Mask.
dd. Scale of adult female always yellowish in color. Insects on
leaves of fruit only. — California, Japan on Euonymus,
Aucuba, orange. aurantii citrina Coq.
cc. Pygidium with second pair of paraphyses longer than third
and distinctly shorter than fourth, longest, fifth and sixth
pairs subequal or shorter than third, first and third pairs sub-
equal; median pair of lobes bluntly rounded, mesal and lateral
notches, broader than long; second pair of lobes narrower
than median, deep lateral notch; third pair of lobes smaller
than second, bluntly rounded, lateral notch distinct; pectinae
arranged 2, 2, 3, 6, those of lateres unapectinae, others dista-
pectinae.— Italy on Taxus taxa Leon,
bb. Pygidium with less than five pairs of paraphyses.
c. Pygidium with four pairs of paraphyses.
444 THE COCCIDAE
d. Pygidium never with long fusiform mesal paraphyses ex-
tending to and sometimes cephalad of anus.
e. Pygidium with second pair of paraphyses longest, twice
as long as third, shortest, first and fourth pairs interme-
diate and subequal; median pair of lobes broadly rounded,
broader than long, entire; second and third pairs of lobes
broader than median, short, emarginate or irregularly
notched; plates short, not longer than lobes, arranged 2,
2, 2, 2; lateres crenulate. — England on pine-apple
bromeliae Leon,
ee. Pygidium with first and second pairs of paraphyses sub-
equal and twice as long as subequal third and fourth
pairs; paraphyses large, conspicuous, knobbed at cephalic
end; median pair of lobes triangular, adjacent, bluntly
pointed, lateral margin longest, deeply notched; second
pair of lobes triangular, minute, entire; third pair of lobes
similar, smaller; pectinae subequal in length to median
pair of lobes, arranged 2, 2, 3, 2-3; setae long, as long as
pectinae; anus small, over five times its width from caudal
margin; mesal calles fused, lateral calles short and dis-
tant from mesal. — Northern Australia on Ficus
subcuticularis Green,
dd. Pygidium always with long fusiform mesal paraphysis ex-
tending to or beyond anus, other paraphyses situated, two
on median lobes, mesal two-thirds length of lateral, one more
or less club-shaped, much longer and stouter than first two,
situated just laterad of median lobe, and similar one at mesal
side of proximal end of second pair of lobes; median pair of
lobes quite broad, united except distal third, distinct lateral
notch; second pair of lobes scarcely broader, oblique, mi-
nutely serrate, bearing about five teeth, separated from me-
dian lobe by space equal to width of both median lobes; third
pair of lobes about two-thirds width of second, similar in
shape, serrate with about three teeth; furcapedtinae ar-
ranged 1, ?5, 3, ?, and several plates of varying length on
lateres; thorax separated from abdomen by lateral inden-
ture.— India. minuta Kotinsky.
cc. Pygidium with less than four pairs of paraphyses.
d. Pygidium with three pairs of paraphyses.
e. Pygidium with pectinae on each lateris, pectinae subequal
in length with lobes, arranged 2, 2, 3, 3, those of each
lateris unapectinae, longer than lobes; median pair of
lobes elongate, constricted on proximal portion, distal por-
tion bluntly rounded, florate, faint mesal and lateral
notches; second and third pairs of lobes similar, third
pair slightly smaller; all three pairs of paraphyses sub-
equal, on mesal side of proximal end of each lobe. — Ceylon
on Phyllanthus. taprohana Green.
ee. Pygidium without pectinae or plates on each lateris.
TRIBE ASPIDIOTINI 445
f. Pygidium with three pairs of lobes; median pair of lobes
narrow, rounded; second and third pairs of lobes broader
than long, lateral margin oblique, notched; pectinac hot
longer than lobes, furcapectinae, arranged 2, 2, 2, 0;
lateres irregularly dentate; paraphyses with second and
third pairs subequal in length and two or more times
as long as first pair. — Massachusetts, Galapagos Islands
on smilax, Croton. smilacis Comst.
ft. Pygidium with four pairs of lobes; median pair of lobes
somewhat triangular, distal end with distinct point,
distinct mesal and lateral notches; second pair of lobes
narrower than median, longer than broad, convergent
caudad, distal end rounded or pointed, distinct lateral
notch; third pair of lobes as broad as long, rounded at
distal end, mesal and lateral notches, mesal sometimes
faint; fourth and fifth pairs of lobes more or less dis-
tinct, triangular, serrated, these probably projections;
pectinae arranged 2, 2, 3, 3, 0; paraphyses short, club-
shaped, pair located opposite second, third, and fourth
incisurae, between incisurae named and between me-
dian pair of lobes with a chitinous knob. — Ceylon on
Pothos. pothi Green.
dd. Pygidium with two pairs of paraphyses.
e. Pygidium with one paraphysis located in each second and
third incisura, mesal paraphyses slightly larger than lat-
eral; three pairs of lobes, median pair broad, truncately
rounded, entire; second pair of lobes smaller than me-
dian and larger than third; third pair bluntly rounded;
pectinae slender, arranged 2, 2, 3, 0; lateres strongly,
irregularly serrate; vulva located some distance cephalad
of anus; calles distinct, mesal calles fused. — Targionia
glomerata Green, — India on Saccharum.__g'Zomerafa Green,
ee. Pygidium with paraphyses located two in each second in-
cisura, mesal about twice as long as lateral and extending
cephalad beyond level of anus; two pairs of lobes; me-
dian pair of lobes large, elongate, mesal margins prac-
tically contiguous, distal end rounded, strong lateral
notch; second pair of lobes of same general shape as me-
dian, very much smaller, more pointed, notch stronger;
each lateris short distance cephalad of second lobe with
two small pointed projections, cephalad of these three
large broad strap-shaped pectinae, cephalad of these two
small pointed projections, margin cephalad of these ser-
rulate; vulva located some distance cephalad of anus. —
Philippine Islands on "rosal" or "campopot."
tayabana Ckll.
SPECIES OF MONAONIDIE-LLA
a. Pygidium with a single pair of paraphyses.
b. Pygidium with median pair of lobes adjacent, ends rounded and
emarginate; plates short and spiniform; each lateris serrate with
446 THE COCCIDAE
large widely forked furcapectinae near median lobe; paraphyses;
two in number, associated with median pair of lobes. — Australia
on Acacia, Templetonia. cerata Mask.
bb. Pygidium with median pair of lobes sharply truncate, without,
lacinations, not emarginate; lateres not with large widely forked
furcapectinae. — West Australia on Acacia. nivea Fuller.
aa. Pygidium with four pairs of paraphyses; median pair of lobes large,
prominent, strongly convergent, prominent lateral notch, lateral
margin oblique, distal end rounded; median incisura narrow, with-
out plates; indentation marking second incisura with two short
plates and two subequal clavate paraphyses, as long as or longer than
median lobe; minute projection cephalad of second incisura, prob-
ably representing lobe, cephalad of this a platey two cephalic para-
physes of each side adjacent, clavate, as long as caudal paraphyses;
single plate cephalad of cephalic paraphyses; anus distant from
caudal margin; dorsal oraceratubae few and inconspicuous. — Aus-
tralia on Acacia. suhfervens Green.
SPECIES OF NEOLEONARDIA
Pygidium with median pair of lobes fused into a single lobe, circular,
entire; each lateris with four stout truncate plates, in two groups of
two each, separated by indentation, another indentation cephalad of
cephalic group of plates, each indentation with two long setae, plates
nearly as long as lobes; paraphyses four in number, cephalic portion
swollen, club-shaped, mesal paraphyses longer than lateral, shortest
three or more times as long as median lobe; caudal portion of pygi-
dium constricted and margins emarginate. — Australia on Eucalyptus
extensa Mask.
SPECIES OF GREENOIDEA
Pygidium with median pair of lobes short, rounded, faint lateral and
mesal notches; second, third, and fourth pairs of lobes similar, round-
ed, smaller than median, lateral notch; paraphyses two or three times
as long as wide, attached mesal margin of second, third, and fourth
pairs of lobes; proximal ends of median pair of lobes prolonged; calles
present; plates or pectinae wanting; anus near middle of pygidium. —
Ceylon on Phyllanthus. phyllanthi Green.
SPECIES OF MASKE-LLIA
Pygidium with median pair of lobes rounded at distal end, entire; sec-
ond and third pairs of lobes short and broad, notched at distal end, dis-
tance between them three times distance between median and second
lobe; each lateris with projections at middle; setae on ventral side of
proximal end of lobes and lobe-like projections and on dorsal side of-
second and third pairs of lobes and lobe-like projections; anus slight-
ly nearer caudal than cephalic end of pygidium ; insects forming galls
on twigs. — ^West Australia on Eucalyptus glodosa Fuller.
SPECIES OF TARGASPIDIOTUS
a. Pygidium with median pair of lobes concave at distal end; thorax
not at all transversely constricted.
TRIBE ASPIDIOTINI 447
b. Pygidium with median pair of lobes quadrangular, concave at
distal end; second pair of lobes as broad at distal end as at proxi-
mal; densariae of second incisurae with mesal margin about twice-
as long as lateral. — Kansas on Andropogon, Panicum, Chrysopo-
gon. marlatti Parr.
bb. Pygidium with median pair of lobes quadrangular, lateral mar-
gin longest, concave; second pair of lobes as broad as median,
similar in shape; third pair of lobes similar in form, much
smaller; second and third incisurae with densariae, mesal margin
longer than lateral; dorsal oraceratubae numerous, arranged in
three rows; anus about seven times its width from caudal margin
and about twice from vulva. — Chrysomphalus tonilensis Ckll.,
Chrysomphalus covilleae Ferris. — New Mexico, Arizona on Yucca,
Bigelowia, Covillea. yuccarum Ckll.
aa. Pygidium with median pair of lobes never concave at distal end.
b. Pygidium with median pair of lobes quadrangular, bluntly rounded,
entire; second and third pairs of lobes similar in form, subequal
in size, all about as broad at distal as at proximal end; meso-
spiracerores 6; segments of preabdomen with numerous large
dorsal oraceratubae. — Australia on Casuarina casuarinae Mask.
bb. Pygidium with median pair of lobes adjacent, longer than wide,
mesal and lateral notches; second pair of lobes narrower, sides
nearly straight, large lateral notch; third pair of lobes small,
bluntly pointed; median and aiecond incisurae narrow, lobes
adjacent third incisurae broad; plates or pectinae wanting; me-
dian setae inconspicuous, others prominent; second incisurae ap-
parently with thickenings. — Australia on Dryandra
dryandrae Fuller.
SPECIES OF PYGIDIASPIS
Pygidium of adult female with median and second pairs of lobes similar,
broad, short, three or more times as broad as long, distal margin
obscurely notched on each side and slightly projecting between notches,
placed close together, with long thickened broad paraphysis-like pro-
jections extending cephalad from proximal ends; each lateris with
three broad projections, similar in form to lobes, broader, distal margin
faintly crenulate, without paraphysis-like projections, cephalic one of
each side smallest, distance between it and median one twice that be
tween caudal and median; conspicuous translucent curved channel
extending cephalad for distance equal to width of lobes; anus minute,
located between channels. — Australia on cedar logs cedri Green.
SPECIES OF TARGIONIA
a. Pygidium with strong tooth-like projections on each lateris.
b. Pygidium with tooth-like projections of lateres arranged in pairs.
c. Pygidium with median pair of lobes with distal end bluntly
rounded, quadrangular, distant, not converging, faint mesal
and prominent lateral notches; setae small. — France, Italy,
Algiers on Vitis vitis Sign.
448 THE COCCIDAE
cc. Pygidium with median pair of lobes projecting, distal end
truncate or subtruncate, faint lateral notch. — Italy on Quercus.__
vitis suberi Leon,
bb. Pygidium with tooth-like projections of lateres not arranged in
pairs.
c. Pygidium with two tooth-like projections on each lateris, simi-
lar to second and third pair of lobes, not projecting beyond
general contour of margin of pygidium; lateres very oblique;
median pair of lobes large, convergent, strongly projecting;
large bifurcate thickening prolonged inside lobes of each side
to anus; marginal setae prominent; pectinae or plates wanting.
— France on Alnus. alni Marchal.
cc. Pygidium with three tooth-like projections on each lateris, be-
coming successively smaller cephalad, located near median
lobes, remainder of each lateris entire; median pair of lobes
small, rounded, distinct lateral and faint mesal notches, near
but not contiguous; anus located four times its width from
caudal margin; pectinae and plates wanting; dorsal oracera-
tubae few in number, arranged near peripheral margin. —
Aonidia juniperi Miarl. — Utah on Juniperus juniperi Marl.
aa. Pygidium without strong tooth-like projections on lateres.
b. Pygidium with median pair of lobes large and projecting,
c. Pygidium with median pair of lobes large, prominent, project-
ing, somewhat convergent, distinct mesal and lateral notches;
setae prominent, two located between median pair of lobes and
five on each lateris in two groups, three near median lobe and
two some distance cephalad of these. — France on Cineraria,
Quercus. nigra Sign.
cc. Pygidium with median pair of lobes prominent, projecting, dis-
tant, subtriangular, distal end bluntly rounded, lateral notch;
second and third pairs of lobes indicated as slight triangular
projections; setae prominent, four on each side, one on each
median lobe and each projection indicating second and third
lobe and one on each lateris; anus distant from caudal mar-
gin two or three times its width. — Argentina on Fabiana
fabianae Leon,
bb. Pygidium with median pair of lobes very short, not projecting,
distinct lateral and mesal notches on caudal margin with mar-
gin bluntly pointed and projecting between them; lateres coarsely
crenulate with these undulations finely crenulate; setae minute;
pygidium of second nymphal female with three pairs of lobes,
median pair large with distinct lateral notch; second pair of lobes
smaller with lateral notch; third pair of lobes small, triangular
with mesal and lateral notches; pectinae longer than lobes, ar-
ranged 2, 2-3-2-3. — Chrysomphalus barbusano Lindgr. — Canary Is-
lands on Phoebe. Mrdusano Lindgr.
SPECTES OF AFFIRMASPIS
a. Pygidium with densariae distinct, caudal one largest, mesal and
lateral margins equal; median pair of lobes large, mesal margins
TRIBE ASPIDIOTINI 449
straight, near to but not touching, distal and lateral margins
convex, lateral angles not conspicuous, faint lateral notch; second
pair of lobes long, longer than broad, two distal notches, mesal mar-
gin nearly straight, distal and lateral contours pointed; setae, par-
ticularly of each lateris, very long, longer than median pair of lobes;
each lateris with broad, blunt; tooth-like projections; anus located
near caudal margin. — Sokotra, Africa on Dracaena, socotrana Lindgr.
aa. Pygidium with densariae distinct, caudal one largest, mesal mar-
gin much larger, nearly twice, than lateral; median pair of lobes
large, mesal margins convex, distant, lateral margin oblique, notched,
distal end bluntly rounded; second pair of lobes similar to median,
much smaller, lateral notch; setae prominent; (jach lateris without
tooth-like projections, entire or finely serrate; anus nine times its
width from caudal margin. — South Africa on Ehretia. __e7ire^iae Brain.
SPECIES OF TARGIONIDEA
a. Pygidium with median pair of lobes with three distinct lateral
notches, large, triangular, subadjacent, convergent; densariae want-
ing in median incisura, large, two margins subequal in second in-
cisurae, smaller in third, margins subequal; setae long and stout,
longer on median pair of lobes, seven on each side, four dorsal and
three ventral; pygidium of first nymph with single pair of lobes and
without pectinae or plates. — Canary Islands on Campylanthus
campylanthi Lindgr.
aa. Pygidium with median pair of lobes with single notch, large, pro-
jecting, subadjacent; second incisurae bounded by small, unsymmet-
rical, triangular, notched, lobe-like projections; densariae of second
incisurae with margins subequal; setae long and stout, subequal or
longer than median pair of lobes, six on each side, three dorsal
and three ventral. — Denmark on Euphorbia, Olea. lenticularis Lindgr,
SPECIES OF CIRCULASPIS
Pygidium with deep mesal indentation and three others on each side;
projections of margin of pygidium between indentations serrate; "con-
spicuous cylindrical canal" extending cephalad from mesal cleft over
half way to anus, terminating in "brush of small tubular ducts;" dor-
sal and ventral surfaces dotted with minute circular pores with thick-
ened rims; anus near cephalic end of pygidium; mesospiracerores and
metaspiracerores numerous. — India on bamboo
canaliculata Green.
SPECIES OF RUGASPIDIS
a. Pygidium with three indentations on each side; mesal projection with
slight mesal groove; margin of pygidium shallowly crenulate; anus
located slightly cephalad of vulva, about fifteen times its width from
caudal margin; mesospiracerores present; dorsal oraceratubae nu-
merous, irregularly placed. — California on grass. --graminis Bremner.
aa. Pygidium with numerous slight indentations, margin appearing to
have three pairs of indefinitely defined lobes with coarse crenula-
tions; dorsal oraceratubae numerous, without definite arrangement,
associated more or less with wrinkles of surface; setae long, slender.
450 THE coccroAE
four ventral and three dorsal, dorsal and ventral seta located on
each lobe-like projection; anus large, about eight times its width
from caudal margin and. three times from vulva. — Diaspis arizonica
Ckll. (Ferris.) — ^Arizona on Prosopis arizonica Ckll.
SPECIES OF OBTUSASPIS
Pygidium without lobes, pectinae, or plates; margin with about thirty
subquadrate, irregular, truncate, closely adjacent projections; meson
of caudal margin slightly incised with largest projection forming lat-
eral limits of incision, three much smaller projections in incision,
others successively smaller cephalad; pygidium short, very broad,
broadly rounded; dorsal oraceratubae small, few in number; anus and
vulva nearly superimposed, located near middle of cephalic half; pygi-
dium with inverted V-shaped thickenings with three large pores or
minute setae laterad of each side of V; scale of adult female dense,
hard, capsulate, two halves slightly separated at caudal end, probably
a puparium. — British East Africa on roots of Chloris
rhizophila Newst.
SPECIES OF LEONARDIANNA
a. Pygidium with four pairs of paraphyses, eight paraphyses, mesal
pair shortest, fourth pair longest, second and third pairs subequal;
margin of pygidium with four rounded lobular projections on each
side, entire, bearing three distant setae on cephalic half; scale dull
grayish brown. — California on Bigelovia Mgeloviae Ckll.
aa. Pygidium with seven subequal paraphyses, one located on meson,
others in pairs; margin of pygidium with three or four rounded lobu-
lar projections on each side; cephalic half of pygidium on each side,
preabdomen, thoracic segments, and margin of head with scattered
setae and numerous small circular pores, latter not extending
cephalad of mesospiracles ; surface of pygidium longitudinally
striate; margin of pygidium on each side finely dentate; mesospira-
cerores 3-8; anus and vulva superimposed, located at middle of pygi-
dium.— Jamaica on Pimenta. pimentae Newst.
SPECIES OF FROGGATTIELLA
a. Pygidium with two pairs of paraphyses, second pair very slightly
longer than first; pygidial margin with broad mesal emargination,
another adjacent to each second paraphysis, two distant deep angu-
lar notches cephalad of these, minutely irregularly serrate; pair of
setae caudad of caudal end of each paraphysis, another caudad of
caudal notch, two distant setae on each side of areas on each side of
cephalic notch; anus small, located cephalad of line drawn through
cephalic notches; spiracerores present. — Ceylon on Arundinaria
inusitata Green.
aa. Pygidium with three pairs of subequal paraphyses, three or four
times as long as broad; pygidium incised on meson with ihree prom-
inent angular equidistant teeth on each side with smaller teeth be-
tween them; mesal incision provided with dense brush of long taper-
ing setae, distal ends meeting in point; anus located near cephalic
end of pygidium; dense chitinous area with numerous minute cir-
cular pores. — Ceylon on Gigantochloa penicillata Green.
TRIBE ASPIDIOTINI 451
SPECIES OF STRINGASPIDIOTUS
a. Pygidium with four pairs of lobes; median pair large, conspicuous,
pointed, mesal and lateral notches; second and third pairs of lobes
narrow, lanceolate, constricted at proximal end; fourth pair of
lobes broad, distinct, mesal and lateral notches; pectinae subequal
in length to lobes, arranged 2, 2, 3, 3, 0; each lateris irregularly-
dentate; mesospiracerores 3-4; anus distant from caudal margin;
genacerores (12-12)12-12. — Java on Curculigo curculiginis Green.
♦aa. Pygidium with three pairs of lobes; median pair of lobes as broad
as long, distal end bluntly rounded, lateral notch deep, mesal notch
faint; second pair of lobes with lateral margin in part oblique, mesal
and lateral notches faint; third pair of lobes three times as long,
pointed, spine-like; pectinae arranged 2, 2, 3, 5, those of incisurae
distapectinae, those of third incisurae very broad, those of each
lateris unapectinae; setae short, not one-half length of lobes; body
with mesal part of cephalic margin toothed or deeply crenulate;
J genacerores (4-9)6-6; anus distant from caudal margin. — Selenaspidus
ferox Lindgr., Selenaspidus articulatus celastri Mask. — Cape of Good
Hope, Gold Coast, Africa on Celastrus, Plumiera-like Euphorbiaceae.
; celastri Mask.
SPECIES OF SEMELASPIDUS
'«,. Pygidium with anus distant from caudal margin about twice its
length; median pair of lobes small, rounded, entire; second and third
pairs of lobes smaller, slightly notched; second and median pairs
of lobes much closer together than second and third; each lateris
irregularly serrate; paraphyses four in number on each side, slender,
median pair longest; pectinae narrow, obscurely lacinate; meso-
spiracerores numerous; genacerores in thre3 groups, median 4-6,
lateral 3-6. — India on leaves of unidentified tree. --triglandulosa Green.
aa. Pygidium with anus distant from caudal margin about three times
its length; median pair of lobes triangular, bluntly pointed, sides
slightly emarginate; second pair of lobes smallest, near median pair;
third pair of lobes triangular, larger than median pair; pectinae
small and inconspicuous; paraphyses four in number on each side,
median pair slightly longer than others, these subequal; each lateris
with prominent serrate projections; genacerores in three groups,
median 2-4, lateral 2-5. — Ceylon on Cinnsunomum.. .cistuloides Green.
Aspidiotus capsulatus Green from Java on Piper has a similar scale
and undoubtedly belong to this genus.
SPECIES OF SELENASPIDUS
a. Body with cephalic margin entire or finely or distinctly serrate,
never crenulate with projections of two sizes.
b. Body with mesothoracic marginal spine suddenly drawn out,
conical, bluntly pointed, length about equal to greatest width;
median pair of lobes entire, rectangular, frequently with faint
lateral notches, distal end bluntly rounded, longer than broad;
second pair of lobes broader than median, subequal in length,
lateral margin obliquely rounded, one or two notches, distal end
452 THE COCCIDAE
rounded; third pair of lobes long, slender, spine-like, distant
from second pair; pectinae as long as lobes, arranged 2, 2, 3, 4-5,
those of median and second incisurae furcapectinae, of third in-
cisurae broad distapectinae, and of lateres unapectinae; gena-
cerores in two groups of 6-8 cerores each; calles present, mesal
fused. — Selenaspidus articulatus simplex Charm. — England, West
Africa, Demerara, West Indies, Brazil, Panama, Costa Rica, Mex-
ico on Pandanus, Dictyosperma, Cordyline, orange, lime, coffee,
Gardenia, Ficus, etc. articulatus Morg.
bb. Body with mesothoracic marginal spine not suddenly drawn out,
twice as long as wide.
c. Body with cephalic margin distinctly serrate; median pair of
lobes quadrangular, entire, with rounded angles, longer than
broad; second pair of lobes broader than long, rounded, distal
margin oblique, sinuate or notched; third pair of lobes dis-
tant from second, spine-like, hooked, cephalic surface convex,
caudal concave, not conical; pectinae as long as lobes, ar-
ranged 2, 2, 3, 4-6, those of each third incisura very broad;
genacerores in two groups; calles present, mesal calles not
fused. — Aspidiotus (Selenaspidus) articulatus magnospinus
Newst. — Uganda, Africa magnospinus Newst.
cc. Body with cephalic margin entire; median pair of lobes
broader than long, broadly rounded, entire or faintly notched;
second pair of lobes similar in size and form to median pair,
entire; third pair of lobes distant from second, spine-like, not
hooked, cephalic and caudal margins concave or straight, as
long as pectinae, length three times greatest width; pectinae
subequal in length to lobes, arranged 2, 2, 3, 2, those of each
third incisura broad; genacerores in two groups, 7-10; dorsal
oraceratubae numerous. — ^South Africa on Euphorbia, Mi-
musops. pertusus Brain.
aa. Body with cephalic margin crenulate with projections of two sizes,
large projections with smaller ones interspersed; median pair of
lobes with distal end bluntly pointed, distal portion of lateral mar-
gin suboblique, mesal and lateral notches faint to distinct, longer
than broad; second pair of lobes longer than broad, distal end
bluntly pointed, lateral margin oblique, mesal notch near distal
end and lateral notch near middle of lateral margin; third pair of
lobes longer than broad, pointed, spine-like; pectinae subequal in
length to lobes, arranged 2, 2, 3, 4-6, those of median incisura dista-
pectinae, others unapectinae, those of each lateris irregular; dorsal
oraceratubae numerous; genacerores in two groups; mesothoracic
marginal spine conical, blunt, greatest width equal to length. —
Kamerun, Africa on palms kamerunicus Lindgr.
SPECIES OF DUPLASPIDIOTUS
a. Pygidium with two pairs of paraphyses, cephalic end enlarged,
knob-like, long pair between first and second pairs of lobes, pair
about one-half this length between second and third pairs of lobes;
median pair of lobes large, rounded at distal end, distinct lateral
TRIBE ASPIDIOTINI 453
notch; second and third pairs of lobes distinct, much smaller than
median, rounded on mesal and distal margins, prominent lateral
notch; each lateris with numerous tooth-like processes; plates short,
arranged 0, 0, 2, 2; mesopsiracerores 11, metaspiracerores 0; gena-
cerores arranged in inverted U-shaped mass, — Ceylon on Acalypha.
irreptus Ruth.
aa. Pygidium with three or four pairs of paraphyses.
b. Pygidium with three pairs of paraphyses, strongly developed,
ending in distinct knob; four pairs of lobes, median pair scarcely
notched on mesal margin; second and third pairs of lobes nar-
rower than median; lobes all of same general character, second
and third pairs much decreased in size; each lateris cephalad of
fourth pair of lobes serrate as if with many small lobes; anus
distant from caudal margin; genacerores present; mesogenacerores
and pregenacerores forming a continuous semicircle. — Natal,
Africa on Camellia. claviger Ckll.
b. Pygidium with four pairs of paraphyses, mesal pair located
cephalad of median incisura, not enlarged at cephalic end, short-
er than others, all other pairs with distinct ball-like enlarge-
ment, second pair longest, third and fourth pairs subequal, longer
than first; median pair of lobes longer than broad, triangular,
mesal and lateral notches; second and third pairs of lobes as long
as median, much narrower, notched; lateres with triangular lobe-
like projections; plates shorter than lobes, slender, arranged
2, 2, 3, 5; mesospiracerores 7-9; genacerores in form of horse-
shoe, slightly interrupted at middle, 17-23 on each side. — South
Africa. niger Brain.
SPECIES OF PSEUDAONIDEA
a. Pygidium with four pairs of subequal lobes.
b. Body of adult female twice as long as wide, tapering uniformly
from constriction to caudal end; pygidium with median pair of
lobes with sides subparallel, distal ends rounded, mesal and lat-
eral notches; second, third, and fourth pairs of lobes similar in
shape, each pair successively slightly smaller; pectinae deeply
lacinate, arranged 2, 2, 3, 3, 0; each lateris with its margin
irregularly serrate, with two deep notches, each bounded on
cephalic side by tooth-like projection; genacerores 0-2(21-24)16-27,
rarely with mesogenacerores, pregenacerores always most numer-
ous; mesospiracerores 12-20; scale of adult female reddish brown,
exuviae yellow. — Ceylon, East Indies, Mauritius, Japan, Java,
Brazil, British Columbia on Nothopegia, Dalbergia, Ficus, orange,
grape-fruit, "umame," mandarin triloMtiformis Green.
bb. Body of adult female oval, not twice as long as wide; genacerores
(23-25)10-11; mesospiracerores 15-17; scale of adult female almost
pure white. — Pseudaonidia trilobitiformis darutyi Charm. — Mauri-
tius, Seychelles, Liberia, Brazil on Mangifera, Murraya, Euphoria.. _
darutyi Charm.
454 THE COCCIDAE
aa. Pygidium with four pairs of very unequal lobes; median pair of
lobes projecting distinctly beyond others, broadly bluntly pointed,
entire, as broad as long; second and third pairs of lobes similar,
elongate, sides subparallel, bluntly pointed, entire, twice as long as
wide; fourth pair of lobes short, as broad as long, bluntly rounded,
entire; pectinae not deeply lacinate, arranged 2, 2, 3, 3, 0, mostly
narrow distapectinae with few teeth; each lateris coarsely serrate
with deep indentations; genacerores (30-42) 20-30; mesospira-
cerores 13-25; scale blackish brown with orange colored exuviae. —
Pseudaonidia duplex CklL, Pseudaonidia rhododendri Green, Pseu-
daonidia rhododendri thearum Fern. — India, Hawaiian Islands,
Japan, California, District of Columbia on tea. Camellia, Eurya,
Myrica, Olea, Rhus, camphor-tree, orange, rhododendron, "Umame,"
Azalea. theae Mask.
SPECIES OF PSEUDAONIDIELLA
Pygidium with four pairs of lobes; median, second, and third pairs with
mesal and lateral notches; fourth pair with single notch; each lat-
eris with two or three pectinae; mesospiracerores about 15; pregena-
cerores and postgenacerores of each side apparently fused to form
single group consisting of 70-76 cerores. — Japan, California on Peony,
Camellia, Rhododendron, Ilex, Thea paeoniae CklL
SPECIES OF PARAONIDEA
Pygidium with four pairs of lobes; median pair of lobes largest, others
successively smaller, all with mesal margin straight, lateral margin
oblique and notched; pectinae not conspicuously lacinate, arranged
2, 2, 3, 3, 0; paraphyses club-shaped, enlarged at cephalic end, three
pairs, first and second pairs subequal, slightly longer than third pair;
each lateris with prominent indentation; mesal portion of dorsum with
large lattice-shaped area; anus distant from caudal margin; genacerores
in two large curved groups of about 45; metathorax with lateral mar-
gins straight, narrower than portions of body cephalad and caudad of
it. — Ceylon on Mimusops. malleola Green.
SPECIES OF ENTASPIDIOTUS
a. Pygidium with median pair of lobes entire.
b. Pygidium with eleven plates or pectinae on each lateris; median
pair of lobes entire, broader than long, broadly rounded; second
pair of lobes as broad as median, entire, slightly oblique, broader
than long; third pair of lobes longer than broad, lateral margin
oblique with faint emargination; pectinae and plates arranged
2, 2, 3, 11, plates in incisurae, dagger-shaped, sometimes finely
serrate on each lateris near third lobe; dorsal oraceratubae nu-
merous; setae minute, shorter than lobes. — Abyssinia on
Euphorbia. magnus Lindgr.
bb. Pygidium always with considerably less than eleven plates or
pectinae on each lateris.
c. Pygidium with two pectinae or plates or less on each lateris.
d. Pygidium with clearly defined more strongly chitinized
tongue-shaped area on ventral aspect, pointing caudad; me-
TRIBE ASPIDIOTINI 455
dian and second pairs of lobes similar, entire, truncately
rounded, constricted, subequal in size; third pair of lobes
minute, described as obsolete, pointed; pectinae subequal in
length to lobes, arranged 2, 2, 3, 1, those of each third in-
cisurae and each lateris broad, unapectinae, consisting of
long distal projection and remainder broad area with short
teeth; dorsal oraceratubae small, numerous, scattered; anus
located slightly caudad of tongue-shaped area; lateres crenu-
late. — South Africa on Euphorbia euphorHae Newst.
dd. Pygidium never with strongly chitinized tongue-shaped area;
median pair of lobes oval, not very large; second pair of
lobes subequal to median; third pair of lobes small; pectinae
or plates arranged 2, 2, 3, 2, those of incisurae furcapectinae,
those of each lateris broad obliquely truncate plates; lateres
without teeth; dorsal oraceratubae numerous, arranged in
rows; calles divided into four parts. — Cape Town, South
Africa on Mesembryamthemum lounsJ)uryi Marl.
cc. Pygidium with four pectinae on each lateris, pectina arranged
2, 2, 3, 4, all trilacinate; lobes four pairs, similar, median pro-
jecting, others decreasing regularly in size; lateris with teeth
in three areas; spiracerores 7-8; dorsal oraceratubae large, con-
spicuous, arranged in rows; calles wanting. — Java on mango-
steen and mango greeni Marl.
aa. Pygidium with median pair of lobes notched.
b. Pygidium with three or more pectinae on each lateris adjacent
to third lobe.
c. Pygidium with median pair of lobes distinctly longer than
broad, distal end bluntly rounded, mesal and lateral notches
broad and shallow; second pair of lobes similar in form,
smaller, distinct mesal and faint lateral notches; third pair of
lobes longer than broad, dagger-shaped; pectinae subequal in
length to lobes, arranged 2, 2, 3, 6-7, mesal projection of third
incisurae spine-like plate, others unapectinae, those of each
lateris irregular unapectinae; dorsal oraceratubae few. —
Kamerun, Africa on Agelaia, Tricalysia. gracilis Lindgr.
cc. Pygidium with median pair of lobes distinctly broader than long,
d. Pygidium with width of third incisurae distinctly greater
than total width of median pair of lobes and median in-
cisura; median pair of lobes broader than long, distal end
bluntly pointed, mesal and lateral notches deep; second pair
of lobes longer than broad, lateral margin slightly oblique,
mesal and lateral notches deep; third pair of lobes sharply
pointed, spine-like; pectinae slightly longer than median
lobes, arranged 2, 2, 3, 4, those of third incisurae, particu-
larly two cephalic, broad distapectinae, those of each lateris
very irregular; dorsal oraceratubae numerous. — Kamerun,
Dutch East Africa and south Africa on Anacardiaceae, Bandei-
raea, Rinorea, Ficus, Aberia, Euonymus, Berberis, Chrysophyl-
lum, tea, citrus, palms. _ silvaticus Lindgr.
456 THE COCCIDAE
dd. Pygidium with width of third incisura distinctly less than
total width of median pair of lobes and median incisura;
median pair of lobes quadrangular, distinct lateral and me-
sal notches, sides straight, subparallel; second pair of lobes
longer than broad, faint mesal and lateral notches; third
pair of lobes sharply pointed, spine-like; pectinae arranged
2, 2, 3, 2-3, those of each third incisura and lateris not or
only slightly longer than third lobe; spiracerores wanting. —
South Africa on Phormium. pumilus Brain.
bb. Pygidium never with more than one pectina on each lateris,
sometimes none.
c. Pygidium with caudal pectina of each third incisura much nar-
rower than others; median and second pairs of lobes similar
in size and shape, distinct mesal and lateral notches, bluntly
rounded between notches, proximal portion more or less con-
stricted; third pair of lobes small, entire, as long as adjacent
pectinae, shorter than other lobes, crenulate; pectinae sub-
equal in length to lobes, arranged 2, 2, 3, 1-0, one on each lat-
eris broad, adjacent to third lobe; setae small, short; dorsal
oraceratubae numerous, tubes very long. — Southwest Africa on
succulent plant. schultzei Newst.
cc. Pygidium with caudal pectina of each third incisura as broad
as one next to it; median and second pairs of lobes subequal,
second pair slightly smaller, faint mesal and lateral notches,
proximal portion more or less constricted; third pair of lobes
pointed, as long as pectinae and other lobes; pectinae dista-
pectinae, arranged 2, 2, 3, 1, subequal in length to lobes; setae
inconspicuous. — South Africa on Arthrosolen griqus Brain.
SPECIES OF SCHIZASPIDIOTUS
Body with deep mesal incision at cephalic end and deep thoracic con-
striction apparently between prothorax and mesothorax and two others
on caudal portion of preabdomen; median pair of lobes quadrangu-
lar, deep lateral and mesal notches with bluntly pointed projection
between them, distant; second pair of lobes smaller than median,
entire, distal end broadly rounded; pectinae deeply lacinate, longer
than lobes, arranged 2, 2, 6 or more, one next each second lobe a
broad pectina, others plates; anus slightly more than its width from
caudal margin; vulva about same distance cephalad of anus. — Philip-
pine Islands on Ficus. lo'bata Ckll. & Robs.
SPECIES OF AMBIGASPIS
Pygidium with median pair of lobes large, subtriangular, bluntly pointed,
both margins crenulate; second pair of lobes bluntly rounded, mesal
margin straight, lateral oblique, crenulate, broader than long; third
pair of lobes twice as broad as long, short, mesal margin straight,
lateral oblique, crenulate; plates dagger-like, longer than lobes, ar-
ranged 0, 0, 3, 5-7, those of each lateris broader, caudal three more
than twice as long as those of third incisurae; anus and vulva situ-
ated at about same level; mesospiracerores and metaspiracerores 2-2;
TRIBE ASPmiOTINI 457
la;ttice-shaped thickenings not strongly chitinized, seen with difficulty.
— South Africa on Lycium. lycii Brain.
SPECIES OF LATTASPIDIOTUS
a. Pygidium with plates or pectinae in median incisura.
b. Pygidium with furcapectinae, wanting on lateres, furcapectinae
distinctly divided, slightly longer than lobes, arranged 2, 2, 3, 0;
median pair of lobes longer than broad, slightly constricted on
proximal portion, mesal and lateral notches distinct, distal end
bluntly rounded between them; second and third pairs of lobes
long, narrow, twice as long as broad, distinct mesal and lateral
notches; mesospiracerores 9-18, metaspiracerores (?)0; oracera-
tubae large, arranged in longitudinal rows; body with cephalic
portion bearing many large, widely separated setae; anus distant
from caudal margin. — Uganda, Africa on Baikea.__6aifceae Newst.
bb. Pygidium with plates, one to three on each lateris, plates as long
as or slightly longer than lobes, arranged 2, 2, 3, 1-3; median
pair of lobes longer than broad, subtriangular, bluntly pointed,
faint lateral notch; second and third pairs of lobes smaller than
median, second smaller than third, subtriangular, faint lateral
notch or entire; lateres with indentations and lobe-like projec-
tions; three pairs of paraphyses with cephalic end with ball-like
enlargements, second and third pairs sometimes represented only
by cephalic enlargements; anus located on line drawn through
cephalic enlargements of second paraphyses, covered by cuticu-
lar flap, caudal margin of flap deeply emarginate. — South Africa
on Acacia. laciniae Brain.
aa. Pygidium without pectinae or plates in median incisura.
b. Pygidium with two plates in each second incisura; median pair
of lobes quadrangular, entire, truncate with rounded angles;
second pair of lobes smaller than median, broader than long,
faint lateral notch, rounded; third pair of lobes broader than
second pair, distinct mesal and lateral notches with rounded pro-
jection between them; each lateris with two pairs of dentate
projections, generally larger than lobes; plates short, subequal in
length to lobes, arranged 0, 2, 3; setae slightly longer than lobes;
paraphyses in three pairs, mesal pair longest, lateral pair short-
est, uniform in width throughout; mesospiracerores 4-4, meta-
spiracerores 1-2. — Barbados on grape fossor Newst.
bb. Pygidium with one plate in each second incisura.
c. Pygidium with single plate on each lateris cephalad of third
lobe, this plate about twice as long as median lobe, other
plates subequal in length to lobes, arranged 0, 1, 3, 1; median
pair of lobes broader than long, faint mesal and lateral
notches; second pair of lobes short, broader than long, very
faintly notched; third pair of lobes irregular in form, some-
times notched; setae very long and slender, especially cephalic
ones; paraphyses in three pairs, distinctly knobbed at cephalic
end, mesal and second pairs subequal, third pair oval in out-
458 THE COCCIDAE
line and one-half length of second. — Antigua, British West
Indies, Java, Mauritius, Mexico on Malvaviscus, Prunus, Vitis—
tesserata Charm.
CO. Pygidium without plates on each lateris, plates inconspicuous,
arranged 0, 1, 1-2, 0; median pair of lobes largest, usually
broadly rounded, sometimes slight mesal and lateral notches;
second and third pairs of lobes similar in shape, second pair
larger than third, rounded at distal end, prominent lateral
notch, slightly inclined caudad; each lateris with several
sharp-pointed processes; paraphyses six in number, mesal and
second pairs subequal, cephalic end of each enlarged, knob-
like, third pair located cephalad of third pair of lobes, incon-
spicuous; oraceratubae arranged in longitudinal rows; setae
long, increasing in length, 'cephalad. — Ceylon on Oreodoxa,
Acalypha. oreodoxae Ruth.
SPECIES OF PARTARGIONIA
Head with cephalic indentation; transverse constriction between meso-
thorax and metathorax, first and second abdominal segments, third
and fourth abdominal segments, fifth segment and pygidium; pygi-
dium with four pairs of tooth-like lobes; small setae between second
and third, third and fourth pairs of lobes, and on each lateris; trans-
verse thickening near middle of pygidium and with two projections
on cephalic side. — India on Artocarpus artocarpi Green.
SPECIES OF PARROTTIA
Pygidium with three pairs of lobes, third pair small and tooth-like;
paraphyses two in number, associated with second incisurae; anal
opening on line drawn through third pair of lobes, and one on each
lateris. — India on Grislea. moorei Green
SPECIES OF NEOMORGANIA
a. Pygidium with distal end of median pair of lobes serrate, oblique,
proximal ends adjacent, distal ends distant, projecting but little
beyond pygidial margin, other lobes obsolete; each lateris with three
indentations, each with an oraceratuba; mesospiracerores 15, meta-
spiracerores 10; plates a single one on each lateris adjacent to
median lobe. — New South Wales, Tasmania on Eucalyptus, Acacia,
Hakea. acaciae Morgan.
aa. Pygidium with median pair of lobes with distal ends entire; dorsal
oraceratubae minute and limited to pygidium.
b. Pygidium with median pair of lobes distinctly separated, oblique,
entire; second and third pairs of lobes represented by mere
teeth; each lateris with setae nearly as long as median pair of
lobes; spiracerores apparently wanting. — Australia on Eucal-
yptus, Casuarina. eucalypti Mask.
bb. Pygidium with median pair of lobes contiguous along mesal mar-
gin, oblique exteriorly, giving appearance of single mesal area or
lobe. — Australia on Acacia. junctiloba Marl.
TRIBE ASPIDIOTINI 459
SPECIES OF BAKERASPIS
Pygidium with median pair of lobes rounded, entire; second pair of lobes
deeply incised, mesal lobelet larger than lateral, lobelets rounded,
entire; third pair of lobes broad, not prominent, deeply concave at
middle of distal margin; plates subequal in length to median pair of
lobes, arranged 2, 2, 2, 2, those of each lateris placed far apart; lat-
eres finely crenulate; preabdomen with lateral margin fringed with
quadrate, serrate, scale-like projections; genacerores in two groups of
about 150 cerores each. — Philippine Islands on Schizostachyum
schizostachyi Ckll. & Robs.
SPECIES OF GREENIELLA
a. Pygidium of adult female triangular, as long as or longer than broad,
b. Pygidium of second nymphal female with median pair of lobes
with mesal and lateral notches.
c. Pygidium of second nymphal female with latapectinae or una-
pectinae, unequal in length; median pair of lobes elongate,
distal end bluntly rounded, mesal and lateral notches; sec-
ond and third pairs of lobes smaller, oblique, bluntly pointed,
lateral margin serrate;^ pecftiiuae longer than lobes:, disitar<
pectinae and unapectinae, arranged 2, 2, 3, 1-2; each lateris
with several, eight to ten, irregular, serrate, pectinae-like pro-
jections; pygidium of adult female with asymmetrically ar-
ranged projections, variable in form, frequently with two or
three pairs of long irregular projections with smaller projec-
tions between them. — Ceylon on Psychotria cornigera Green.
CO. Pygidium of second nymphal female with broad distapectinae,
subequal in length; median pair of lobes elongate, much long-
er than wide, distinct mesal and lateral notches, distal end
prolonged and rounded between notches; second pair of lobes
similar in size and shape to median; third pair of lobes
smaller, similar in size and shape to second pair; pectinae
slightly longer than lobes, arranged 2, 2, 3, 6-7, those of me-
dian incisura furcapectinae, of second and third incisurae
narrow distapectinae, of lateres broad distapectinae; incis-
urae with seven altaceratubae-like oraceratubae ; pygidium of
adult female triangular, elongate; median pair of lobes
elongate, parallel sides, distinct lateral and mesal notches;
second pair of lobes elongate, dagger-like, pointed; plates and
pectinae arranged 2, 1, 0, those of median incisura plates or
furcapectinae, others distapectinae; lateres undulate an(i
finely serrate; setae minute. — Kamerun, Africa on Schotia,
Crudia, Cynometra. Hafrae Lindgr.
bb. Pygidium of second nymphal female with median pair of lobes
entire, elongate, distal end oblique, and continuous with lateral
margin; second pair of lobes short and broad, bluntly pointed^
subtriangular, entire; pectinae short with shallow lacinations, ar-
ranged 2, 2, 3, those of each lateris placed in an indentation
cephalad of each second lobe; lateres with prominent serrate pec-
tina-like or plate-like projections; pygidium of adult female with
460 THE COCCIDAE
pygidial margin with asymmetrical small projections, never with
long paired projections. — Ceylon on Mesua. messuae Leon.
aa. Pygidium of adult female not triangular, distinctly broader than
long.
b. Pygidium of second nymphal female with pectinae or plates want-
ing in median incisura.
c. Pygidium of adult female with distinct serrate or deeply laci-
nate pseudolobes.
d. Pygidium of adult female with narrow, distinctly lacinate
pectina-like pseudolobes, pseudolobes arranged in groups
with two or three small tooth-like projections between them;
setae long, subequal in length to pseudolobes; pygidium of
second nymphal female with three pairs of similar, sub-
equal, elongate lobes, mesal notch wanting, lateral notch
deep and distinct; pectinae as long as or longer than lobes,
arranged 0, 2, 2, 8. — India on Walsura dentata Lindgr.
dd. Pygidium of adult female with three pairs of pseudolobes,
distinctly projecting, broader than long, all similar in size
and form, distal margin uniformly deeply serrate; other
smaller pseudolobes placed between larger, arranged 2, 2,
3, 2; lateres except pseudolobes named, without projections;
pygidium of second nymphal female with median pair of
lobes short, twice as broad as long, distal margin with five
tooth-like projections; second pair of lobes similar to me-
dian, shorter; third pair of lobes short, smaller than sec-
ond pair, irregular; pectinae arranged 0, 2, 2, 1, broad dista-
pectinae; incisurae with six altaceratuba-like oraceratubae.
— South Australia on Casuarina. paradoxa Lindgr.
cc. Pygidium of adult female not with lacinate pseudolobes, with
three pairs of projections, lobe-like in form and arrangement,
margin of each with two prominent indentations; each side
with five fiask-shaped plate-like projections, arranged one in
second and third incisurae and three equidistant on lateres,
attached by broad base; setae particularly of lateres, long and
slender, longer than lobe-like projections; pygidium of second
nymphal female with three pairs of lobes, median pair large,
subadjacent, deep lateral notch, distal end bluntly pointed;
second and third pairs of lobes smaller, with deep distal notch;
plates or pectinae wanting; setae longer than median lobes. —
Burma on Miliusa. ta7'gioniopsis Lindgr.
bb. Pygidium of second nymphal female with pectinae or plates in
median incisura.
c. Pygidium of adult female with lobes or pseudolobes.
d. Pygidium of adult female with two pairs of more or less dis-
tinct lobes.
e. Pygidium of adult female with pair of short, crenulate,
lobe-like projections; each lateris with one or two short,
lobe-like extensions, each bearing setae, margin of lateres
finely crenulate; pectinae much longer than median lobe-
TRIBE ASPIDIOTINI 461
like projections, arranged 1, 1; setae long, particularly-
median pair; pygidium of second nymphal female- with
median pair of lobes projecting, bluntly rounded, distinct
lateral notch; second pair of lobes subtriangular, blunt,
lateral noitch; third pair of lobes triangular, pointed;
pectinae long, deeply lacinate, much longer than lobes, ar-
ranged 2, 2, 3, 3-4, those of lateres smaller than those of
incisurae; incisurae with altaceratbuae-like oraceratubae ;
anus twice its width from caudal margin. — Natal on
Euphorbia. austro-africana Lindgr.
ee. Pygidium of aduHt female with distinct lobe-like lobes;
median pair of lobes subadjacent, mesal margins sub-
parallel, bluntly rounded, lateral margin oblique, two or
three lateral notches; second pair of lobes broadly round-
ed, twice as broad as long, slight mesal and lateral emar-
ginations; plates shorter than lobes, truncated, arranged
0, 2, 8, those of each lateris arranged 3-1-1-1-1-1; setae
long and stout; pygidium of second nymphal female
with median pair of lobes longer than broad, rounded,
lateral notch; second pair of lobes similar in form and
subequal in size; third pair of lobes much smaller, bluntly
rounded, longer than broad, lateral notch; pectinae about
as long as lobes, arranged 2, 2, 3, 7, ithose of median
incisura furcapectinae, those of other incisurae and adja-
cent to third lobe broad distapectinae, others more point-
ed and irregular in form, those of each lateris arranged
3-2-2; setae as long as lobes; incisurae with altaceratuba-
like oraceratubae. — India on Mimusops
spinosissima Lindgr.
dd. Pygidium of adult female without lobes, but with pectina-
like pseudolobes.
e. Body of adult female entire, not crenulate; pygidium of
adult female hyaline, without lobes, provided with twenty-
four long pectina-like pseudolobes; anus located about
three times its length from caudal margin; anus and vulva
superimposed; pygidium of second nymphal female with
pectinae arranged 2, 2, 3, 3, equal in length with lobes;
each lateris with an acute lobe-like projection cephalad of
pectinae; median pair of lobes widest at middle, lateral
and slight mesal notch; second pair of lobes with lateral
notch; third pair of lobes with lateral and mesal notches. —
Ceylon on Mesua. ferreae Ruth.
ee. Body of adult female with margin with deep rounded
crenulations; pygidium without lobes, but with about
seven elongate deeply lacinate pectina-like pseudolobes,
one located on meson; margin of pygidium between
pseudolobes disitinctly dentate; pygidium of second nymph-
al female with median pair of lobes elongate, longer than
broad, rounded, mesal and lateral notches; second and
third pairs of lobes similar in form, larger; fourth pair
462 THE COCCIDAE
of lobes subequal in size, somewhat irregular in form;
fifth pair of lobes triangular, pointed; pectinae longer
than lobes, deeply lacinate, arranged 2, 2, 3, 3, 3, 2; setae
minute, not as long as lobes; margin apparently with eight
altaceratuba-like oraceratubae. — India on Aglaia
viridis Lindgr.
cc. Pygidium of adult female without lobes or pseudolobes, with
about five slight undulations, margin serrate; setae minute,
inconspicuous; pygidium of second nymphal female with me-
dian pair of lobes large, elongate, subquadrangular, rounded,
disltinct lateral notch; second pair of lobes similar in size
and form to median lobes; third pair of lobes one-third size
of second pair, bluntly pointed; pectinae about as long as lobes,
arranged 2, 2, 2, 2, those of each lateris adjacent third lobe
and broader than others; lateres irregularly dentate. — New
Caledonia on Podocarpus. Tonga Lindgr.
SPECIES OF AONIDIA
a. Pygidium of adult female without lobes and with pseudolobes, sec-
ond nymphal female with three pairs of lobes.
b. Pygidium of second nymphal female known.
c. Pygidium of second nymphal female with pectinae arranged
2, 2, 3, 0; pectinae with sides parallel, subequal in length to
lobes; lobes conspicuous, three pairs, lateral and mesal mar-
gins parallel, distal margin crenulate, pygidium of adult fe-
male with mesal depression on caudal margin on each side of
which occasionally short wide faint lobes can be identified. —
West Australia on Banksia. Mnksiae Fuller.
cc. Pygidium of second nymphal female with pectinae arranged
0, 2, 2, 12, each lateris completely fringed with narrow dista-
pectinae, pectinae not longer ithan lobes; median pair of lobes
elongate, much longer than wide, mesal margin straight, distal
and lateral margins continuous curve, notched; second and
third pairs of lobes similar to median; pygidium of adult
female with numerous asymmetrical narrow lobe-like projections
with plate-like projections at their distal end. — Ceylon on
Ficus. planchonioides Green.
bb. Pygidium of second nymphal female unknown.
c. Pygidium of adult female always with lobe-like or plate-like
projections, although sometimes small.
d. Pygidium of adult female with mesal lobe-like projection.
e. Pygidium of adult female with truncate mesal projection
bearing three small spine-like setae; two conical projec-
itions on each side cephalad of mesal projection, equidis-
tant from each other and mesal projection, each bearing
one comparatively long and two short setae; margin of
pygidium with low, broad, inconspicuous, lobe-like projec-
tion just cephalad of each projection, minutely serrate;
* anus and vulva located near middle of pygidium. — Ceylon
on Mesua. perplexa Green.
TRIBE ASPIDIOTINI 463
ee. Pygidium of adult female with mesal truncate projection
not bearing setae, sides constricted, much smaller triangu-
lar lobe-like projection on each side and adjacent tto mesal
projection; margin of pygidium adjacent to these latter
projections with three or four rounded prominences; anus
large and prominent, not adjacent to margin; scale yellow
with a whitish secretion which persists only as a marginal
fringe; first nymphal exuvia deciduous. — Ceylon on
Carissa. pusilla Green.
dd. pygidium of adult female never with mesal projection.
e. Pygidium of adult female with margin with six minute
thorn-like projections, arranged hook-like, three on each
side of meson extending laterad; margin cephalad of pro-
jections entire or gently crenulate; setae minute; anus
located slightly caudad of middle of pygidium; scale dull
reddish-brown, roughened with innumerable, slender,
curved, spine-like projections that are not soluble in caustic
potash. — Ceylon on Hemicyclia echinata Green.
ee. Pygidium of adult female never with minute hook-like pro-
jections with long slender pseudolobes.
f. Pygidium of adult female not with lacinate pseudolobes.
g. Pygidium of adult female with pseudolobes clavate,
pair on meson, pair on each side of these and distant
from mesal pair, adjacent to and cephalad of each
of these on each side, a broad triangular lobe-like
projection, then three clavate projections, a broad
triangular lobe-like projection, and finally three
irregular clavate projections; anus located about
midway between vulva and caudal margin, four times
its width from caudal margin; caudal end of pygi-
dium truncate. — Ceylon on Psychotria
spatulata Green,
gg. Pygidium of adult female with large number of long,
slender, dagger-shaped pseudolobes, distal end al-
ways pointed, never clavate; setae of one surface
nearly as long as pseudolobes and of other surface
minute, hardly projedting beyond margin; anus lo-
cated near and caudad of vulva. — Spain on Billartia.__
picea Leon,
ff. Pygidium of adult female with median pair of pseudo-
lobes prolonged into lacinate processes with deeply ser-
rate distal ends, other lobes and here and there on
margin with smaller similar projections; setae spine-
like, few in number; females noit excreting any definite
and separate scale, congregated in numbers under gen-
eral mass of thin, whitish, curling, cottony threads;
second exuvia much larger than adult female. — Australia
on Syncarpia. syncarpiae Mask.
CO. Pygidium of adult female without lobe-like or plate-like pro-
464 THE COCCIDAE
jections; margin of pygidium irregularly crenulate; dorsal
and ventral surface with few irregularly distributed oracera-
tubae; anus large, circular, located near middle of pygidium;
pygidium of second nymphal female resembling species of
Parlatoria or Gymnaspis; scale of male and first nymphal fe-
male, when present, with number of long curling glassy brittle
processes of wax. — Australia on Callistemon pulchra Green.
aa. Pygidium of adult female with symmetrically arranged projections,
apparently lobes and not pseudolobes.
b. Pygidium of adult female with more than two pairs of lobes.
0. Pygidium of adult female with four pairs of lobes, pygidium
transverse with caudal margin subtruncate; median pair of
lobes narrow, subquadrangular, mesal and lateral notches;
second pair of lobes projecting farther caudad than median
lobes, similar in shape, mesal and lateral notches; third and
fourth pairs of lobes successively shorter than each dther and
second pair of lobes, mesal and laternal notches; plates slender,
hardly longer than lobes, arranged 2, 3, 5, 5, 6, those of lateres
adjacent to fourth pair of lobes; cephalic end of body with
deep mesal cleft. — Ceylon on Loranthus loranthi Green.
CO. Pygidium of adult female with three pairs of lobes and pygi-
dium semicircular, caudal margin uniformly convex, never
subtruncate; median pair of lobes distant, bluntly pointed,
subtriangular, entire; second pair of lobes similar in shape
to median pair, more than twice as far from median as dis-
tance between median pair of lobes, entire; third pair of
lobes minute, similar to second pair; plates of second and
third incisurae twice or more as long as lobes, arranged 2, 3,
3, 2, those of lateres adjacent to third pair of lobes; each lat-
eris with two indentations; cephalic end of body entire, not
cleft. — Ceylon on Loranthus. obscura Green.
bb. Pygidium of adult female with two pairs of lobes.
c. Pygidium of adult female without pectinae or plates; median
pair of lobes contiguous at proximal end, bluntly pointed, tri-
angular, lateral margin with two notches; second pair of lobes
small, triangular, entire; each lateris with three equidistant
setae and two broad emarginations or indentations; pygidium
of second nymphal female with three pairs of lobes, median
pair not contiguous, mesal margins straight, distal and lateral
margins continuously rounded, notched; second pair of lobes
broader, deeper lateral notch; ithird pair of lobes bluntly
pointed, triangular, lateral margin serrate; pectinae narrow
distapectinae, arranged 2, 2, 2, 2, those of each lateris distant
from third lobe width of lobe; each lateris coarsely crenulate.
— Europe on Laurel. lauri Bouche.
cc. Pygidium of aduHt female with median pair of lobes separated
by pair of long, slender, plate-like pseudolobes, twice as long
as lobes; median and second pairs of lobes long, slender, sides
parallel, two or three times as long as wide; second incisura
with long, broad, dagger-like plate and one or two cephalad
of each second lobe; plate-like projections variable in size
TRIBE ASPIDIOTINI 465
and form; setae long, three on each side, one near to and
cephalad of each lobe, and near middle of each lateris; anus
located near cephalic part of pygidium; margin of segments
of preabdomen distinctly toothed. — Java on Myristica
javanensis Green.
SPECIES OF XEROPHILASPIS
Pygidium of adult female with two pairs of small subequal lobes, broader
than long, margin rounded, entire; median incisura normal, one-half
width of second; pectinae or plates wanting; setae minute, shorter than
lobes; ceratubae small, nolt abundant; anus four times its width from
caudal margin and once from vulva; calles obsolete; pygidium of
second nymphal female with median pair of lobes large, subtruncated,
angles rounded, as long as broad, protuberant, entire; second pair of
lobes similar in form to median, smaller; third pair of lobes four or
five times as broad as long, mesal margin straight, lateral margin
oblique, emarginalte, slightly crenulate; plates small, shorter than
lobes, arranged 0, 1, 1, 0; anus over eight times its width from caudal
margin. — Arizona on Prosopis. prosopidis Ckll.
SPECIES OF PHAULASPIS
a. Pygidium of adult female transverse, truncated, hardly projecting
beyond the margin; preabdomen fringed with six small equidistant
setae, minutely serrate; pygidium of second nymphal female with
single pair of lobes; median pair of lobes elongaJte with slightly
converging sides, bluntly pointed, entire, distant; pectinae or plates
wanting; each lateris with three deep indentations limiting areas
with prominent caudal angles; each median lobe and each of two
caudal projedtions bearing two setae; mesospiracerores present. —
Australia on Hakea. hakeae Mask.
aa. Pygidium of adult female transverse, truncated, distinctly project-
ing beyond the margin, entire, deep mesal incision; lobes, pectinae
or plates, and setae wanting; pygidium of second nymphal female
with four simple lobes, scarcely projecting, extending cephalad
paraphysis-like, adjacenit, inconspicuous; cephalad of lobes two large
deep inconspicuous clefts; pectinae or plates wanting; pygidium
of first nymph with two lobes with single deep cleft on each side. —
Gymnaspis grandis Green. — Seychelles on Lodoicea.- -grrandis Green.
SPECIES OF ANCEPASPIS
Pygidium of adult female strongly narrowed at middle; lobes and
pectinae or plates wanting; caudal margin with two deep, narrow,
parallel emarginaitions, extending for one-half length of narrowed por-
tion; mesal portion between emarginations with rounded angles,
bearing single seta on each side near caudal end of emarginations;
each lateral portion bordering emargination narrower than mesal,
angulate at caudal end; anus about eighteen times its width from
caudal margin and once from vulva; body cephalad of constricted
portion broadly rounded; pygdidium of second nymphal female rounded,
entire, two undulations where median lobes usually located with
emargination between; setae minute, more numerous than in adult;
anus six times its width from caudal margin; lobes and pectinae or
plates wanting; pygidium of first nymph without lobes, short setae,
and anal setae. — Arizona on Prosopis tridentata Ferris..
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470 THE COCCIDAE
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472 THE COCCIDAE
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Japan. Ent. Month. Mag., 33, 1897, 239-244.
On a new species of coccid on fern-roots. Proc. Lin. Soc.
New South Wales, 8, 1894, 225-226; pi. 7.
Matheson, R. — Number of moults of the female Dactylopius citri.
Can. Entom., 39, 1907, 284-287.
Maxwell-Lefroy, H. — Notes on Indian scale insects (Coccidae).
Mem. Dept. Agr. India., Entom., 2, 1908, 111-137; pis. 10-12.
Mayet, V. — La cochenille des vigues du Chili (Margarodes vitium
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12, 1907, 1-26; pis. 1-4.
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Daven. Acad. Sci., 2, 1879, 293-347; pis. 12-13.
Riley, C. V. — The cottony cushion-scale. Rept. U. S. Dept. Agr.
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Sanders, J. G. — The Coccidae of Ohio, I. Proc. Ohio State Acad.
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The identity and synonomy of some of our soft scale-
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m
THE COCCIDAE
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CHANGES
The discovery that certain names used on previous pages were
preoccupied was not made until too late to insert other names in their
place. The following need to be changed: —
Douglasia to Douglariella
Greenella to Greenacoccus
Paragreenia to Neogreenia
Hemiaspis to Hemaspidis
INDEX
Abdomen, 35, 221
Abdominal cerari, 122
Abdominal setae, 163
Abdominal spiracles, 37, 108
Aberemoae, 254
Aberrans, 279
Abietis, 272, 399
Acaciae, Ceroplastodes, 178
Acaciae, Coccomytilus, 294
Acaciae, Epicoccus, 104
Acaciae, Gymnaspis, 256
Acaciae, Neomorgania, 458
Acaciae, Trullifiorinia, 376
Acericola, 176, 177
Aclerda, 175, 182
Actenaspis, 263
Actenaspis, Species of, 267
Active period, 8
Acuminata, 339
Adenostomae, 145
Adiopugnatorische organ, 123
Adiscodiaspis, 313
Adiscodiaspis, Species of, 366
Adiscofiorinia, 372
Adiscofiorinia, Species of, 378
Adonidum, 137
Aechmeae, 257
Aegyptiaca, 73
Aesculi, 414
Affinis, 266
Affirmaspis, 393
Affirmaspis, Species of, 448
Africana, Adiscofiorinia, 378
Africana, Diaspis, 319
Africana, Gramenaspis, 354
Africana, Gymnaspis, 255
Africana, Monophlebus, 71
Africana, Neosignoretia, 425
Africanus, Hendaspidiotus, 439
Africaspis, 307
Africaspis, Species of, 338
Agavis, 421
Agavium, 141, 142
Agonis, 335
Agrifoliae, Essigaspis, 324
Agrifoliae, Pseudococcus, 137
Akermes, 175, 182
Alaeodendri, 386
Alatae, 340
Alate setae, 201
Alatus, 386
Albata, 176
Albataspis, 275
Albataspis, Species of, 290
Albida, 154
Albiventer, fernaldi, 410
Albizzae, 343
Albolutea, 73
Albopictus, 418
Albus, Coccomytilus, 293
Albus, juglans-regiae, 409
Alecanopsis, 173, 178
Aleyrodidae, 5
Alichtensia, 171, 178
Alienus, 418
Allan tomytilus, 276
Allantomytilus, Species of, 295
Allaudi, Amelacoccus, 189
Allaudi, Aspidiotus, 386
Alni, Hendaspidiotus, 440
Alni, Targionia, 448
Alni, Xylococcus, 86
Altaceratubae, 224, 301
Alveolae, 163
Amaniensis, 359
Ambigaspis, 394
Ambigaspis, Species of, 456
Ambigua, Ceroputo, 144
Ambigua, Scrupulaspis, 288
Ambigua, Unachionaspis, 337
Amelococcus, 188, 189
Americana, Fundaspis, 338
Americana, Orthezia, 113
Americanae, Phenacoccus, 142
Americanus, Trionymus, 138 •
Americoccus, 78
475
476
THE COCCIDxiE
Ametrochaspis, 311
Ametrochaspis, Species of, 359
Amorphococcus, 147, 189, 190
Ampelodesmae, 280
Amygdali, .176
Anacerores, 27, 84
Anal aperature, 153
Anal cerari, 121
Anal chalazae, 64
Anal cleft, 160
Anal Lobe Possessors, 48
Anal lobes, 38, 166, 186, 227
Anal operculum, 161
Anal plate, 109, 161, 222
Anal process, 151
Anal ring, 3 8, 161
Anal Ring Conservers, 49
Anal Ring Loosers, 49
Anal ring setae, 38, 161, 186
Anal Ring Type, 47
Anal setae, 166, 186
Anal spine, 151
Anal tube, 36, 84, 161, 186
Anamaspis, 263
Anamaspis, Species of, 267
Anamefiorinia, 372
Anamefiorinia, Species of, 377
Ancepaspis, 395
Ancepaspis, Species of, 465
Ancylus, 411
Andaspis, 275
Andaspis, Species of, 292
Andreae, 74
Andrei, 196
Angraeci, 216
Angulata, 255
Angusta, 351
Annae, 113
Anomala, 365
Anomalacoccus, 187, 189
Anomaladensae, 235
Anonae, 441
Anoplura, 3
Ansei, 415
Antennae, 30, 158, 221
An|;ennal formula, 30, 158
Antennata, 144
Anterior group, 23 6
Anterior-lateral group, 236
Anterior laterals, 236
Antonia, 132, 145, 211
Antonia, Species of, 146
Anus, 36, 162, 233
Aonidia, 229, 395
Aonidia, Species of, 462
Aonidiella, 392
Aonidiella, Species of, 442
Aonidiformis, 252
Aonidoides, 438
Aonidomytilus, 275
Aonidomytilus, Species of, 292
Aonidum, 416
Aperature sessuale feminae, 233
Apex, 161
Aphididae, 5
Aphylonis, 133
Apical setae, 163
Apicata, 441
Apiococcus, 210, 211
Apiomorpha, 201, 204
Apiomorphinae, 54, 57, 61
Apiomorphinae, Subfamily, 199
Apiomorphinae, Genera of, 204
Apodema, 33
Apteronidia, 247
Araucariae, 145
Arctorthezia, 111, 113
Arctorthezia, Species of, 114
Arctostaphyli, 404
Ardisiae, 114
Arecae, dictyospermi, 416
Arenae, 144
Arenosus, 145
Argentata, Lichtensia, 175
Argentata,' Niveaspis, 296
Argentina, Hemiberlesia, 436
Argentina, Pulvinaria, 176
Argentina, Tacbardiella, 154
Argentosis, 356
Ariditalis, 182
Arii, 280
Arizonensis, 141
Arizonica, 450
Armatum, Erium, 140
Armatus, Aspidoproctus, 74
Armatus, Eriococcus, 145
Armatus, Pseudokermes, 178
Armored scales, 217
INDEX
477
Arrangement, pectinae, 229
Artemisiae, Eriococcus, 145
Artemisiae, Orthezia, 113
Artemisiae, Phenacoccus, 143
Arthrocnemi, 344
Articulatus, 452
Artocarpi, 458
Arundinariae, 353
Ascelis, 2 01, 204
Asper, 74
Asperatus, 211
Aspidiella, 3 87
Aspidiella, Species of, 403
Aspidiotini, 244
Aspidiotini, Genera of, 387
Aspidiotini, Tribe, 380
Aspidiotus, 387
Aspidiotus, Species of, 396
Aspidistrae, 343
Aspidoides, 387
Aspidoides, Species of, 406
Assimilis, Ultracoelostoma, 87
Assimilis, Unachionaspis, 337
Aspidoproctus, 70, 74
Aster, 141
Asterolecaniinae, 56, 61
Asterolecaniinae, Genera of, 187
Asterolecaniinae, Subfamily, 183
Asterolecanium, 188, 189
Asymmetraspis, 311, 360
Atalantiae, Adiscofiorinia, 379
Atalantiae, Syngenaspis, 252
Atlantica, 322
Atherospermae, 395
Atripennis, 71
Atriplicia, 131, 145
Atriplicis, 364
Attenuata, 178
Aucubae, 346
Augulaspis, 309
Augulaspis, Species of, 353
Aulacaspis, 305
Aulacaspis, Species of, 316
Aurantia, 141
Auranticolor, pentagona, 315
Aurantii, 443
Auratilus, 3 62
Aurescens, 145
Aureum, 189
Auriculata, 282
Aurilanatus, 135
Austini, 197
Australis, Callipappus, 98
Australis, Diaspis, 320
Australis, Palaeococcus, 72
Austriaca, 330
Austro-africana, 461
Austrolichtensia, 169, 175
Auxiliary setae, 122
Axin, 71
Azaleae, 145
Aztectus, 178
Baccharidis, 321
Bahiae, Ceroputo, 144
Bahiae, Eriococcus, 145
Baikeae, 457
Bakeraspis, 395
Bakeraspis, Species of, 459
Bambusae, Asterolecanium, 189
Bambusae, Chaetococcus, 147'
Bambusae, Chionaspis, 304
Bambusae, Ischnafiorinia, 378
Bambusae, Leucaspis, 264
Bambusae, Pinnaspis, 290
Bambusarum, 424
Bambusicola, 294
Banksiae, Aonidia, 462
Banksiae, Triaspidis, 277
Banksiae, Syngenaspis, 249
Barberi, Ceroputo, 144
Barberi, Pseudaulacaspis, 316
Barbusano, 448
Barrancorum, 360
Basal thickenings, 235
Base, 161
Basement membrane, 25
Bavaricus, 410
Berlese, 162, 225
Berlese and Leonardi, 42
Berlesaspidiotus, 389
Berlesaspidiotus, Species of, 423
Berlesaspis, 274
Berlesaspis, Species of, 289
Berlesii, 352
Betheli, 144
Betulae, 86
Beyeriae, 296
478
THE COCCIDAE
Biafrae, 459
Biclavis, 363
Bicornis, Termiticoccus, 141
Bicornis, Triaspidis, 277
Bidens, Chorizaspidiotus, 434
Bidens, Fiorinia, 373
Biformis, 407
Bigeloviae, Leonardianna, 450
Bigeloviae, Pulvinaria, 176
Bilobis, 333
Bilobis, 386
Bipartita, 178
Bipindensis, 291
Bituberculatum, 179
Blanchardii, Ripersia, 141
Blanchardi, Parlatoria, 253
Blankenhorni, 304
Boguei, 197
Bohemicus, 142
Boisduvalii, 319
Borealis, 145
Bornmulleri, 428
Bossiae, 386
Boutelouae, 146
Bouvari, 71
Bouvieri, 74
Bowreyi, 422
Brachypodii, 176
Brachyscelis, 204
Bracteae, 224
Braggi, 141
Brainaspis, 390
Brainaspis, Species of, 427
Branagani, 198
Brasilensis, Cryptokermes, 87
Brasiliensis, Eriococcus, 145
Brasiliensis, Icerya, 73
Brasiliensis, Megasaissetia, 182
Braziliensis, Palaeococcus, 72
Brevaceratubae, 224, 302
Brevipes, 133
Britannica, Kuwania, 79
Britannicus, Aspidiotus, 402
Broadwayi, 176
Bromeliae, Aonidella, 444
Bromeliae, Diaspis, 321
Bromeliae, Pseudococcus, 133
Bromii, 140
Bruneri, 177
Bufo, 98
Bulla, 223, 225
Bullata, 256
Bupleuri, 350
Burmeisteri, 74
Bussi, 348
Buxi, 290
Buzenensis, 284
Cacti, Coccus, 103
Cacti, Llaveia, 71
Caerulea, 154
Caffra, 333
Calceolariae, 140
Calcitectus, 144
Caldesii, 386
Californica, Aclerda, 182
Calif ornica, Orthezia, 113
Calif ornicus, Aspidiotus, 396
Californicus, Trionymus, 140
Calles, 235
Callipappinae, 53, 56, 59
Callipappinae, Subfamily, 94
Callipappus, 98
Callococcus, 147, 188, 189
Calurna, 441
Calyptroides, 320
Calyx, 27
Camelicola, 176
Camelliae, Fiorinia, 367
Camelliae, Hemiberlesia, 435
Campylanthi, Targionidea, 449
Canaliculata, Lepidosaphes, 2 80
Canaliculata, Circulaspis, 449
Canariensis, Chorizaspidiotus, 434
Canariensis, Duplachionaspis, 332
Canaspis, 308
Canaspis, Species of, 352
Candida, Duplachionaspis, 332
Candida, Icerya, 73
Candida, Phenacaspis, 352
Candidula, 412
Canella, 151, 159
Capparis, 335
Capensis, Coccus, 104
Capensis, Dentachionaspis, 358
Capensis, fimbriatus, 402
Capensis, Margarodes, 93
Capensis, Truncaspidiotus, 431
INDEX
479
Capreae, 180
Capsulatus, 451
Capulina, 210, 211
Caricis, 175
Cardiococcus, 170, 177
Carinata, 292
Carissae, 356
Carpochloroides, 210, 211
Carteria, 149
Caruleli, Diaspis, 313
Carulaspis, 305
Carulaspis, Species of, 313
Caryae, Eulecanium, 179
Caryae, Fundaspis, 338
Cassiae, 339
Cassinae, 284
Castaneae, 140
Castilloae, 181
Casuarinae, Anamefiorinia, 378
Casuarinae, Cylindrococcus, 211
Casuarinae, Frenchia, 190
Casuarinae, Ourococcus, 211
Casuarinae, Poliaspis, 304
Casuarinae, Scrupulaspis, 288
Casuarinae, Sphaerococcus, 147
Casuarinae, Targaspidiotus, 447
Catalinae, 145
Cataphracta, 114
Cattleyae, biformis, 407
Caudalabiae, 123
Caudal postanal setae, 125
Caudal setae, 186
Caudo-lateral group, 236
Caudolateral group, 236
Caudo-lateral margin, 161
Caustic potash, 15
Cavaera, 37, 124, 159
Cecconi, 425
Cocropiae, 145
Cedri, 447
Celastri, 451
Cells, 163
Cellulae, 163
Cellular areas, 163
Celtidis, Lecaniodiaspis, 189
Celtidis, Diaspis, 304
Cephalabiae, 123
Cephalaspis, 274
Cephalaspis, Species of, 286
Cephalic postanal setae, 125
Cephalo-lateral group, 236- -
Cephalolateral group, 236
Cephalo-lateral margin, 161
Cephalothorax, 28
Ceraran setae, 122
Cerari, 121
Cerarii, 121
Cerata, 446
Ceratoniae, Lepidosaphes, 272
Ceratoniae, Phenacoccus, 350
Ceratubae, 26, 185, 193, 223, 225
385
Ceravitreous Coccids, 183
Ceriferus, 196
Cerococcus, 187, 189
Ceronema, 169, 176
Ceroplastes, 165, 170, 177
Ceroplastidia, 177
Ceroplastina, 177
Ceroplastodes, 172, 178
Ceroputo, 130, 144
Cerores, 26, 230
Cevalliae, 142
Chaetachmae, 334
Chaetococcus, 133, 147
Chalaza, 64
Chamaeropsis, 386
Championi, 72
Charmoyi, 430
Cheilanthi, 113
Chemnaspidiotus, 391
Chemnaspidiotus, Species of, 439
Chenopodii, 434
Chentraspis, 391
Chentraspis, Species of, 434
Chilaspidis, 178
Chilensis, 304
Chilina, 313
Chilopsidis, 278
Chinensis, 254
Chionaspiformis, 339
Chionaspis, 299, 307
Chionaspis, Species of, 324
Chitinosa, Mytilaspis, 272
Chitinosus, Coccomytilus, 294
Chitinous cradle, 32
Chorizaspidiotus, 391
Chorizaspidiotus, Species of, 432
480
THE COCCIDAE
Chrysomphalus, 388
Chrysomphalus, Species of, 414
Cicatrices, 123, 240
Cicatrix, 240
Cinerea, 249
Cingala, 250
Cinnamomi, Aulacaspis, 317
Cinnamomi, Proceraspls, 363
Circulaspis, 393
Circulaspis, Species of, 449
Circumferential lamellae, 110
Circumgenital gland openings,
236
Circumgenital gland orifices, 236
Circumgenital glands, 163, 236
Cirripediformis, 177
Cisanal setae, 125
Cissococcus, 131, 145
Cistudiformis, 177
Cistuloides, 451
Citricola, Lecanium, 178
Citricola, Takahashia, 177
Citrina, aurantii, 443
Citri, Prontaspis, 359
Citri, Pseudococcus, 134
Citrophilus, 138
Cladii, 442
Clarifying, 15
Classification, 40
Clavaceratubae, 224
Clavaspis, 391
Clavaspis, Species of, 441
Clavate setae, 158
Claviger, Duplaspidiotus, 453
Claviger, Pseudococcus, 133
Clavus, 159
Clearing, 20
Club-shaped thickenings, 232
Clypeococcus, 74
Clypeus, 159
Cobbii, 211
Coccidae, 5
Coccidae, Subfamilies of, 52
Adult females, 57
Female nymphs, 55
First nymphal stage, 52
Coccidae, Synopsis of the, 45
Coccinae, 53, 56, 58
Coccinae, Genera of, 102
Coccinae, Subfamily, 99
Coccineus, 145
Coccomytilus, 276
Coccomytilus, Species of, 292
Cocculi, 286
Coccura, 129, 142
Coccus, 103
Coccus, Species of, 103
Cochineal Insects, 99
Cockerell, 41, 42
Coclcerellae, .Ripersia, 141
Cockerellaspis, 306
Cockerellaspis, Species of, 318
Cockerellella, 71, 75
Cockerelli, Kermes, 197
Cockerelli, Leucaspis, 267
Cockerelli, Phenacaspis, 345
Cockerelli, Phenacoccus, 142
Cockerelli, Pseudococcus, 133
Cockerelli, Pulvinaria, 176
Cocotiphaga, 408
Coelostomidea, 86
Colemani, Phenacoccus, 143
Colemani, Unachionaspis, 337
Colimensis, 175
Colinensis, 73
Colobopyga, 154
Coloratus, 397
Coloradensis, Eriopeltis, 176
Coloradensis, Physokermes, 182
Colvei, 304
Comari, 142
Compacta, 74
Comperei, 406
Compressis, 86
Comstock, 40
Comstockaspis, 391
Comstockaspis, Species of, 438
Comstocki, Aspidiella, 405
Comstocki, Pseudococcus, 135
Comstockiella, 388
Comstockiella, Species of, 423
Conchaspinae, 54, 57, 61
Conchaspinae, Genera of, 215
Conchaspinae, Subfamily, 212
Conchaspis, 213, 215
Conchaspis, Species of, 215
Concinnum, 366
Concinulus, 198
INDEX
481
Concolor, 292
Condaliae, 361
Confusus, Aspidiotus, 386
Confusus, Coccus, 104
Coniferarum, 439
Conjunctiva, 1
Conservans, 357
Conspicus, 433
Contigaspis, 309
Contigaspis, Species of, 354
Contrahens, 74
Convexa, Protopulvinaria, 176
Convexus, Coccomytilus, 294
Cooley, 302
Cooleyaspis, 308
Cooleyaspis, Species of, 353
Cooleyi, 176
Coralinus, 386
Cordaliae, 154
Cordiae, 304
Cordylinidis, Pusilaspis, 289
Cordylinidis, Leucaspis, 264
Coriae, 1, 10, 222
Corni, Chionaspis, 331
Corni, Eulecanium, 179
Cornigera, 459
Cornuaspis, 274
Cornuaspis, Species of, 286
Cornuparvum, 178
Corokiae, 406
Coronaspis, 312
Cornaspis, Species of, 362
Coronifera, 362
Cornuta, 154
Corpulenta, 74
Corrugata, 283
Corticis, 189
Corticis-pini, 406
Cortrioides, Mytilaspis, 272
Corticoides, subrubescens, 396
Corticosus, 419
Coryphae, 401
Coulteri, 176
Coursetiae, 413
Covilleae, 423
Coxacoila, 33
Coxartis, 33
Cradle, 159
Crassa, 272
Crateraformis, 211
Crawfordi, 74
Crawii, Antonia, 146
Crawii, Lepidosaphes, 280 ~ ~
Crawii, Pseudaulacaspis, 314
Crawii, Pseudococcus, 134
Credodiaspis, 313
Credodiaspis, Species of, 366
Cremastogastri, 189
Crenulaspidiotus, 389
Crenulaspidiotus, Species of, 426;
Crenulata, 257
Crenulate, 228
Crescentiae, 175
Cribiform plates, 186
Cristata, 313
Cristicola, 190
Crocea, 73
Crotonis, 295
Crudiae, 278
Crumena, 32, 218
Cryptaspidus, 309
Cryptaspidus, Species of, 357
Cryptaspidotus, 389
Cryptaspidiotus, Species of, 42 S
Cryptes, 167, 174, 181
Crypthemichionaspis, 372
Cryptinglisia, 171, 178
Cryptococcus, 132, 147
Cryptodiaspis, 309
Cryptodiaspis, Species of, 357
Cryptokermes, 86, 87
Cryptomeriae, 400
Cryptoparlatoria, 248
Cryptoparlatoria, Species of, 253
Cryptophyllaspis, 390
Cryptophyllaspis, Species of, 42T
Cryptoripersia, 128, 141
Cryptoxanthus, 411
Ctenochiton, 165, 171, 178
Cualatensis, 133
Cuculus, 433
Cueroensis, 433
Cupaniae, 176
Cupidaspis, 312
Cupidaspis, Species of, 363
Cupressi, Cupidaspis, 363
Cupressi, Ehrhornia, 144
Cupressi, Hendaspidiotus, 440
482
THE COCCroAE
Cupressi, Puto, 144
Cupressicolus, 138
Curculiginis, 451
Cuticle, 25
Cuticular plates, 152
Cuticular tubes, 223
Cyanogena, 341
Cyanophylli, 407
Cycadis, 356
Cydoniae, Aspidiotus, 412
Cydoniae, Tachardiella, 154
Cyllndrococcinae, 53, 56, 59
Cylindrococcinae, Genera of, 210
Cylindrococcinae, Subfamily, 205
Cylindrococcus, 206, 210, 211
Cyst, 90
Cystococcus, 202, 204
Dalbergiae, 75
Daleae, 178
Darutyi, 453
Darwiniensis, japonica, 266
Dasylirii, 133
Dearnessi, Phenacoccus, 144
Dearnessi, Remotaspidiotus, 434
Decurvata, 348
Defecta, 2 88
Degeneratus, 419
Dehydration, 20
Deltoid Scales, 380
Dendrobii, 189
Dendropthorae, 176
Densariae, 227
Densifloriae, 405
Dentacerores, 125
Dentachionaspis, 310
Dentachionaspis, Species of, 358
Dentaspis, 312
Dentaspis, Species of, 362
Dentata, 460
Denticulate pores, 125
Dentiloba, Pseudodiaspis, 364
Dentilobis, Remotaspis, 360
Depressa, Chionaspis, 304
Depressa, Pulvinaria, 176
Dermal cells, 163
Dermal plate, 187
Dermal pores, 163
Destefanii, 289
Destructor, Aspidiotus, 396
Destructor, Cryptoparlatoria, 253
Destructor, Pseudococcus, 134
Diaspicera, 225
Diaspidinae, 55, 57, 61
Diaspidinae, Subfamily, 217
Diaspidinae, Tribes of, 243
Diaspidini, 244
Diaspidini, Genera of, 304
Diaspidini, Tribe, 297
Dispinae, 300
Diaspini, 300
Diaspis, 299, 306
Diaspis, Species of, 319
Diaspidistis, 305
Diaspidistis, Species of, 314
Diaspiformis, 374
Diaspidiotus, 388
Diaspidiotus, Species of, 411
Dictyospermi, 416
Difficilis, 327
Diffinis, 437
Digitules, 159
Dilata, 351
Dimorphus, 75
Dinaspis, 276
Dinaspis, Species of, 295
Diosmae, 364
Discaloca, 65
Discal setae, 163
Disculi ciripari, 236
Disculi ciripari perivulvarea, 236
Dishi ciripari, 236
Dispar, 272
Distapectinae, 230
Distichlii, 139
Disticlium, 147
Distinctissima, 256
Distorta, 360
Divergens, 334
Diversispinus, 145
Dorsal gland orifices, 223
Dorsal glands, 223
Dorsal lamellae, 109
Dorsal lip, 186
Dorsal ostioles, 123
Dorsal pilacerores, 109
Dorsal plates, 109
Dorsal pores, 223
INDEX
483
Dorsal scale, 239
Dorsal setae, 122, 231
Dorsal tubercles, 158
Dorsal tubular spinnerets, 223
Double-shielded Scales, 297
Doughy wax, 164
Douglasia, 110, 111, 474
Douglasiella, 474
Dracaenae, 340
Drimydis, 293
Drosicha, 70, 74
Dryandrae, Syngenaspis, 251
Dryandrae, Targaspidiotus, 447
Dubius, Monophlebus, 71
Dubius, Eriococcus, 145
Dugesi, Monophlebus, 71
Dugesii, Ceroplastodes, 178
Duplachionaspis, 307
Duplachionaspis, Species of, 332
Duplaspidiotus, 394
Duplaspidiotus, Species of, 452
Duponti, 2 86
Dura, 361
Dysoxyli, Aspidiotus, 386
Dysoxyli, Chionaspis, 331
Echancrueres, 226
Echinata, 463
Echiniformis, 204
Edentata, 365
Edwallia, 171, 178
Eglandulosus, 386
Ehretiae, 449
Ehrhornia, 130, 144
Ehrhorni, Diaspidiotus, 413
Ehrhorni, Mycetococcus, 189
Ehrhorni, Pulvinaria, 176
Ekebergiae, lounsburyi, 349
Elaeagni, 329
Elaeidis, 397
Elastica, 144
Elegans, 435
Elongata, Cryptophyllaspis, 428
Elongata, Greenaspis, 339
Elongata, Triaspidis, 277
Elongated pores, 223
Elongate thickenings body-wall,
232
Eloti, 141
Endoskeleton, 32, 159, 221
Ensign Coccids, 105
Entaspidiotus, 394 ^
Entaspidiotus, Species of, 454
Entire, 228
Ephedrae, Genaparlatoria, 255
Ephedrae, Leucaspis, 265
Ephedrae, Philephedra, 176
Ephedrae, Pseudococcus, 134
Ephedrarum, 429
Epicoccus, 103, 104
Epidendri, 189
Epidiaspis, 306
Epidiaspis, Species of, 318
Epigaeae, 410
Epiphytidis, 2 89
Episternum, 33
Equipment, 12
Ericerus, 167, 174, 181
Ericicola, 3 66
Eriochitin, 168, 175
Eriococcinae, 54, 57, 60
Eriococcinae, Genera of, 127
Eriococcinae, Subfamily, 118
Eriococcus, 131, 145
Eremicoccus, 147
Eriogoni, Eriococcus, 145
Eriogoni, Erium, 140
Eriogoni, Phenacoccus, 144
Eriogoni, Pseudococcus, 138
Eriopeltis, 169, 176
Erium, 140
Erythraspidis, 420
Erythrocephala, 93
Essigaspis, 306
Essigaspis, Species of, 324
Essigi, 197
Etheliae, 335
Etrusca, 332
Eucalymnatus, 174, 181
Eucalypti, Mytilaspis, 272
Eucalypti, Neomorgania, 458
Eucalypti, Olliffia, 190
Eucalypti, Ourococcus, 211
Eugeniae, 176
Eulecanium, 173, 180
Eulecanium, Species of, 179
Euleucaspis, 262
Euonymi, Aspidotus, 435
484
THE COCCIDAE
Euonymi, Chionaspis, 325
Euonymi, theae, 250
Euparlatoria, 247
Euphillipia, 168, 175
Euphorbiae, Duplachionaspis, 335
Euphorbiae, Entaspidiotus, 455
Euphorbiae, Walkeriana, 74
Euphoriae, Chionaspis, 304
Euryae, 285
Ewarti, 72
Exaeretopus, 168, 175
Exalbida, 333
Excisus, 403
Exocarpi, 355
Expansa, 379
Extensa, 446
External anatomy, 24
Exuvia, 239
Exuviae, 239
Exuviae, position, 240
Exuvialess Scales, 212
Exuvium, 239
Eye-like glands, 123
Eyes, 29
Fabernii, 438
Fabianae, 448
Fagi, 147
Fagisuga, 213, 215, 216
Falcifer, 141
Fall ax, 175
Farinosus, 98
Fasciata, 281
Fasciculeusis, 78
Fast method, 17
Females, 9
Fernald, 43
Fernaldanna, 276
Fernaldanna, Species of, 296
Fernaldi, 411
Ferox, 75
Ferreae, 461
Ferrisaspis, 388
Ferrisaspis, Species of, 422
Festucae, Eriopeltis, 176
Festucae, Trionymus, 140
Fici, Duplachionaspis, 334
Fici, Hemichionaspis, 340
Ficifolii, 283
Ficus, Lepidosaphes, 286
Ficus, Pulvinaria, 176
Fifth incisura, 227, 228
Fifth lobe, 22 8
Fifth pair lobes, 228
Filamentosus, 133
Filicum, Alecanopsis, 178
Filicum, Saissetia, 181
Filiere, 121
Filiere aggregate, 236
Filiere isolate, 223
Filiere separee, 223
Filieres, 236
Filieres isolee, 223
Fillipia, 169, 176
Filuri, 121
Fimbriata, 216
Fimbriate plates, 22 9
Fimbriatula, 141
Fimbriatum, Asterolecanium, 189
Fimbriatus, Aspidiotus, 402
Fiorinia, 367, 372
Fiorinia, Species of, 372
Fioriniae, 374
Fiorineides, 431
Fioriniini, 244
Fioriniini, Genera of, 372
Fioriniini, Tribe, 367
First exuvia, 238
First incisura, 226
First lateral lobe, 228
First nymphal stage, 219
Fissidens, 428
Fissus, 429
Flabellae, 158
Flabelliform marginal hairs, 158
Flacourtiae, 316
Flava, Ametrochaspis, 359
Flaveola, Ripersia, 141
Flaveolus, Lecanium, 178
Flavus, Ctenochiton, 178
Fletcheri, 179
Floccifera, 176
Floccosa, 114
Florenciae, 399
Floridensis, 177
Floriger, 74
Fluminiensis, 216
Fodiens, Chrysomphalus, 417
INDEX
485
Fodiens, Nudachaspis, 366
Fodiens, Ortheziola, 114
Fonscolombia, 129, 141
Forbesaspis, 388
Forbesaspis, Species of, 422
Forbesi, 422
Formicarii, Pseudococcus, 133
Formicarius, Stictococcus, 75
Formicarum, Margarodes, 93
Formicarum, Newsteadiella, 75
Formiceticola, 175
Formosa, Phenacaspis, 352
Formosus, Trichomytilus, 276
Fortis, 72
Fossette ostioliform labiate, 123
Fossor, 457
Fourth lobe, 228
Fourth incisura, 227, 22 8
Fourth pair lobes, 22 8
Foveola labiate, 122
Fraxini, 141
Frayed setae, 158
Frenchi, 333
Frenchia, 189, 190
Fringe, 222
Fringed plates, 229
Fringe scales, 245
Fringe setae, 162
Froggatt, 149, 201
Froggattiella, 3 93
Froggattiella, Species of, 450
Frons, 28
Fulgens, 154
Fulleri, Cissococcus, 145
Fulleri, Natalensia, 141
Fulleri, Pseudaulacaspis, 315
Fulvoradiata, 154
Fundaspis, 307
Fundaspis, Species of, 337
Funtumiae, 35 8
Furca, 33, 66
Furcate plates, 229
Furcate setae, 158
Furcaspis, 388
Furcaspis, Species of, 406
Furcapectinae, 230
Furchadaspis, 310
Furchadaspis, Species of, 358
Furcillae, 396
Furcraeicolus, 429
Furfura, 331 __
Fuscipennis, 72
Fuscum, Globulicoccus, 180
Fuscus, Monophlebulus, 75
Fuse marginales, 223
Fusi, 223, 236
Fusilaspis, 275
Fusilaspis, Species of, 289
Fusi piliformis, 230
Fusi spiniformis, 230
Gage's Saurefuchsin, 20
Gallica, 93
Gallicola, 145
Galliformens, 354
Galliformens, allaudi, 386
Galliformis, 196
Ganogene cells, 225
Garryae, 112
Gascardia, 154
Gelonii, saprosmae, 377
Gemmifera, 154
Genacerores, 236, 303
Genacerores confluent, 237
Genacerores, formula, 236
Genaparlatoria, 248
Genaparlatoria, Species of, 255
Generalized Coccidae, 46
Generalized Diaspidinae, 241
Genistae, 73
Genital aperature, 233
Genital orifice, 233
Gennadii, 315
Geococcus, 129, 142
Ghiandole ganogene, 225
Ghiandola sericipara, 225
Giant Coccids, 62
Gidgei, 386
Gigas, 264
Gilletti, Eriococcus, 145
Gilletti, Kermes, 196
Glandiferous spines, 106
Glands of spiracles, 159
Gland spines, 230
Glandular pores, 159
Glanduliferus, 411
Glandulosa, 426
Glassy, 165
486
THE COCCIDAE
Gleditsiae, 328
Globosa, Maskellia, 446
Globosum, Erium, 140
Globosus, Apiococcus, 211
Globosus, Unachionaspis, 337
Globulicoccus, 180
Glomerata, 445
Glomerella, 154
Gloverii, 283
Gnidii, 145
Gonaspidiotus, 390
Gonaspidiotus, Species of, 431
Gorodetskia, 79
Gossyparia, 131, 145
Gossypii, 142
Gowdeyi, 425
Gracilis, Cylindrococcus, 211
Gracillis, Entaspidiotus, 455
Gracilis, Eucalymnatus, 181
Graminellus, 432
Gramenaspis, 309
Gramenaspis, Species of, 353
Graminis, Duplachionaspis, 334
Graminis, Ehrhornia, 144
Graminis, Orthezia, 112
Graminis, Rugaspidis, 449
Grandilobis, 288
Grandis, Kermes, 196
Grandis, Phaulaspis, 465
Grandis, Pulvinaria, 176
Graphaspis, 310
Graphaspis, Species of, 359
Green, 42, 89, 150, 238, 302
Greenacoccus, 474
Greenaspis, 307
Greenaspis, Specis of, 339
Greeniella, 70, 75, 474
Greeniella, 395
Greeniella, Species of, 459
Greeni, Entaspidiotus, 455
Greeni, Mytilaspis, 272
Greeni, rossi, 421
Greeni, Syngenaspis, 250
Greenoidea, 392
Greenoidea, Species of, 446
Gregarius, 211
Grindeliae, 139
Griqus, 456
Grisea, 288
Grossulariae, 374
Ground Pearls, 88
Grouped glands, 236
Grouped orifices spiracles, 159
Guerinella, 69, 72
Guerinii, 74
Guilding, 89
Gularostria, 4
Gularostria, Families of, 4
Guterriziae, Chorizaspidiotus, 432
Gutierreziae, Pseudococcus, 138
Gymnaspis, 247, 248
Gymnaspis, Species of, 255
Gymnococcus, 129, 141
Gymnococcus, Species of, 142
Gymnosporiae, 403
Haematochrous, 430
Hairs, 232
Hakeae, 465
Hakearum, 175
Halimicoccus, 211
Hamata, 358 :
Hameli, 93
Hardening, 19
Hartii, 404
Hastata, 251
Hawaiiensis, 292
Head, 28
Head cerari, 121
Head pilacerores, 109
Head plates, 109
Head skeleton, 32
Hederae, 400
Hedyotidis, 327
Helianthi, Phenacoccus, 144
Helianthi, Rhizaspidiotus, 431
Heliococcus, 130, 142
Hellenicus, 72
Hemaspidis, 474
Hemiaspis, 275, 474
Hemiaspis, Species of, 291
Hemiberlesia, 391
Hemiberlesia, Species of, 434
Hemichionaspis, 308
Hemichionaspis, Species of, 340
Hemichionaspiformis, 291
Hemilecanium, 174, 181
Hemiptera, 3
INDEX
487
Hemipter, Suborders of, 4
Hemisphaericum, 181
Hemmicoccidinae, 193
Hempeli, 74
Hendaspidiotus, 391
Hendaspidiotus, Species of, 439
Herculeana, 441
Herbae, 330
Herrerae, 178
Hesperidum, 178
Hesperius, fernaldi, 410
Heterococcus, 130, 144
Heterophyllae, 347
Heteroptera, 4
Hibisci, angraeci, 216
Hibisci, Mytilaspis, 272
Hiemalis, 93
Hikosani, 361
Hirsuta, 378
Hispidus, 175
Homoptera, 4
Honey dew glands, 27
Horny, 165
Houardia, 174, 181
Howardia, 312
Howardia, Species of, 3 62
Howardi, Duplachionaspis, 336
Howardi, Eriococcus, 145
Howardi, Quadraspidiotus, 410
Hubbard, 81
Humilis, 351
Hunteri, Diaspidiotus, 412
Hunteri, Pulvinaria, 176
Hymenocleae, 133
Hymenantherae, 293
Hypodermis, 2 5
Hypopharynx, 159
Hypopygial setae, 162
Icerya, 69, 73
Icerya, Species of, 73
Iceryoides, 133
Ichesii, 29 5
Idiococcidae, 206
Illigeri, 74
Imbricans, 181
Imbricatum, Neolecanium, 178
Imbricata, Protodiaspis, 365
Immaculata, 424
Immanis, 98
Implicata, 406
Inchoaspis, 310
Inchoaspis, Species of, 359
Incisaspis, 311
Incisaspis, Species of, 361
Incised notches, 226
Incision, 226, 228
Incisions thickened edges, 227
Incisor, 292
Incisurae, 226
Incisus, 178
Indentata, 296
Indiae-orientalis, 268
Indica, Suturaspis, 267
Indicus, Coccus, 103
Indicus, Margarodes, 93
Inflata, 182
Inflation, 17
Inflatipes, 211
Infunda, 159
Ingae, 154
Inglisia, 171, 178
Inner angle, 161
Inner margin, 161
Innumerabilis, 176
Insignicola, 182
Insignis, 112
Interantennal setae, 124, 158
Intercalary plates, 110
Interlobular incisions, 226
Intermedia, Poliaspis, 355
Intermedia, Scrupulaspis, 287
Introduction, 1
Inusitata, 450
Irishii, Erium, 140
Irishi, Pseudococcus, 136
Irregularis, 177
Irreptus, 453
Ischnafiorinia, 372
Ischnafiorinia, Species of, 378
Ischnaspis, 271, 275
Ischnaspis, Species of, 291
Ixorae, 2 93
Jaboticabas, 211
Jacobsoni, 73
Jaliscensis, 177
Janeirensis, 423
488
THE COCCIDAE
Japonica, fioriniae, 374
Japonica, Leucaspis, 266
Japonica, Takahashia, 177
Javanensis, Aonidia, 465
Javanensis, Aulacaspis, 317
Javanensis, Opuntiaspis, 295
Javanica, Pinnaspis, 291
Jordani, 407
Juglandis, Aspidiella, 405
Juglandis, Furcaspis, 409
Juglandis, Mytilaspis, 272
Juglans-regiae, 409
Junctilobius, Aspidiotus, 386
Junctiloba, Neomorgania, 458
Juniperi, Carulaspis, 313
Juniperi, Fiorinia, 373
Juniperi, Lepidosaphes, 282
Juniperi, Pseudococcus, 136
Juniperi, Targionia, 448
Kafkae, juglans-regiae, 409
Kamerunensis, 277
Kamerunicus, 452
Kelloggi, Chrysomphalus, 418
Kelloggi, Ripersiella, 141
Kelloggi, Suturaspis, 268
Kelly i, 427
Kemptoni, 145
Kennedyae, 386
Kermanensis, 54, 57, 61, 267
Kermesiinae, Subfamily, 191
Kermes, 192, 195
Kermes, Species of, 195
Kermicus, 132, 147
Kewensis, 379
Kibariae, 178
Kiggelariae, 356
Kingii, Kermes, 187
Kingii, Ripersia, 141
Kinshinensis, 347
Koebelei, Ceroputo, 144
Koebelei, Chrysomphalus, 415
Koebelei, Icerya, 73
Krauhniae, 134
Kuwanae, 142
Kuwanaspis, 311
Kuwanaspis, Species of, 261
Kuwania, 78
Kuwaniinae, 53, 56, 59
Kuwaniinae, Genera of, 77
Kuwaniinae, Subfamily, 76
Kuwanina, 132, 147
Labelling, 22
Labiae, 12 3
Labial setae, 31
Labiatarum, 398
Labrum, 29
Lac, 150
Lacca, 154
Laciniae, 457
Lac Insects, 148
Lachnodiella, 131, 145
Lachnodius, 131, 145
Lacquer, 150
Lactea, 290
Lagosinia, 171, 178
Lahillei, Ceroplastina, 177
Lahillei, Dinaspis, 295
Lamaceratubae, 224
Lamellae, 230
Lamelles, 227
Lampas, 211
Laniger, Sclopetaspis, 324
Lanigerus, Mallococcus, 175
Larreae, Eriococcus, 145
Larreae, Protargionia, 324
Larreae, Pseudodiaspis, 364
Larreae, Tachardiella, 154
Larval pellicle, 239
Lasianthi, 281
Lasii, 141
Lassiorum, Ceroputo, 144
Lasiorum, Orthezia, 113
Last segment, 222
Latadentes, 226, 384
Latadiscaloca, 65
Latagenacerores, 237
Lataniae, 412
Latapectinae, 230
Latastei, 405
Lateral callis, 235
Lateral cerari, 121
Lateral keels, 110
Lateral lobes, 227
Lateral margin, 226
Lateral notch, 228
Lateral orbacerores, 125
INDEX
489
Lateral plates, ilO
Lateral pllacerores, 110
Lateral spinae, 160
Lateres, 226
Latilobis, ancylus, 411
Lattaspidiotus, 394
Lattaspidiotus, Species of, 457
Lattice-shaped thickenings, 234,
386
Lauretorum, 402
Lauri, 464
Laurinus, 432
Leachii, 74
Lecaniinae, 54, 57, 61
Lecaniinae, Genera of, 167
Lecaniinae, Subfamily, 155
Lecaniodiaspis, 187, 189
Lecanium, 173, 178
Lecanochiton, 172, 178
Lecanodiaspidites, 184
Lecanopsis, 169, 176
Ledi, 133
Lefroya, 128, 140
Leg formula, 34
Legs, 34
Lenticularis, 449
Leonardaspis, 274
Leonardaspis, Species of, 287
Leonardi, 226, 247, 258, 368
Leonardianna, 323
Leonardianna, Species of, 450
Lepidaspidis, 275
Lepidaspidis, Species of, 292
Leperii, 318
Lepidosaphes, 274
Lepidosaphes, Species of, 280
Lepiodosaphini, 244
Lepiodosaphini, Genera of, 273
Lepidosaphini, Tribe, 269
Leptospermi, Amorphococcus, 190
Leptospermi, Sphaerococcus, 147
Leptopsermi, Triaspidis, 277
Leucadendri, 336
Leucaenae, 178
Leucaspidini, 243
Leucaspidini, Genera of, 263
Leucaspidini, Tribe, 258
Leucaspis, 253, 260, 263
Leucaspis, Genera of, 263
Lichenoidea, 74
Lichtensia, 167, 168, 175
Leucosoma, 141
Lichtensioides, 140
Lichtensteinii, 176
Lidgetti, Anamefiorinia, 378
Lidgetti, Triaspidis, 280
Ligulaspis, 388
Ligulaspis, Species of, 423
Liliacearum, 133
Lilacina, 442
Limuloides, 366
Linaceratubae, 224
Lindingaspis, 388
Lindingaspis, Species of, 422
Lindinger, 262
Lindingeria, 248
Lindingeria, Species of, 254
Linearis, 421
Lineaspis, 3 08
Lineaspis, Species of, 339
Lineolatae, 176
Lintneri, 328
Liquidambaris, 439
Littoralis, 73
Litzeae, 3 45
Llaveia, 69, 71
Llaveia, Species of, 72
Lobata, Obluctaspis, 361
Lobata, Schizaspidiotus, 456
Lobelets, 22 9
Lobes, 227
Lobulatus, 272
Lobules, 227
Loewi, 267
Longa, 462
Longiloba, 32 9
Longirostris, 291
Longisetosa, Icerya, 73
Longisetosus, Pseudococcus, 136
Longispina, Morganella, 426
Longispina, Parlatoreopsis, 355
Longispinus, Pseudococcus, 137
Longissima, 422
Longivalvata, 176
Longula, Scrupulaspis, 287
Longulum, Lecanium, 179
Lophococcus, 70, 74
Loranthi, Aonidia, 464
490
THE COCCIDAE
Loranthi, Aulacaspis, 317
Loranthi, Macrocepicoccus, 142
Lounsburyi, Cryptinglisia, 178
Lounsburyi, Entaspidiotus, 455
Lounsburyi, Phenacaspis, 349
Lower lateral group, 236
Lubberly Cocclds, 205
Lucumae, 435
Luntii, hartil, 404
Lutea, Licbtensia, 175
Lutea, Pbenacaspis, 347
Luzulae, 175
Luzulaspis, 168, 175
Lycii, Ambigaspls, 457
Lycii, Licbtensia, 175
Lycii, Pbenacoccus, 143
Lycii, Tacbardiella, 154
Macgregori, 283
Macbili, 288
Maclurae, 176
Macrocarpae, 86
Macrocepicoccus, 130, 142
Macrocerococcus, 130, 142
Macropores, 224
Macroprocta, 377
Maculata, 432
Madagascariensis, Gascardia, 154
Madagascarensis, Tylococcus, 140
Madiunensis, 324
Madreporiform bodies, 186
Maeandrius, 430
Maenariensis, 111
Magna, 141
Magnoliarum, 181
Maideni, 295
Magnani, 154
Magnolicida, 133
Magnospinus, 452
Magnus, Entaspidiotus, 454
Magnus, ostreaeformis, 410
Major, 337
Malleola, 454
Males, 8
Mallococcus, 168, 175
Malloti, 323
Malvacearum, 178
Mammeae, 176
Mamillariae, 133
Mammillaris, 437
Mandibles, 31
Mangiferae, Aulacaspis, 317
Mangiferae, Cbrysompbalus, 415
Mangiferae, Genaparlatoria, 255
Mangiferae, Lecanium, 179
Manihotis, 189
Manni, 349
Manzanillense, 178
Manzanitae, 322
Marchalaspis, 312
Marchalaspis, Species of, 363
Marcbali, 341
Margaritae, 336
Margarodes, 92, 93
Margarodinae, 52, 55, 59
Margarodinae, Genera of, 92
Margarodinae, Subfamily, 88
Margin, 22 6
Marginal gland openings, 223
Marginal lunar pores, 223
Marginal plates, 110
Marginal pores, 223
Marginal scales, 160
Marginal setae, 157, 160, 194
Marianum, 181
Maritima, Ripersiella, 141
Maritimus, Pseudococcus, 137
Marlattaspis, 3 87
Marlattaspis, Species of, 406
Marlatti, Pboenicoccus, 211
Marlatti, Targaspidiotus, 447
Marsupial Coccids, 94
Marsupium, 95, 106
Maskell, 41
Maskellana, 276
Maskellana, Species of, 296
Maskellia, 3 92
Maskellia, Species of, 446
Maskelli, Anamefiorinia, 377
Maskelli, Drosicba, 74
Matsucoccus, 77, 78
Matsumurae, 78
Mauritianus, 426
Maxillae, 31
Maximus, 74
Mealy Bugs, 118
Media, 357
Median group, 236
INDEX
491
Median incisura, 226
Median lamellae, 109
Median lobe, 227
Median notch, 228, 234
Median pair lobes, 226, 227
Median spina, 160
Mediterraneus, Cryptaspidiotus,
426
Mediterraneous, Margarodes, 93
Megaloba, 345
Megasaissetia, 175, 182
Melaleucae, Lecanium, 179
Melaleucae, Lepidosaphes, 280
Melratuban Coccids, 80
Memaleucae, 280
Mendozinus, 133
Meridionalis, Triaspidis, 278
Mesal callis, 235
Mesal cerari, 122
Mesal group, 236
Mesal lobes, 227
Mesal margin, 161
Mesal notch, 228
Mesal orbacerores, 125
Mesal plates, 110
Mesal setae, 162
Mesocoria, 1, 33
Mesodiscaloca, 65
Mesofurcae, 34
Mesogenacerores, 236, 237
Mesolecanium, 173, 180
Mesospiracles, 36
Mesothoracic spiracles, 152, 221
Mesuae, Amorphococcus, 190
Messuae, Greenella, 450
Metacoria, 33
Metafurcae, 1, 34
Metamorphosis, 7, 9
Metaspiracles, 36
Metatentoria, 159
Metathoracic spiracles, 152, 221
Metrosideri, 386
Mexicana, sabalis, 423
Mexicana, Schizochlamidia, 178
Mexicana, Tachardiella, 154
Mexicana, Triaspidis, 278
Mexicanorum, 72
Mexicanus, Palaeococcus, 72
Meyeri, 404
Micrococcus, 131, 145
Micropores, 224, 232
Micropori, 325
Migration period, 8
Miliaris, 189
Mimosae, Aonidiella, 443
Mimosae, Lichtensia, 175
Mimosarum, Triaspidis, 279
Mimosicerya, 70, 74
Miniatae, 424
Minima, Diaspis, 304
Minima, Hemichionaspis, 343
Minima, Lepidosaphes, 282
Minima, Ripersia, 141
Minima, Trullifiorinia, 376
Minimum, Lecanium, 179
Minimus, Gonaspidiotus, 431
Minimus, Phenacoccus, 142
Minor, fioriniae, 374
Minor, Hemiberlesia, 343
Minor, Icerya, 73
Minuta, Aonidiella, 444
Minuta, Tenuiaspis, 339
Mirabilis, Kermes, 196
Mirabilis, Lophococcus, 74
Mirabilis, Toumeyella, 181
Mirabilis, tricornis, 74
Miranda, 313
Mischocarpi, 346
Missionum, 133
Mitchelli, 435
Mitulaspis, 310
Mitulaspis, Species of, 358
Monaonidiella, 392
Monaonidiella, Species of, 445
Monophlebinae, 52, 55, 58
Monophlebinae, Genera of, 69
Monophlebinae, Subfamily, 62
Monophlebus, 69, 71
Monophlebulus, 71, 75
Monticola, 112
Monserati, 304
Montserratensis, 73
Montanus, 319
Moorei, 458
Moreirai, 400
Morganella, 389
Morganella, Species of, 426
Mori, 413
492
THE COCCIDAE
Morrilli, 72
Mounting, 21
Mulgae, 272
Multilobia, 314
Multipora, Lepidosasphes, 2 85
Multipora odinae, 37 3
Multipora, Psudodiaspis, 364
Multispinosus, Ericoccus, 145
Multispinosus, Stictococcus, 75
Mussaendae, 344
Mycetaspis, 392
Mycetaspis, Species of, 442
Mycetococcus, 188, 189
Myopori, 3 87
Myristicae, 318
Myrmecophila, 141
Myrthi, Chionaspis, 304
Myrthi, Lepidosaphes, 272
Myrtus, Syngenaspis, 249
Mytilaspiformis, Duplachionas-
pis, 333
Mytilaspiformis, Syngenaspis, 251
Mytilella, 271, 275
Mytilella, Species of, 291
Myxilecanium, 172, 178
Nanus, 179
Natalaspis, 309
Natalaspis, Species of, 354
Natalensia, 141
Natalensis, Chionaspis, 304
Natalensis, Phenacaspis, 348
Nativus, 141, 142
Neglectus, 145
Neo-caledonia, 375
Neogreenia, 474
Neolecanium, 172, 178
Neoleonardia, 392, 446
Neomargarodes, 92
Neomexicanus, 133
Neomorgania, 394
Neomorgania, Species of, 458
Neosignoretia, 389
Neosignoretia, Species of, 424
Nephelii, 378
Nerii, 32 5
Newstead, 43
Newsteadia, 111, 114
Newsteadiella, 71, 75
Newsteadii, Coccus, 104
Newsteadii, Conchaspis, 215
Newsteadi, Lepidosaphes, 2 85
Nidularia, 132, 147
Niger, Duplaspidiotus, 453
Niger, Margarodes, 93
Nigerensis, 336
Nigra, Lepidosaphes, 281
Nigra, Tachardiella, 154
Nigra, Targionia, 448
Nigra, Trullifiornia, 376
Nigrocincta, 112
Nigrofasciata, 179
Nigroupunctatus, Chrysomphalus,
421
Nigroupunctatus, Kermes, 198
Nipa, 135
Nipple, 223
Nipponorthezia, 111, 114
Nitens, Poliaspis, 357
Nitens, Pseudokermes, 178
Nitida, 304
Nitrariae, 437
Nivalis, 195
Nivea, Albataspis, 290
Nivea, Monaonidiella, 446
Niveaspis, 27 6
Niveaspis, Species of, 296
Noacki, Platinglisia, 178
Noacki, Pseudoparlatoria, 314
Nopal plant, 100
Nortoni, 147
Notched, 228
Notched plates, 229
Notches, 226
Nucum, 357
Nuda, 111
Nudachaspis, 312
Nudachaspis, Species of, 365
Nudata, Augulaspis, 353
Nudatus, Palaeococcus, 72
Number of cerores, 238
Number pectinae, 230
Number plates, 230
Nymphal exuvia, 229
Nymphal pellicle, 2 39
Nymphs, 7
Nyssae, 351
INDEX
493
Oahuensis, 145
Obanal setae, 125
Oblongus, ostreaeformis, 410
Obluctaspis, 311
Obluctaspis, Species of, 361
Obscura, Aclerda, 182
Obscura, Aonidia, 464
Obscurus, Chrysomphalus, 420
Obscuratus, 147
Obtusaspis, 393
Obtusaspis, Species of, 450
Obtusum, 180
Occidentalis, Arctorthezia, 114
Occidentalis, Kermes, 197
Occidentalis, Pulvinaria, 176
Occulta, 427
Oceanicus, 430
Ocellala, 29
Ocellana, 29, 221
Ocellara, 2 9
Ocellata, 286
Ocelli, 30
Octacerores, 27, 184
Octaspidiotus, 387
Octaspidiotus, Species of, 395
Odinae, 373
Odonaspis, 388
Odonaspis, Species of, 423
Odontoglossi, 407
Ohioensis, 412
Okadae, 73
Olea, Diaspis, 304
Olea, Syngenaspis, 248
Oleae, Pillipia, 176
Oleae, pinnaeformis, 284
Oleae, Saissetia, 180
Olivacea, Orthezia, 113
Olivaceus, Pseudococcus, 133
Olliffia, 188, 190
Olliffiella, 188, 190
Olivina, 175
Omnagenacerores, 237
Opercularia, 160
Opercula, 166
Operculum, 161
Opisthoscelis, 201, 204
Opuntiaspis, 271, 276
Opuntiaspis, Species of, 295
Oraceratubae, 223
Oraceroris, 27
Orbacerores, 125, 161
Oreodoxae, 45 8
Organo retrattile anali, 161
Orientalis, 408
Orlando, 344
Ornata, Pulvinaria, 177
Ornatum, Stictolecanium, 180
Orthezia, 110, 111
Orthezia, Species of, 111
Ortheziinae, 53, 56, 60
Ortheziinae, Genera of, 110
Ortheziinae, Subfamily, 105
Orthezioides, 144
Ortheziola, 111, 114
Ortholobis, 327
Osborni, Diaspidiotus, 413
Osborni, Phenacoccus, 143
Osmanthi, 387
Ostreata, 304
Ostreaeformis, 410
Ourococcus, 210, 211
Outer angle, 161
Outer margin, 161
Oval pores, 223
Ovatus, 145
Oxycoccus, 404
Oyster-shell Scales, 269
Paenoniae, 454
Palaelecanium, 180
Palaeococcus, 69, 72
Palette, 227
Pallae, 227
Pallens, 2 89
Pallida, gloverii, 283
Pallidus, 71
Palmae, Asterolecanium, 189
Palmae, Furcaspis, 407
Palmeri, Eriococcus, 145
Palmeri, Icerya, 73
Paludicoccus, 132, 147
Pandani, 387
Pangoenensis, 429
Papillosus, 93
Paraceroris, 27
Parachionaspis, 309
Parachionaspis, Species of, 354
Paradensae, 235
494
THE COCCIDAE
Paradoxa, 460
Parafairmairia, 172, 178
Parafiorinia, 372
Parafiorinia, Species of, 378
Paragreenia, 78, 474
Paralecanium, 174, 181
Paranal plates, 109
Paranewsteadia, 391
Paranewsteadia, Species of, 432
Paraonidia, 394
Paraonidia, Species of, 454
Paraonidiella, 392
Paraonidiella, Species of, 442
Paraphyses, 227, 232
Parasitica, 3
Parastigmatic glands, 159
Parastigmatic pores, 159
Parigenitals, 236
Parischnaspis, 276
Parischnaspis, Species of, 295
Parcispinosus, 145
Parlatoroides, Pseudoparlatoria,
314
Parlatoreoides, Syngenaspis, 252
Parlatoreopsis, 309
Parlatoreopsis, Species of, 354
Parlatoria, 2 48
Parlatoriae, 2 49
Parlatoria, Species of, 252
Parlatoriini, 243
Parlatoriini, Genera of, 248
Parlatoriini, Tribe, 245
Parrotti, Antonia, 146
Parrotti, diffinis, 43 8
Parrottia, 394
Parrottia, Species of, 458
Partargionia, 394
Partargionia, Species of, 458
Parthenolecanium, 180
Parva, Diaspis, 323
Parva, Tachardiella, 154
Parvicornis, 181
Parvula, Lictensia, 175
Parvula, Protodiaspis, 365
Parvus, Kuwanina, 147
Patavina, 408
Paulistus, 416
Pectinae, 224, 229
Pectinae, formula, 230
Pectinata, 408
Pedronis, 419
Pegtop Coccids, 199
Pe-la, 181
Peli semplici, 232
Pellicles, 239
Pelomphala, 392
Pelomphala, Species of, 441
Penicillata, 450
Pentagona, 315
Penzigi, 318
Percerorsus, 133
Perconvexum, 178
Perforatus, 181
Pergande, 81
Pergandei, Porococcus, 142
Pergandei, Syngenaspis, 250
Perissopneumon, 71, 75
Peritreme, 37, 124
Perlonga, 279
Perlusus, 452
Permutans, 359
Perniciosa, 43 8
Perplexa, Aonidia, 462
Perplexus, Eriococcus, 145
Perpusilla, 256
Perrisii, Margarodes, 93
Perrisii, Trionymus, 138, 140^
Perryi, 196
Perseae, 420
Persearum, 403
Persimilis, 316
Personata, 442
Pertinax, 74
Petinii, 229
Petrophilae, 250
Pettiti, Kermes, 198
Pettiti, Phenacoccus, 143
Phantosura, 373
Phaulaspis, 395
Phaulaspis, Species of, 465
Phaulomytilus, 276
Phaulomytilus, Species of, 296"
Phenacaspis, 308
Phenacaspis, Species of, 344
Phenacoleachia, 117
Phenacoleachiinae, 53, 56, 60-
Phenacoleachiinae, Subfamily,.
115
INDEX
495
Phenacoccus, 130, 142
Phenacoccus, Species of, 142
Phenax, 442
PhilepheSra, 170, 176
Philococcus, 295
Phoenicococcus, 211
Phoradendri, Diaspis, 320
Phoradendri, Mesolecanium, 180
Phoradendri, Pseudococcus, 133
Phormii, 387
Phyllanthi, Greenoidea, 446
Phyllanthi, pergandei, 250
Phymatodidid, 289
Physokermes, 175, 182
Physopoda, 3
Picea, Aonidia, 463
Piceae, Physokermes, 182
Piceus, Diaspidiotus, 413
Pilacerores, 27, 106
Pili simplices, 232
Pilosa, Coelostomidea, 86
Pilosa, Icerya, 73
Pilosellus, 144
Pimentae, 450
Pinifoliae, 326
Pini, Leucaspis, 265
Pini, Poliaspis, 355
Pinnaeformis, 284
Pinnaspis, 275
Pinnaspis, Species of, 290
Pinnuliferus, 416
Piperis, 284
Piricola, 318
Pirogallis, 147
Pisai, 400
Pistaciae, 268
Pittospori, 253
Planchonii, 304
Planchonioides, 462
Plaque de filieres, 236
Platani, Phenacaspis, 345
Platani, Stomacoccus, 79
Plate, formula, 230
Plates, 224, 230
Platinglisia, 172, 178
Platysaissetia, 174, 181
Plebeium, 178
Plucheae, Palaeococcus, 72
Plucheae, Pulvinaria, 177
Pluridentatus, fissidens, 428
Poleii, 74
Poliaspis, 309
Poliaspis, Species of, 355
Poliaspoides, 309
Poliaspoides, Species of, 357
Pollini, 189
Pollinia, 188, 189
Polonicus, 93
Polls, 232
Pols, 229
Polygon!, 326
Popularum, 414
Porococcus, 129, 142
Porogymnaspis, 248
Porogymnaspis, Species of, 254
Porterae, 141
Portoricensis, 427
Postanal plates, 110
Position anus, 233
Position vulva, 233
Posterior lateral group, 236
Posterior lateral plates, 110
Posterior laterals, 236
Postgenacerores, 236
Postparadensa, 236
Pothi, 445
Praelonga, Cooleyaspis, 353
Praelonga, Orthezia, 112
Preabdomen, 35, 222, 300
Preanal plates, 109
Pregenacerores, 236
Preparadensa, 236
Preparation, 11
Primary lobes, 227
Primitiva, 72
Privignus, 399
Proboscidariae, 374
Proceraspis, 312
Proceraspis, Species of, 363
Processes, 232
Processi chitenosi, 232
Procoria, 1
Prontaspis, 311
Prontaspis, Species of, 359
Propinqua, rubra, 378
Proposimus, 415
Prosopidis, Lecaniodiaspis, 189
Prosopidis, Pseudococcus, 138
496
THE COCCIDAE
Prosopidis, Xerophilaspis, 465
Protargionia, 306
Protargionia, Species of, 324
Proteae, Separaspis, 427
Proteus, Syngenaspis, 251
Protodiaspis, 312
Protodiaspis, Species of, 364
Protopulvinaria, 170, 176
Provincialis, 436
Pruinosa, 189
Pruni, juglans-regiae, 409
Pruni, Pulvinaria, 177
Pseudaonidia, 394
Pseudaonidia, Species of, 453
Pseudaonidiella, 394
Pseudaonidiella, Species of, 454
Pseudaspidiotus, 255
Pseudaspidistrae, 341
Pseudaulacaspis, 305
Pseudaulacaspis, Species of, 314
Pseudinglisia, 213
Pseudischnaspis, 388
Pseudischnaspis, Species of, 421
Pseudococcus, 127, 133
Pseudococcus, Species of, 134
Pseudodiaspis, 312
Pseudodiaspis, Species of, 363
Pseudogall Coccids, 191
Pseudohesperidum, 179
Pseudokermes, 172, 178
Pseudolobes, 227, 229
Pseudonipiae, 133
Pseudoparlatoria, 305
Pseudoparlatoria, Species of, 314
Pseudophillipia, 169, 176
Pseudopulvinaria, 169, 176
Pseudoripersia, 129, 141
Pseudospinosa, 404
Pseudotargionia, 389
Pseudotargionia, Species of, 426
Psidii, 177
Psyllidae, 5
Pubescens, 195
Pugionifera, 361
Pulchella, Pulvinaria, 177
Pulchella, Pulvlnella, 176
Pulchellus, Callococcus, 147, 189
Plucher, Palaeococcus, 72
Pulchra, Aonidia, 464
Pulchra, Protodiaspis, 365
Pulvinaria, 176
Pulvinaria, Species of, 177
Pulvinata, Nidularia, 147
Pulvinella, 169, 176
Pumilus, 456
Punicae, 405
Puparium, 229, 238, 240
Purpurea, 146
Purchasi, 63, 73
Pusilla, Actenaspis, 267
Pusilla, Aonidia, 463
Pusillaspis, 2 62
Pustulans, Aspidiotus, 398
Pustulans, Asterolecanium, 189
Pustulata, 154
Putearia, 427
Puto, 130, 144
Pygidial formula, 231
Pygidial fringe, 222, 226, 303
Pygidial incision, 234, 301
Pygidial margin, 226
Pygidial setae, 231
Pygidial Type, 47
Pygidiaspis, 392
Pygidiaspis, Species of, 447
Pygidium, 35, 222, 300, 383
Pygidium deeply concave, 234
Pygidium emarginate, 234
Pygidium indented, 2 34
Pygidium notched, 234
Pyri, Cryptoparlotoria, 254
Pyri, Epidiaspis, 318
Pyriformis, Pulvinaria, 177
Pyriformis, Triaspidis, 277
Quadraspidiotus, 3 88
Quadraspidiotus, Species of, 409
Quadriclavatus, 414
Quadrifasciata, 181
Quaintancii, Pseudococcus, 133
Quaintancii, Pseudophillipia, 176
Quercicola, 189
Quercifex, 180
Quercus, 192
Quercus, Cerococcus, 189
Quercus, Ericoccus, 145
Quercus, Fundaspis, 338
Quercus, Kuwania, 78
INDEX
497
Quercus, Xylococcus, 86
Quiescent period, 8, 10
Raddoni, 74
Radiata, 189
Radicum, 142
Radicum-gramina, 141
Rap ax, 435
Rapid method, 22
Rattani, 197
Rectal spinnerets, 27
Recurvata, Mytilaspis, 272
Recurvatum, Hemilecanium, 181
Regius, 403
Regularis, 323
Remotaspidiotus, 391
Remotaspidiotus, Species of, 434
Remotaspis, 311
Remotaspis, Species of, 360
Reniformis, 417
Replicata, 439
Reticulatus, 430
Retigera, 332
Rhizaspidiotus, 390
Rhizaspidiotus, Species of, 430
Rhizococcus, 130, 145
Rhizoecus, 128, 141
Rhizophila, 450
Rhizophorae, 419
Rhododendri, 342
Rhois, 177
Rhus venix, 150
Riccae, 265
Rileyi, Icerya, 73
Rileyi, Margarodes, 93
Ringeyed Coccids, 115
Ripersia, 128, 141
Ripersiella, 128, 140
Ripersioides, 142
Riverae, 423
Rodrigueziae, 213
Rombica, 2 90
Rosae, Aulacaspis, 316
Rosae, Chionaspis, 304
Rosae, Palaeococcus, 72
Rosae, Tachardiella, 154
Roseotinctus, 133
Rossi, 421
Rostral fold, 32
Rostral furrow, 31
Rostralis, 32, 159, 221
Rostrum, 31, 159, 221
Rotundata, 154
Rubellus, 179
Ruber, Gymnococcus, 141, 142
Rubi, Tetrura, 142
Rubiginosus, 98
Rubivorum, 142
Rubra, Parafiorinia, 378
Rubra, Tachardiella, 154
Rubribullata, 387
Rubrolineata, 377
Rubrovittatus, 282
Rubsaameni, 428
Rufescens, 189
Rufa, Porogymnaspsis, 254
Rufus, Tollaspidiotus, 426
Rugaspidiotus, 393
Rugaspidiotus, Species of, 449
Rugosa, Edwallia, 178
Rugosa, Lecaniodiaspis, 189
Rumicis, 140
Rutherfordia, 306
Rutherfordia, Species of, 323
Ryani, 135
Sabalis, 423
Sacchari, 405
Sacchari-folii, 304
Salicicola, 262, 263
Salicicola, Species of, 267
Salicis, 329
Salicis-nigrae, 330
Salinus, Cryptoripersia, 141
Salinus, Ericoccus, 145
Salinus, Pseudococcus, 138
Saissetia, 173, 180
Saissetia, Species of, 180
Sallei, Capsulina, 211
Sallei, Neolecanium, 178
Salmonacea, 141
Samoana, Lindingaspis, 422
Samoana, Phenacaspis, 346
Santali, 387
Saprosmae, 376
Sassceri, Chionaspis, 329
Sassceri, Kermes, 198
Saundersii, 71
498
THE COCCIDAE
Saurefuchsin, 20
Scales, 158, 201, 213, 238, 298,
381
Scaly hairs, 229
Schizaspis, 3 94, 456
Schizaspls, Species of, 456
Schizochlamidia, 172, 178
Schizostachyi, 459
Schrottkyi, 73
Schultzei, 456
Sclopetaspis, 307
Sclopetaspis, Species of, 324
Scobinaspis, 274
Scobinaspis, Species of, 286
Scrobicularum, Hemichionaspis,
341
Scrobicularum, Trullifiorinia, 377
Scrupulaspis, 274
Scrupulaspis, Species of, 287
Scutare, 213, 215, 216
Scutiae, 354
Scutiformis, 418
Secreta, Adiscofiorinia, 379
Secreta, Odonaspis, 423
Secretus, Pseudococcus, 133
Second exuvia, 239
Second exuvium, 239
Second incisura, 227
Second lateral lobe, 228
Second lobe, 228
Second pair lobes, 228
Segmentation, 27, 35
Segregatus, 133
Selenaspidus, 385, 394
Selenaspidus, Species of, 451
Semelaspidus, 393
Semelaspidus, Species of, 451
Semiocculata, 190
Semilunar marginal pores, 223
Senegalensis, 322
Senex, 74
Sentali, 304
Separaspis, 390
Separaspis, Species of, 427
Separata, 340
Sequoiae, 136
Serrate, 22 8
Serrated ducts, 229
Serrate plates, 229
Serratulae, 73
Serratus, ancylus, 411
Serrata, Aspidiotus, 387
Serrata, Ripersia, 141
Serrei, 74
Serrifrons, 287
Serrulata, 314
Serrulate plates, 229
Setae, 26
Setiger, 415
Setosus, 133
Seurati, 438
Sexes, 7
Seychellarum, 73
Shastae, 432
Shastensis, 196
Signata, 375
Signoret, 40
Signoreti, Leucaspis, 265
Signoreti, Ortheziola, 114
Sikkimensis, 176
Silvaticus, 455
Silvestri, 145
Simillimus, Aspidiotus, 401
Simillima, Lichtensia, 175
Simillis, 376
Simplex, articulatus, 452
Simplex, hederae, 400
Simplex, Natalaspis, 354
Simplex, Phenacoccus, 142
Simplex, Poliaspoides, 357
Simplex, Pseudococcus, 133
Simulans, 177'
Silveirai, 178
Sinensis, 175
Singularis, 211
Sinistraspis, 309
Sinistraspis, Species of, 353
Sinoran Coccids, 76
Siphones, 89
Siphonodontis, 290
Situlaspis, 311
Situlaspis, Species of, 361
Sjostedti, Monophlebus, 71
Sjostedi, Stictococcus, 75
Slow method, 17
Smilacis, 445
Smithii, 139
Socialis, Antonia, 146
INDEX
499
Socialis, Conchaspis, 216
Socotrana, 449
Solani, Phenacoccus, 143
Solani, Pseudococcus, 133
Solacerores, 237
Solenopsis, 142
Solus, 414
Solidaginis, 113
Sonorensis, Orthezia, 112
Sonorensis, Toumeyella, 181
Sophorae, 387
Sorghiellus, 135
Spartinae, 325
Spathulata, 2 96
Spatulata, 463
Specialized Coccidae, 47
Specialized Diaspidinae, 241
Spermococcus, 168, 175
Sphaerococcopsis, 210, 211
Sphaerococcus, 132, 147, 206
Sphaerolecanium, 180
Sphaerioides, 420
Spinae, 159, 187
Spinaspidiotus, 390
Spinaspidiotus, Species of, 428
Spine ducts, 230
Spines, 106, 232
Spinifera, Berlesaspis, 289
Spiniferus, Cylindrococcus, 211
Spiniger, 433
Spinnerets, 159, 223, 236
Spinnerets of spiracles, 159
Spiniferus, 142
Spinomarginata, 256
Spinosissima, 461
Spinosa, Berlesaspis, 2 89
Spinosus, Aspidiotus, 398
Spinosus, Eriochitin, 175
Spinulae, 26
Spinulosa, Luzulaspis, 175
Spinulosa, Pulvinaria, 177
Spiracerores, 151, 159
Spiracles, 36
Spiracular depressions, 160
Spiracular grooves, 160
Spiracularia, 37, 64
Spiracular setae, 160
Spiracular spines, 160
Sporoboli, 141
Spuria, 145
Squamae, 229
Squameaux, 229
Squames, 229
Squamosus, 304
Stachyos, 142, 143
Staining, 19
Stantophri, 326
Stebbingii, 75
Steelii, 136, 140
Steingelia, 78
Stellifera, 177
Stick-lac, 152
Stictococcus, 71, 75
Stictolecanium, 173, 180
Stigmacoccus, 70, 74
Stigmata, 3 6
Stigmatic aperatures, 153
Stigmatic clefts, 160, 221
Stigmatic process, 151
Stigmatic spines, 160
Stilosa, 323
Stomacoccus, 78, 79
Stotzia, 168, 175
Strachani, Lagiosinia, 178
Strachani, minor, 343
Striata, Lineaspis, 340
Striata, Phaulomytilius, 296
Striata, Stotzia, 175
Stricta, 264
Stringaspidiotus, 393
Stringaspidiotus, Species of, 451
Strobilanthi, 350
Subandina, 73
Subapical setae, 163
Subcircular scars, 65
Subcorticalis, 352
Subcuticularis, 444
Subdorsal keels, 109, 110
Subdorsal plates, 109, 110
Suberi, vitis, 448
Submarginal setae, 231
Submarginal tubercles, 158
Subnudata, 354
Subrubescens, 395
Subsimilis, 440
Substriata, 362
Subterranea, Epidiaspis, 304
Subterraneus, Pseudococcus, 133
500
THE COCCIDAE
Superbus, 142
Supplementary scales, 110
Suturaspis, 262, 264
Suturaspis, Species of, 267
Sutures, 222
Sylvatica, 338
Sylvestrls, 147
Syncarpiae, 463
Syngenaspis, 246, 247
Syngenaspls, Species of, 248
Synopsis of the Diaspidinae, 241
Syriaca, 320
Tachardia, 153, 154
Tachardiella, 153, 154
Tachardiinae, 53, 56, 61, 148
Tachardiinae, Genera of, 153
Tachardiinae, Subfamily, 148
Tachardina, 154
Tafiranus, 406
Takahashia, 170, 177
Tamarinda, 75
Tangalla, 73
Tangana, 348
Taorensis, 436
Taprobana, 444
Targaspidiotus, 392
Targaspidiotus, Species of, 446
Targionia, 393
Targionia, Species of, 447
Targionidea, 393
Targionidea, Species of, 449
Targioniopsis, 460
Targioni Tozzetti, 40
Tasmaniae, 398
Taxa, 443
Taxicola, 304
Taxifoliae, 182
Tayabana, 445
Tectaria, 425
Tectococcus, 131, 145
Tectonae, 436
Tectopulvinaria, 170, 176
Tegalensis, 304
Telson, 110
Temnaspidiotus, 387
Temnaspidiotus, Species of, 403
Tenebricosa, 443
Tentorium, 159
Tenuiaspis, 308
Tenuiaspis, Species of, 339
Tenuidisculis, 360
Tenuior, 2 81
Tenuipes, 141
Tenuis, 375
Terminaliae, 179
Termitococcus, 129, 141
Terrestris, 141
Tessalata, Lecaniodiaspis, 189
Tesselatus, Eucalymnatus, 181
Tesserae, 163
Tesserata, 458
Tessarobelus, 70, 74
Test, 151, 152, 165, 185
Tetrura, 142
Texanus, Rhizococcus, 145
Texensis, Diaspis, 322
Texensis, Pseudococcus, 133
Theae, Duplaspidiotus, 454
Theae, Eriochitin, 175
Theae, Fiorinia, 375
Theae, Hemichionaspis, 342
Theae, Syngenaspis, 249
Thickened lateral margin, 226
Thickenings body-wall, 232
Third incisure, 227
Third lateral lobe 228
Third lobe, 22 8
Third pair lobes, 228
Theobromiae, 72
Thoracic cerari, 122
Thoracic spiracles, 221
Thorax, 33
Tiliae, 177
Timberlakei, 134
Tinctorius, 142
Tinerfensis, 437
Tinsleyi, Eriococcus, 145
Tinsleyi, Pulvinaria, 17,7
Tokionis, 182
Tollaspidiotus, 389
Tollaspidiotus, Species of, 425
Tomentosus, 103
Tools, 14
Tortoise Scales, 155
Toumeyella, 175, 181
Toumeyi, 316
Townsendi, Asterolecanium, 189
INDEX
501
Townsendi, Cockerellella, 75
Townsendi, Diaspis, 321
Townsendi, Hemichionaspis, 344
Townsendi, Palaeococcus, 72
Townsendi, Pseudococcus, 133
Townsendi, Quadraspidiotus, 409
Townsendiana, 293
Trabuti, Hemiberlesia, 437
Trabutina, 130, 144
Transcaspiensis, 440
Translucens, 401
Transparens, 401
Transverse pilacerores, 109
Transverse plate, 109
Transverse thickenings, 235
Travancorensis, 279
Triaspidis, 273
Triaspidis, Species of, 276
Trichomytilus, 273
Trichomytilus, Species of, 276
Tricerores, 27, 121
Trichura, 141
Tricolor, 439
Tridentata, 465
Trifolii, 139
Triglandulosa, 451
Triloba, 216
Trilobitiformis, 453
Trimeni, 93
Trinacis, 304
Trinotatus, 198
Trionymus, 127, 138
Trionymus, Species of, 139
Trivittata, 141
Troglodytes, 181
Trullae, 227
Trullifiorinia, 372
Trullifiorinia, Species of, 375
Truncaspidiotus, 390
Truncaspidiotus, Species of, 431
Tsugae, 407
Tubercularis, Aulacaspis, 317
Tuberculatum, Neolecanium, 178
Tuberculatus, Pulvinaria, 177
Tubular dermal glands, 185
Tubular glands, 185
Tubular spinnerets, 223
Tubuli, 223
Tuliparum, 133
Tumida, 374
Turgida, 182
Turgipes, 141
Tylococcus, 127, 140 ^
Uberifera, 2 53
Uhleri, 72
Uhleria, 3 69
Uhlerian, Scales, 367, 368
Ulmi, Hendaspidiotus, 440
Ulmi, Lepidosaphes, 285
Ulmicola, ficifolii, 283
Ultima, 112
Ultracoelostoma, 86, 87
Umbaspis, 306
Umbaspis, Species of, 323
Umbonatus, 177
Umboniferus, 417
Unachionaspis, 307
Unachionaspis, Species of, 337
Unacoria, 1
Unapectinae, 230
Unaspidiotus, 3 87
Unaspidiotus, Species of, 405
Unaspis, 308
Unaspis, Species of, 339
Undulatus, 432
Ungulaspis, 274
Ungulaspis, Species of, 286;
Ungulata, 286
Ungual digitules, 159
Unicolor, 2 86
Unilateralis, 353
Uniloba, 292
Unilobus, 434
Upper group, 236
Upper lateral group, 236
Urbicola, 177
Urichi, Neolecanium, 178
Urichi, Asterolecanium, 189
Urticae, 113
Uviae, 412
Uvariae, 342
Vaginal aeroles, 65
Vaginal discs, 65
Vaginal opening, 233
Valve anali, 161
Valve of operculum, 161
502
THE COCCIDAE
Varians, 397
Varicaspis, 390
Varicaspis, Species of, 431
Varicosa, 346
Variolosum, 189
Vagabundus, 402
Vejdovskyi, 114
Ventralabia, 123
Ventralis, 179
Ventral glands, 236
Ventral grouped glands, 236
Ventral pilacerores, 109
Ventral plates, 109
Ventral scale, 239
Ventral scars, 65
Ventral setae, 231
Ventral thickenings, 162, 235
Versiculaspis, 312
Versiculaspis, Species of, 364
Viburni, 175
Victoriae, intermedia, 2 87
Villosa, Ripersia, 141
Villosa, Trionymus, 139
Vinsonia, 171, 177
Virescens, 396
Virgatus, 133
Viridis, Carpochloroides, 211
Viridis, Greeniella, 462
Viridis, Lecanium, 179
Viridis, Syngenaspis, 251
Viridis, theae, 249
Viridula, Ripersia, 141
Viridulum, Asterolecanium, 189
Visci, Diaspis, 304
Visci, Phenacaspis, 350
Vitiensis, Aspidiotus, 399
Vitiensis, Diaspis, 304
Vitis, Chionaspis, 326
Vitis, Pulvinaria, 177
Vitis, Targionia, 447
Vitium, Margarodes, 93
Vuilleti, Hendaspidiotus, 440
Vuilleti, Lophococcus, 74
Vuilleti, Marchalaspis, 363
Vulva, 36, 162, 233
Wairoensis, 86
Waldeni, 197
Walkeriana, 70, 74
Washing, 18
Wax, 223
Wax ducts, 223, 224
Wax glands, 236
Websteriella, 247
Wedge-shaped plates, 110
Westwoodii, 98
Wheeleri, 133
White-shield Scales, 258, 260
Wilga, 287
Wilmattae, 142
Wistariae, 349
Wroughtoni, 147
Xanthopthalma, 312
Xanthopthalma, Species of, 366
Xanthorrhoeae, 336
Xerophilaspis, 395
Xerophilaspis, Species of, 465
Xylococcinae, 53, 55, 59
Xylococcinae, Genera of, 85
Xylococcinae, Subfamily, 80
Xylococcus, 85
Xylococcus, Species of, 86
Yuccae, Ceroputo, 144
Yuccae, Ponscolombia, 141
Yuccae, Lecaniodiaspis, 189
Yuccae, Neosignoretia, 424
Yuccarum, 447
Yulupae, 424
Zamiae, 358
Zapotlanum, Erium, 140
Zapotlana, Lichtensia, 175
Zealandica, Coelostomidea, 86
Zealandica, Phenacoleachia, 117
Zeteki, 73
Zeylanica, 78
Zimmermanni, 75
Zizyphi, 2 52
Zonata, 409
i
THE UNIVERSITY LIBRARY
UNIVERSITY OF CALIFORNIA, SANTA CRUZ i
SCIENCE LIBRARY
This book is due on the last DATE stamped below.
N.B.-HOLD
R£C'0 StP ^ . idbb
MAY 2 3 1979
REC'DJUN4 1979
MAR 16 '81
rec:dmar6 1981
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MAY 4 1983REC'D
OCT 23 1985
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OL523.C7M3 Sci
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