UNIVERSITY OF CALIFORNIA AT LOS ANGELES Agric. Dept. 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 believe 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 COMPA> CHAMPAIGN, ILLINOIS ' ' ' ' : Agric. Dept TO MY FORMER STUDENTS OF THE COCCIDAE WHO THROUGH THEIR INSISTENCE ARE RESPONSIBLE FOR THE COMPLETION AND PUBLICATION OF THIS VOLUME 268019 PREFACE The following materials were originally collected for the 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 and hence unpublished.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- 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, URBANA, ILLINOIS, July 15, 1920. CONTENTS PAGE CHAPTER I. INTRODUCTION 1 CHAPTER II PREPARATION 11 CHAPTER III EXTERNAL ANATOMY 24 CHAPTER IV. CLASSIFICATION 40 CHAPTER V. SUBFAMILY MONOPHLEBINAE 62 CHAPTER VI. SUBFAMILY KUWANIINAE 76 CHAPTER VII. SUBFAMILY XYLOCOCCINAE 80 CHAPTER VIII. SUBFAMILY MARGARODINAE _. -88 CHAPTER IX. SUBFAMILY CALLAPAPPINAE 94 CHAPTER X. SUBFAMILY COCCINAE 99 CHAPTER XI. SUBFAMILY OETHEZIINAE 105 CHAPTER XII. SUBFAMILY PHENACOLEACHIINAE 115 CHAPTER XIII. SUBFAMILY ERIOCOCCINAE 118 CHAPTER XIV. SUBFAMILY TACHARDIINAE 148 CHAPTER XV. SUBFAMILY LECANIINAE 155 CHAPTER XVI. SUBFAMILY ASTEROLECANIINAE _. ._ 183 PAGE CHAPTER XVII. SUBFAMILY KEBMESHNAE 191 CHAPTER XVIII. SUBFAMILY APIOMOBPHINAE 199 CHAPTER XIX. SUBFAMILY CYLINDBOCOCCINAE 205 CHAPTER XX. SUBFAMILY CONCHASPINAE 212 CHAPTER XXI. SUBFAMILY DIASPIDINAE 217 CHAPTER XXII. TBIBE PABLATOBHNI 245 CHAPTER XXIII. TBIBE LEUCASPINI 258 CHAPTER XXIV. TBIBE LEPIDOSAPHINI 269 CHAPTER XXV. TBIBE DIASPIDINI 297 CHAPTER XXVI. TBIBE FIOBINHNI 367 CHAPTER XXVII. TBIBE ASPIDIOTINI 380 BlBLIOGBAPHY 466 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 1 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 eoccids 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 eoccids 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 eoccids 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 always 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 wins; labium a conical tube enclosing one mandible and two maxillae; mazillae 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 finser 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, Homoptcra, 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 4 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. HETEBOPTEBA. 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 HOMOPTEBA. 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. GULABOSTBIA. The suborder Heteroptera as here denned includes the super- families and families usually assigned to it. The suborder Homoptera includes the superfamilies Cicadoidea, Jassoidea, Ful- goroidea, and Membraeoidea, 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 5 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 always 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 short, rarely reaching mesocoxae or as far as metathoracic spiracles if mesocoxae are wanting in fe- male, wanting in older nymphal stages and always in cdult 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 coceids 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 family, 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 fix 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 ease 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 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 coccid 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 insectean 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 COCDIDAE 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 wfib 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 coccids. 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 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 sc 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 first, 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 wateh-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 PREPAKATION 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 few 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 mark the slide in such a way that the mounted specimens can be con- nected up with the original lot of material from which they were 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 : Remove Insect from Host-plant Treat with Caustic Potash. Wash in Distilled Water. Harden in 95 per cent Alcohol. Stain in Alcoholic or Aqueous Stain. I 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 endoskeleton 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 borne 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 euticular 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 euticular 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 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 lare 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 CALLIPAPPIXAE 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 segment 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 invagination 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 Tepper 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, y THE 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 westwoodii 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 flat 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 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 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 war; 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, similar 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 coceids, 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 Linnaeus. 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 Linnaeus. 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 OBTHEZIINAE 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 flat 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 coccids. 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 Coccids, 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 oraeeroris, 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 alnd the sutures between the trochanter and femur and often also between the tibia and the tarsus of each of the legs are frequently wanting, or 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 Cockerel!, 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 tc 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 ORTHEZIINAE 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. b. Antennae of adult female with four or more segments and 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. Donglasia 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, strawberry, 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 second and 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 Ckl!. ff. 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 Ringeyed 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, Phenacoleachia, 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 ER1OCOCCINAE The Mealy Bugs The body of the adult female varies from convex to subrotund, 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. The mouth- parts, rostrum and rostralis, 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 trochanto-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 of 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. 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 118 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, sc 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 antenna] 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 wingless 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 eerores 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 t-ricerores 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 ceraran 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, fossette 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 without, 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 ring 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 end 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 ERIOCOCCIKAE 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 setae 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. i. 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 1. 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. Riper sia Sign. oo. Antennae of adult female with six segments; anal cerari only present; ceraran setae setiform. Cryptoripersia 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 cerores 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. 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. Ceroputo 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. Trabutina 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 with mealy 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. i. 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 Antonio, 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 Sphaerococcus 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. bb. 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. Pseudococcus Westw. This is the largest genus of the subfamily. Over one-half of the described 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 COCCIDAE 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, ccraran 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. krauhniae 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. d. 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. dd. 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 L,a.sius.sorghiellus Forbes. 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 longisetosna 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 few 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. ee. 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, Mauritius, 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, Flcug, 138 THE coccroAE 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 scainus 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: SUBFAMILY ERIOCOCCIXAE 139 SPECIES OF TRIONYMUS a. 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. californicus 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.