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MORPHOLOGY AND TAXONOMY (; *"”) OF ADULT MALES OF THE ““"" FAMILY COCCIDAE (HOMOPTERA : COCCOIDEA)
J. H. GILIOMEE
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ENTOMOLOGY Supplement 7 LONDON : 1967
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MORPHOLOGY AND TAXONOMY OF ADULT MALES OF THE FAMILY COCCIDAE (HOMOPTERA : COCCOIDEA)
BY
J. H. GILIOMEE y, ,
Dept. of Entomology, University of Stellenbosch) South Africa
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Supplement 7 LONDON : 1967
THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), <¢nstituted im 1949, 15 issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department.
This paper is Supplement No. 7 of the Entomolog- ical series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals.
© Trustees of the British Museum (Natural History) 1967
DRUSDEES OF THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 20th January, 1967 Price fase
MORPHOLOGY AND TAXONOMY OF ADULTE MALES OF THE FAMILY COCCIDAE (HOMOPTERA : COCCOIDEA)
By J. H. GILIOMEE*
CONTENTS Page
ACKNOWLEDGEMENTS ; : 4 INTRODUCTION : ; : , ‘ : : : ; : 4 REVIEW OF THE LITERATURE 5 MATERIAL AND TECHNIQUE 7
KEY TO LETTERING ON FIGURES - : ; ; P : z 9 ABBREVIATIONS USED IN THE FIGURES : : : . ‘ } 9 GENERAL MORPHOLOGY . ; ; . : : ‘ ; : II General Characteristics 3 : . 3 : ‘ ; : 12 The Head : - : 3 : : : : : é 14 Head Capsule : : : ‘ i ; : : : 14 Antennae . . ; ‘ F : ; : ; : 19 The Thorax , ‘ ' . F : : ; ‘ ; 21 Prothorax . é : , y : P : : ; 21 Mesothorax : : ; . : , _ : 5 2 Metathorax : 4 : d - ; : : ; 28 Wings and Halteres_ - : : : : : : 30 Legs . , : ; ‘ : ‘ ‘ : * ; 31 The Abdomen . : : ; ; " ‘ PF 3 33 Pregenital Seemients ; - ; : : : 33 Genital Segment and External Genitalia : ‘ ; ; 36 DESCRIPTION OF THE SPECIES . : ; : : : 2 : 37 The EULECANIUM Group é 3 ‘ : : ; . 37 The ERIOPELTIS Group . ‘ ~ . , ‘ . : 7hG | The INGLISIA Group F ; ; : : 5 : ; 88 The COCCUS Group . : : : : ? ; : 5 92 DISCUSSION : : : 125 Taxonomic Sieairibanidd of the € haracters : 126 Classification and Interrelationships of the Coccidae based on the Males . : - F 3 : ’ : ‘ ; 139 Groups of Genera ; ; g . : : : F 139 Genera : : : ; : : : : : : 146 Species : I51 Relationships of the isecekiae ith ie eae Subdivisions a the Coccoidea . 2 . : : . ; : E . 153 KEYS é ; ‘ , . . ; 3 : : . : 159 IREFERENCES . ; : : , : : : - : : 164 INDEX . ‘ : : : : : . : : : : 168
* The information contained in this work was submitted in the form of a thesis for the degree of Doctor of Philosophy in the University of London, June, 1964.
4 MORPHOLOGY AND TAXONOMY OF ADULT MALES
SYNOPSES
The males of 23 species (representing 19 genera) of the family Coccidae have been described and illustrated in detail and a general account of the external morphology of male Coccidae is given. A number of structures present in other male Coccoidea but not hitherto observed in the Coccidae have been recorded. The relationships of the lecanoid type of male with the margaroid and diaspidoid types have been discussed and the males of two families of the lecanoid type (Coccidae and Pseudococcidae) have been compared with each other. Within the Coccidae the males were often found to reveal different relationships from the female and a classification is suggested which differs from the classifications based on female characters. The results of this study is in accordance with recent discoveries that the characters of the male are valid at all taxonomic levels, including genera and species. Detailed keys to groups of genera, genera and species have been provided.
ACKNOWLEDGEMENTS
I WouLpD like to express my gratitude to Dr. K. Boratynski for suggesting the problem and supervising the work, Mr. R. G. Davies for advice and constructive criticism, and Prof. O. W. Richards, F.R.S. for providing facilities at the Imperial College of Science and Technology, London. I also wish to thank Dr. D. J. Williams, Commonwealth Institute of Entomology, London and Mr. G. De Lotto, Plant Pro- tection Research Institute, Pretoria, who supplied and identified some of the mater- ial, and the large number of other workers who so generously made material avail- able for this study. They are : Prof. N. S. Borchsenius, Zoological Institute, Academy of Sciences, Leningrad ; Dr. J. M. Cherret, University College of North Wales, Bangor, Wales ; Dr. M. S. K. Ghauri, Commonwealth Institute of Entom- ology, London ; Prof. M. Kosztarab, Virginia Polytechnic Institute, Blacksburg, Virginia ; Prof. Z. Kawecki, Agricultural University, Warsaw ; Dr. G. Matesova, Institute of Zoology, Academy of Sciences, Alma-Ata, Kazakhstan ; Ing. Mitié- Muzina, Institute for Plant Protection, Belgrade ; Mr. J. Munting, Plant Protect- ion Research Institute, Pretoria; Dr. J. Rehaéek, Virological Institute, Bratislava ; Dr. J. G. Theron, University of Stellenbosch, Stellenbosch.
I am also indebted to the Trustees of the British Museum (Nat. Hist.) for per- mission to study material in their collection.
Finally, I wish to thank the authorities of the Commonwealth Scholarship and Fellowship Plan for financial assistance, and the University of Stellenbosch and the Government of the Republic of South Africa for granting me leave to undertake this study.
INTRODUCTION
As far as the Coccidae are concerned it is true to say that this large and important family is still very inadequately known, especially as regards the interrelationships of its members. This is partly due to the fact that satisfactory preparations of the adult females can usually only be obtained from freshly moulted specimens, which are seldom available ; preparations of the old, frequently heavily sclerotized females show only a very limited number of characters and the identity of many species described from such specimens is uncertain. A few workers (Steinweden, 1929 ; Sulc, r941 ; Bodenheimer, 1953) indicated the close relationship of a small number
OF THE FAMILY COCCIDAE 5
of genera, but the only comprehensive classification which has so far been published, is that of Borchsenius (1957), who divided the family into 3 subfamilies, and one of the subfamilies into 2 tribes. His classification was based mainly on a small number of characters of the adult female, among which he used also the way in which the body of the female and the eggs are protected. It was considered that a study of the male would contribute substantially to our knowledge of this family and make it possible to support or modify the classification suggested by Borchsenius, just as Ghauri’s work has done with regard to the Diaspididae. Thus the scope of the present work was : (i) to make a detailed study of the morphology of a represent- ative sample of the family Coccidae ; (i1) to describe these species in detail ; (iii) to determine what characters are of taxonomic importance and on what levels they can be used, and (iv) to advance our understanding of the relationships of members within the family, and of this family with other subdivisions of the Coccoidea.
REVIEW OF THE LITERATURE
The literature pertaining to male Coccoidea has been adequately reviewed by Ghauri (1962) and one need only mention a number of papers, particularly referring to the Coccidae, that were not discussed by him or were published subsequent to his review.
An early publication, not mentioned by Ghauri, is Newstead’s (1g01, 1903) monograph on the British Coccoidea. Newstead described the males whenever they were available, but mentioned only the most obvious features. He pointed to the fact that the number and position of the eyes might be of generic importance. Another early paper is that of Moulton (1907), who studied the monterey pine scale, Physokermes insignicola (Craw). Following the pattern established by Putnam (1879), he gave equal attention to all stages and included some information on their internal anatomy. The morphology of the adult male was only briefly outlined, but special attention was given to the eyes and terminal antennal segments.
Silvestri (I9g1ga, Ig1gb, 1920) published three papers dealing with Sphaerolecanium prunastri (Fonsc.), Eulecanium coryli (L.) and Ceroplastes sinensis D. Guerc. respectively. His descriptions and illustrations of the males lack detail, but such useful information as the number of eyes, and the structure of the genital and pre- genital segments is given. These papers are included as an appendix to Leonardi’s (1920) monograph on the Coccoidea of Italy. Leonardi briefly described the males of a wide variety of Coccoidea of which the following are only a few examples (names as given by Leonardi): Aspidiotus hederae (Vall.), Lichtensia viburni Signoret, Aclerda berlesit Buffa, Pseudococcus citri (Risso), Eriococcus auricariae Mask., Micrococcus silvestrit Leon., Trabutina leonardi Silv. and Ceroputo superbus Leon. In all these descriptions only the more obvious features were mentioned, but information such as the length of the body, antennae, hind legs and wings is included.
In a paper by Smutterer (1954), the males of Eulecaniuwm corni and E. crudum were illustrated, but not described. Kawecki’s (1958b) paper on Eulecanium coryli (L.) also contains a brief reference to the male ; he suggested that the term “ pseudo- halteres ’”’ should be used for the reduced hind wings.
6 MORPHOLOGY AND TAXONOMY OF ADULT MALES
A number of Russian workers have given attention to male Coccidae. Hadzibejli (1955), when describing a new species (Neopulvinaria imeretina), gave an account of the male. Borchsenius (1957), in a monograph on the Coccidae of the USSR, included original descriptions and illustrations of the males of 11 species, short notes on others, and repeated some of the published descriptions ; he also (1960) briefly described the male from each of the families Kermococcidae, Asterolecaniidae and Aclerdidae. Bustshik & Saakjan-Baranova (1962) dealt with some aspects of the life history, morphology and internal anatomy of the male of Coccus hesperidum. The descriptions of these workers follow the same general pattern, i.e. the morphology is described in general terms only, but some attention is given to the details of the head, antennae, scutellum, as well as the genital and pregenital segments. The main contribution of these papers collectively, and that of Borchsenius in particular, is that they clearly show the availability of characters in the male which can be used for taxonomic purposes.
Apart from Bustshik and Saakjan-Baranova, Bielenin (1962, 1963, 1963a) also studied the internal anatomy of a male soft scale, Parthenolecanium pomeranicum (Kaw.).
As far as families other than the Coccidae are concerned, mention can be made here of a paper on the Aclerdidae by McConnel (1954), in which the male of the family is briefly described, with the statement that “ considerable diversity of form existed among the few species available ”’
As pointed out by Ghauri (1962), a new standard of detail and accuracy was reached by Theron (1958) and the latter’s study should form the foundation on which any study of male Coccidae is based. Using Theron’s interpretations and termin- ology, Giliomee (1961) gave a detailed account of the morphology of 3 species of the genus Pseudococcus (Pseudococcidae) ; he also studied the chaetotaxy and discussed a number of characters which can be used to separate the 3 species. One of the species (P. maritimus Ehrhorn) is described as consisting of two “‘ types ’’ which show small differences, rather smaller than those observed between the other two species of Pseudococcus or the interspecific differences recorded by Beardsley (1960). However, in view of Wilkey & McKenzie’s (1961) finding (from a study of the females) that more than one distinct, but very similar species have been involved under the name P. maritimus, it now seems likely that the two “types ’”’ of male described were in fact two species.
Ghauri (1962), in an excellent paper on the males of the Diaspididae, critically examined and amended Theron’s (1958) definition of the male of this family ; he studied 24 species (representing 4 tribes and 16 genera) and proved convincingly that male characters could be used at all taxonomic levels.
A few papers on male Coccoidea have appeared since the publication of Ghauri’s work. Beardsley (1962) published a paper in which he described the males of another 5 species of the Pseudococcidae, including the interesting species Puto yuccae (Coquillet) and Rhizoecus falcifer Kunckel d’Herculais. Husseiny & Madsen (1962) dealt with the life history of Lecaniwm kunoensis Kuw. and included a des- cription of the adult male. This description is very inadequate and shows that the
OF THE FAMILY COCCIDAE 7
authors were not aware of Theron’s paper. Theron (1962), in a paper on the same lines as his earlier publication, gave an account of the structure and relationships of the male of Phenacoleachia zealandica (Mask.) and stated that it “ ostensibly belongs to the margaroid group’, showing certain similarities to the male of Steimgelia.
MATERIAL AND TECHNIQUE
The males of 23 species, belonging to Ig genera were studied. It was found that the classification of Borchsenius (1957), based upon female characters, was not corroborated by the characters of the male, which revealed different relationships within the family. This will be discussed later and in the list below the species studied are arranged into the suggested four new groups.
The EULECANIUM Group
Eulecanium Cockerell, 1896.
E. tiliae (Linnaeus, 1758). Nemolecanium Borchsenius, 1955.
N. abietis Borchsenius, 1955. Physokermes Targioni Tozetti, 1868.
P. piceae (Schrank, 1801). Rhodococcus Borchsenius, 1953.
R. spiraeae (Borchsenius, 1949). Palaeolecanium Sulc, 1908.
P. bituberculatum (Targioni Tozetti,
1868).
Phyllostroma Sulc, 1942.
P. myrtilli (Kaltenbach, 1874). Filippia Targioni Tozetti, 1868. F. viburm (Signoret, 1873). Ctenochiton Maskell, 1879.
C. species.
Ericerus Signoret, 1874.
E. pela (Chavannes, 1848). Genus A species. Sphaerolecanium Sulc, 1908.
S. prunastri (Fonscolombe, 1834).
The ERIOPELTIS Group
Eniopeltis Signoret, 1871. E. species. E. ?festucae (Fonscolombe, 1834).
Luzulaspis Cockerell, 1902. L. luzulae (Dufour, 1864).
The INGLISIA Group
Inglisia Maskell, 1879. I. theobromae Newstead, 1917.
The COCCUS Group
Coccus Linnaeus, 1758.
C. hesperidum Linnaeus, 1758. Genus B species (nr. Pulvinaria). Pulvinaria Targioni Tozetti, 1868.
P. 2betulae (Linnaeus, 1758).
P. acericola (Walsh & Riley, 1868).
Parthenolecanium Sulc, 1908.
P. cornt (Bouché, 1844).
P. pomeranicum (Kawecki, 1954). Ceroplastes Gray, 1830.
C. berliniae (Hall, 1931).
C. species.
8 MORPHOLOGY AND TAXONOMY OF ADULT MALES
The two species, Genus A sp. and Genus B sp., are apparently both new species and genera, but no definite statement can be made here as the females are still being studied. In the JNGLISIA group only one species (I. theobromae) was studied in detail, but 3 specimens of another (Ceroplastodes chiton Green) were compared with it, and their characters used in the discussion.
Most of the species were obtained from workers all over the world, who generously made material from their collections available, or collected material specially for the purpose of this study. The specimens were usually received in 70% alcohol, but specimens of 4 species received from J. Rehdéek were remounted from Swann’s mountant. In most cases the material received was already identified ; unidentified material obtained from various sources was identified by K. Boratynski, G. De Lotto and D. J. Williams. The males of six species were collected by myself, one of them (Ctenochiton sp.) in Stellenbosch, South Africa, and the rest at the Imperial College Field Station, Sunninghill, Berks., where their habitats were known to Dr. Boratynski.
In nearly all cases, 10 specimens of each species were examined, the various measurements taken and the setae counted. For each sample the range of variation was recorded and the average calculated. The specimens were prepared for micro- scopic study according to the method described by Ghauri (1962). It was found, however, that 45 minutes in KOH was usually the shortest period needed to clear the specimens and often several hours were necessary for the larger specimens. Only one stain, Chlorazol Black E, was used and the specimens were stained for one hour.
In making the illustrations, the same techniques and procedure were followed as in Giliomee (1961).
In order to standardize the measurements and prevent repetition in the individual descriptions, the way in which the various structures were measured is explained in detail below (see also Text-fig. I).
The body length was measured from the anterior margin of the head to the apex of the penial sheath.
The head exhibits no definite structure posterodorsally and its length was there- fore measured from the anterior margin to the first ridge on the thorax, the pronotal ridge ; its width was measured across the genae. The external margin of the cornea is not very distinct and the internal diameter was therefore taken. Antenna: The length of the scape was measured along the dorsal margin, the width across the middle of the segment ; the length of the pedicel was measured from the articular process posteroventrally to the apex of the segment, and the maximum width was taken ; the width of segments III and X was measured where they are widest, but that of segments IV-IX across the middle because they are sometimes rather wider at the apex.
The length of the thorax was measured from the pronotal ridge to the posterior margin of the mesopostphragma (the latter was also used by Ghauri (1962) in estimating the length of the thorax of the Diaspididae). The length of the prescutwm
OF THE FAMILY COCCIDAE 9
was measured from the anterior margin (topographically) to the prescutal suture, without including the broad internal ridge of the latter ; the width was measured across the middle of the sclerite, including the lateral mdges. The maximum length and width of the median membranous area of the scutum, the length and width across the middle of the scwtellum, and the maximum length and width of the basisternum (but not including the ridges) was taken. The length of the wing was measured from the base of the costal complex of wing veins to the tip of the wing, and the width across its maximum expansion. The length of the segments of the leg was measured as in Pseudococcus (Giliomee, 1961) ; the tarsus, however, was measured along the outer margin instead of the inner, the former being the more accurate. The width of each segment was measured at its maximum, with the ex- ception of the tibia which was measured across the middle because it is usually somewhat wider at the apex.
The length of the abdomen was measured from the mesopostphragma to the anus, and the width across segment III. The penial sheath may be somewhat curved and for accuracy its length was measured in pleural view, following the curve ; its width was measured at the level of the base of the aedeagus.
The material used in this study has been deposited in the collections of the British Museum (Natural History), London, the Imperial College of Science and Technology, London and the Department of Entomology, University of Stellenbosch, South Africa.
KEY TO LETTERING ON FIGURES
The lettering of all figures is uniform and is as follows :
A Dorsal and ventral aspects of body IL Hind claw, posterior view AI Lateral aspect of body M Part of inner margin of fore coxa, B Head, dorsal view showing setae and coxal bristle(s) (% Head, ventral view N Articulation of fore wing, showing D Mesoprephragma, anterior view pteralia E Mesopostphragma, posterior view O 3rd axillary wing sclerite, posterior F 3rd segment of left antenna, view ventral view 12. Subalare, dorsal view G roth segment of left antenna, O Caudal extension of 8th abdominal ventral view segment, dorsal view H Membranous area of scutum R Apex of penial sheath, ventral I 3rd axillary wing sclerite, dorsal view view Ss: Apex of penial sheath, ventral J Fore claw, posterior view view K Middle claw, posterior view at Tentorium and cranial apophysis ABBREVIATIONS USED IN THE FIGURES aas ante-anal setae als alar seta ab antennal bristles ams antemetaspiracular setae ads abdominal dorsal setae amss anterior metasternal setae aed aedeagus an anus
al alar lobe anp anterior notal wing process
MORPHOLOGY AND TAXONOMY OF ADULT MALES
anterior postalar ridge
abdominal sternite
additional sclerite
apical seta
anteprosternal seta
abdominal tergite
anterior tentorial arm
anterior tentorial pit
abdominal ventral setae
first axillary wing sclerite
second axillary wing sclerite
third axillary wing sclerite
basalare
basal membranous area
basal rod of aedeagus
sensilla basiconica
cicatrix
cranial apophysis
coxal bristle(s)
costal complex of wing veins
caudal extension
claw
coxa
dorsal head setae
dorsomedial part of midcranial ridge
dorsal ocular setae
dorsopleural setae
dorsal simple eye
dorsospiracular setae
mesepimeron
metepimeron
mesepisternum
metepisternum
postmetaspiracular setae
furca
segments of flagellum, 3rd to 1oth
femur
furcal pit
fleshy seta
gena
seta of glandular pouch
glandular pouch
genal setae
setae of genital segment
haltere
hair-like seta
interocular ridge
lateral branch of midcranial ridge
lateropleurite
lateral simple eye
mc mdr med mo mpns mr mts
median crest
median ridge
media
mouth opening
medial pronotal setae
marginal ridge
metatergal setae
ocellus
ocular sclerite
postalare
precoxal ridge of mesothorax
vestigial precoxal ridge of meta- thorax
pedicel
proepisternum + cervical sclerite
propleural apophysis
mesopleural apophysis
vestigial metapleural apophysis
propleural ridge
mesopleural ridge
metapleural ridge
postmesospiracular setae
posterior metasternal setae
mesopostnotum
metapostnotum
postnotal apophysis
posterior notal wing process
pronotal ridge
postocular ridge
posterior postalar ridge
prealare
lateral pronotal sclerite
pronotal ridge
preocular ridge
preoral ridge
prescutum
penial sheath
prescutal ridge
prescutal suture
post-tergite
posterior tentorial arm
posterior tentorial pit
peritreme of mesothoracic spiracle
peritreme of metathoracic spiracle
post-tergital setae
mesopleural wing process
vestigial metapleural wing pro- cess
radius
subalare
OF THE FAMILY COCCIDAE 11
set. scla subapical seta t tendon-like apodeme scl scutellum tar tarsus
sclf scutellar foramen tb tentorial bridge
scls scutellar setae tdgt tarsal digitule
scp scape teg tegula
set scutum tegs tegular setae
sctse scutal setae tib tibia
ser subepisternal ridge tibs tibial spur
SPy mesothoracic spiracle tp triangular plate
SP3 metathoracic spiracle tr trochanter
spl sensillum placodeum udgt ungual digitule
ss suspensorial sclerite vhs ventral head setae stn, prosternum vmcr ventral part of midcranial ridge stn, basisternum of mesosternum vps ventropleural setae stn, metasternum vs ventral sclerite
stn,s prosternal setae vse ventral simple eye stn,s basiternal setae
GENERAL MORPHOLOGY
The first serious attempt to study the morphology of the male of the Coccidae was made by Putnam (1879) in his paper on Pulvinaria innumerabilis. Apart from morphological observations, he suggested that the shape and proportions of the scutellum (which he called apodema) would be of some value in distinguishing the species of Pulvinaria. After Putnam’s work, attention was shifted almost com- pletely to the female, but some authors, e.g. Moulton (1907), Silvestri (IgIga, Ig1gb, 1920), Cusciana (1931), Hadzibejli (1955), Kawecki (1958b) and Husseiny & Madsen (1962) included brief descriptions of the male when describing the females of single species. Newstead (1903), Green (1904-1909), Leonardi (1920), Sule (1932) and Borchsenius (1957) each dealt with a series of females from definite localities ; they included short descriptions of the males available and usually gave a general account of the male of the family. Of all these, the works of Sulc and Borchsenius are the most significant. Sulc’s interpretation of the thoracic structures is fairly accurate, and he pays special attention to the eyes, halteres and chaetotaxy of the head in differentiating the species studied. Borchsenius’ paper contains a number of inaccuracies concerning the homologies of the various structures, but he describes and illustrates the differences in the head, 3rd and roth antennal segments, scutellum and terminal abdominal segments (including the genital segment) in the species that he studied. A small number of workers, namely Pesson (1941), Dtirr (1954), Habib (1956) and Bustshik & Saakjan-Baranova (1962) devoted papers to the description of the males of individual species, but their descriptions are rather superficial and contain many inaccuracies. Jancke (1955), Ezzat (1956) and Theron (1958) made comparative studies of the males of a number of families ; each of them included one member of the Coccidae : Physokermes piceae, Pulvinaria ericicola and Parthenole- canium pomeranicum (described as Eulecanium taxt) respectively. While Ezzat and Jancke contributed little that is morphologically significant, Theron gave a very detailed and accurate account, and was the first to make a comprehensive study of the pleural region.
12 MORPHOLOGY AND TAXONOMY OF ADULT MALES
In the present investigation it was possible, by studying a more representative sample (23 species), to substantiate Theron’s findings, to record a number of characters which have hitherto been unknown or overlooked, and to gain information on the range of variation within the family. In the general description of the morphology of male Coccidae that follows, Theron’s terminology is followed except where otherwise stated ; the account is illustrated by a generalized figure (Text- fig. 1) and all abbreviations in brackets refer to this figure, unless otherwise indicated.
General Characteristics
Compared with other Coccoidea, the males of the Coccidae are medium-sized, being smaller than the Margarodidae and larger than the Diaspididae. Among the species studied the smallest was L. luzulae (1020-1290, average 1141 yp long) and the largest EF. pela (2500-3100, average 2864 » long); L. /uzulae had the shortest wing-span (2090-2350, average 2213 uw) and E. pela the longest (5330-5700, average 5563 wu). Some species are slender in appearance (e.g. C. hesperidum) while others are rather robust (e.g. Cevoplastes spp.).
The body colour of living specimens varies from light reddish brown (E. festucae) to purplish (P. bituberculatum) ; the sclerotized areas are darker, brown to black, and the legs and the antennae yellowish ; the wings are hyaline, often with a purplish tinge between anterior margin and first wing vein.
In some species (e.g. the COCCUS group, Text-figs. 31-43) the head carries numerous setae, which give it a “ hairy’ appearance ; when the setae are few in number (the EFULECANIUM group, Text-figs. 2-23) the head looks rather bare.
The body is divided into the head, thorax and abdomen ; the head is separated from the thorax by a distinctly constricted “ neck’, characteristic of the Coccidae and Pseudococcidae.
The head (Text-fig. 1B, C) is irregular in shape, generally wide near the base dorsally, and narrowed anteriorly and ventrally. In dorsal view it is somewhat cone-shaped, broad and rounded posteriorly, with a tapering, more or less pro- truding apex. In lateral view the anteroventral surface slopes backwards towards the conspicuous, conical, medioventral bulge, which carries a pair of ventral eyes. The head is comparatively well sclerotized, with fewer and less developed secondary ridges than in the Diaspididae (see Ghauri, 1962) ; it carries 2-5 pairs of simple eyes and a pair of lateral ocelli. Mouth parts are absent, but an oval, non-functional mouth opening is present behind the medioventral bulge. The antennae are long, filiform and ten-segmented.
Thorax. The prothorax is considerably desclerotized, with a few more or less developed ridges and small sclerites. The mesothorax is well developed, sclerotized and with strong supporting ridges ; prescutum, scutum, scutellum and postnotum are all distinct, the latter curving down into the metathoracic cavity ; pleural sclerites are well developed, the pleural ridge strong ; basisternum large and usually with a median ridge. The fore wings are well developed, with two veins. The hind pair is reduced to halteres or absent ; when present each haltere has 1-4, apically hooked setae. The legs are long and slender with a one-segmented tarsus and a single claw. The meso- and metathorax each carry a spiracle laterally.
13
COCCIDAE
FAMILY
THE
4
OT
4\d---
uowp sy sy
Generalized drawing of the male of the Coccidae.
FIG. 1.
14 MORPHOLOGY AND TAXONOMY OF ADULT MALES
The abdomen is parallei-sided, with a slightly tapering posterior end. It consists of 8 pregenital segments and the genital segment ; the latter is narrow, elongated, partly sclerotized, and carries the genital organs. The pregenital segments are usually almost entirely membranous (e.g. P. pomeranicum), but sometimes with a small tergite and sternite on each segment (e.g. EF. tiliae). Sometimes (the COCCUS group) segments VII and VIII are each produced laterally into a finger- or lobe-like caudal extension ; posteriorly segment VIII usually carries a subdorsal glandular pouch, from the bottom of which two long setae arise ; they serve as a supporting core for a long wax filament.
The Head Head Capsule
Theron (1958) showed that, of the three regions of the generalized homopteran head (Weber, 1928 ; 1935), only one of them, i.e. the epicranium, makes up almost the entire head capsule in the Coccoidea ; of the other two regions the so-called “ Vorderkopf ”’ is reduced to a small area around the non-functional mouth opening, while the third, the labium, is absent altogether. Compared with other families, the head of the Coccidae is peculiar in having the anterodorsal part of the head capsule considerably expanded, with the apex sometimes produced into an antero- dorsal bulge. Apparently the enlarged dorsal part has shifted forwards and the ventral part backwards ; this process of transformation appears to resemble rotation about a point near the lateral ocellus. As a result some structures, which in the other Coccoidea are situated on the dorsal surface of the head, have been trans- located forwards or even to the ventral side. Thus the antennae occupy a ventral position, the median crest extends anteriorly over the apex of the head and termi- nates between the antennae, and the ventral part of the midcranial ridge does not reach the apex of the head. At the same time the dorsal membranous area of the posterior part of the head and neck has become extended. The degree of this deformation of the head varies within the family from a condition that is com- paratively normal and similar to the Pseudococcidae (e.g. E. tiliae, Text-fig. 3) to the extreme as shown by C. hesperidum (Text-fig. 32).
The dorsomedial part of the epicranium, which corresponds to the median crest (mc) of the Diaspididae (see Theron, 1958 ; Ghauri, 1962), is slightly raised, weakly sclerotized, and usually polygonally reticulated, but in a few cases (F. viburni, Ctenochiton sp. ; Text-figs. 14, 16 ; B) with only weak striations. The posterior margin is usually broadly rounded, but sometimes obtuse (J. theobromae, Text-fig. 29, B), and somewhat more heavily sclerotized, although no distinct ridge is present as found by Theron (1958) and Giliomee (1961) in the Pseudococcidae. The heavier sclerotization probably serves to strengthen the epicranium in a region where, according to Theron (1958), the mesothoracic muscles are attached. Borchsenius (1957) referred to the median crest as the “ cephalic longitudinal plate ’’.
In the anterior part of the median crest there is sometimes a linear vestigial ridge (dmcr), which corresponds to a similar structure found in Planococcus citri (Theron, 1958) and interpreted by Theron as a detached dorsal part of the midcranial ndge.
OF THE FAMILY COCCIDAE 15
It is best developed in N. abietis (Text-fig. 4, B), where it stretches anteriorly from behind the level of the posterior margin of the eyes to the anterior margin of the head, but even in this case it is not joined to the ventral part of the midcranial ridge (vmcr). The latter is usually well developed and Y-shaped. The median part of the ridge merges posteriorly into the ocular sclerite, but in some cases (J. theobromae, Text-fig. 29, C; the ERJOPELTIS group, Text-figs. 24, 26, 27, C) it fades away before doing so and in Ceroplastodes chiton Green it is vestigial. The lateral arms (lmcr) run to the base of each scape, but do not articulate with it. The arms are long in some species (J. theobromae, Text-fig. 29, C), short in others (Eriopeltis sp., Text-fig. 24, C), or even absent (E. ?festucae, Text-fig. 26, C). The area around the posterior section of the median part of the midcranial ridge is usually membranous and weakly polygonally reticulated. Rarely, however, the reticulation is entirely absent (Ctenochiton sp., Text-fig. 16, C ; Genus A, Text-fig. 20, C) or the area is both reticulated and sclerotized (Eviopeltis spp., P. myrtilli, I. theobromae ; Text-figs. 24 & 26, 14, 12, 29 ; C). This area corresponds to the ventromedial part of the epicranium of the more primitive margaroid Coccoidea, where it is sclerotized. The ventral part of the midcranial ridge was described as the ‘“‘chitinous impression’’ by Pesson (1941), the ‘‘ mesantennal plate ’’ by Borchsenius (1957) and the “ sclerotized fork ’’ by Bustshik & Saakjan-Baranova (1962). The Russian workers suggested, quite erroneously, that it might represent a rudiment of the mouth apparatus.
The large ocular sclerite (ocs), dorsally separated from the median crest by a membranous band surrounding the latter, constitutes most of the ventral and part of the lateral surface of the head capsule ; ventrally it extends uninterrupted from one side of the head to the other. It is distinctly polygonally reticulated. The ocular sclerite is partly bounded anteriorly by the preocular ridge (procr), which provides a process for articulation with the scape. In some species (the ERJO- PELTIS group, Ctenochiton sp., E. pela, Genus A ; Text-figs. 24, 26, 27 and 16, 18, 20 ; C) it is fused with, or closely approximates its opposite number, where it may also be joined by the median bar of the midcranial ridge. More often, however, it fades away at some distance from the articulating process.
The posterior margin of the ocular sclerite is bounded for the most part by the well developed postocular ridge (pocr). Dorsally the ridge originates behind the dorsal eye at about the level of the posterior margin of the median crest, passes behind the ocellus and extends posteromedially across the lateroventral surface of the head. Near the median line it curves backwards and extends for a short distance beyond the anterior end of the proepisternum + cervical sclerite, either as a definite ridge or as a small sclerite. In most species the ridge forks below the ocellus, with the short anterior branch surrounding or partly surrounding the ocellus. In the COCCUS group the postocular ridge is strong and thick through- out, but in the other groups the dorsal part of it is weaker, uniformly thin or gradually narrowing ; occasionally the part of the ridge immediately behind the ocellus is missing or very weak (R. spiraeae, Genus A ; Text-figs. 9, 21). In the literature the ridge has been illustrated, but not discussed by Leonardi (1920),
16 MORPHOLOGY AND TAXONOMY OF ADULT MALES
Silvestri (Ig1ga, Ig1gb, 1920), Sulc (1932) and Jancke (1955) ; it has been called the chitinous apodeme (Pesson, 1941) and lateral arches (Borchsenius, 1957 ; Bustshik & Saakjan-Baranova, 1962).
Sometimes the pre- and postocular ridges are joined together below the ocellus by a strong ridge ; this ridge has not been observed before and is here called the interocular ridge (ior). Its presence is constant in a few species (the ERIOPELTIS group ; Text-figs. 25, 28), where it is a broad ridge ; in one other species (Genus A ; Text-figs. 20, C ; 21) its occurrence is irregular : it is very narrow if present, some- times present on one side only and occasionally absent on both sides. On the preocular ridge in J. theobromae (Text-fig. 30) a small posteriorly directed process below the articular process apparently represents a rudiment of this ridge. The homology of the interocular ridge and its possible relation to the conditions found in the Pseudococcidae will be discussed later.
The ocular sclerite bears a number of simple eyes and a pair of lateral ocelli. The simple eyes comprise a pair of dorsal and a pair of ventral eyes, while 1-3 pairs of additional lateral ones may be present.
The dorsal eyes (dse) are situated on the anterolateral part of the head above the bases of the antennae and are widely separated from each other. The ventral eyes (vse) are located on the medioventral bulge close to each other in a submedian position ; the area between the ventral eyes is somewhat raised (e.g. FE. tiliae, Text-fig. 3) or flat (e.g. L. luzulae, Text-fig. 28). The lateral eyes (Ise) occur on each side of the head, more or less in line with the dorsal and ventral eyes. The dorsal and ventral eyes are usually large and subequal in size, while the lateral ones are considerably smaller. Sometimes (J. theobromae, Text-fig. 29, 30 ; Ceroplastodes chiton Green) the lateral eyes are only slightly smaller or as large as the others. The corneae of all these simple eyes are circular, deeply produced into the head- capsule and surrounded by a narrow membranous ring. The dorsal and ventral eyes are always present and are the only ones which occur in the ERIOPELTIS and COCCUS groups and some species of the EULECANIUM group (Text-figs. 24-43, 6, 10, 22) ; some other species have one (N. abietis, Text-fig. 4), two (R. spiraeae, P. myrtilli, Ctenochiton sp., Genus A, I. theobromae ; Text-figs. 8, 12, 16, 20, 29) or three (FE. tiliae, F. viburm, E. pela ; Text-figs. 2, 14, 18) additional lateral eyes. The lateral ocellus (0) usually appears as a weakly sclerotized spot on a membranous bulge, which is somewhat conical in Eviopeltis spp. (Text-figs. 24, 26). It is situated posterolateral to the dorsal simple eye, immediately anterior to the postocular ridge. The structure of the eyes and their innervation in P. corm were studied by Pflugfelder (1936). From his work it seems certain that the simple eyes on one side represent the isolated facets of a single compound eye. He also claimed that the lateral ocelli are persisting larval ocelli, a view already held by Putnam (1879) and Moulton (1907).
The large lateral bulge posterior to the postocular ridge corresponds to the gena (g). It is weakly sclerotized and in most species it is distinctly reticulated (e.g. Eviopeltis spp., Parthenolecanium spp. ; Text-figs. 24 & 26, 38 & 39). The sclerotization and
OF THE FAMILY COCCIDAE 17
reticulation were not described by Theron (1958). The deep cervical groove immediately behind the genae indicates the posterior margin of the head.
Ventromedially, immediately behind the ocular sclerite, the preoral ridge (pror) is situated. It has the form of an inverted V and is very narrow. It fuses posteriorly with the postocular ridge. Sometimes it is completely absent (the ERIOPELTIS group, Genus A ; Text-figs. 24, 26, 27 and 20 ; C).
The preoral ridge gives support to the cramial apophysis (ca), which is a strong scoop-like structure (Text-figs. 24, 39; T). Its apex is usually bifurcate, but sometimes trifurcate (Eviopeltis sp., Text-fig. 24, C) or truncate (E. pela, E. ?festucae, L. luzulae ; Text-figs. 18, 26, 27; C). In some cases the apex also carries an irregular central lobe, which is very pronounced in Genus A (Text-fig. 20, C). Theron (1958) found that in P. pomeranicum eight antennal muscles are attached to the apex of the cranial apophysis. The length of the cranial apophysis varies within the family. It is long in some species (e.g. C. hesperidum, Genus B ; Text- figs. 31, 33 ; C), reaching the level of the anterior margin of the ventral eyes, while in others (e.g. E. tiliae, P. piceae and E. pela ; Text-figs. 2, 6, 18) it does not extend beyond the posterior level of these eyes.
An irregular mouth opening (mo) is situated on a slight membranous bulge behind the cranial apophysis. On each side of the mouth opening, immediately median to the junction of the preoral and postocular ridges, a tendon-like apodeme (t) is present. According to Theron (1958) it serves for the attachment of a muscle which extends to the posterior margin of the median crest. In R. spiraeae this apodeme has a broad base and in Genus A it arises from the anterior part of an elongated sclerite (Text-fig. 20, vs) which appears to represent the ventral sclerite described by Theron (1958) in the margaroid Pseudaspidoproctus ?fulleri. The ventral sclerites were regarded by Theron as vestiges of the ventral plate found in Margarodes. They are also present in some Diaspididae (Ghauri, 1962).
The tentorial pits are situated in the membrane around the mouth opening. Ina few species (e.g. the ERIOPELTIS group ; Text-figs. 24, 26, 27 ; C) four tentorial pits are present. The two anterior ones (atp) are situated anterolateral to the mouth opening, near the preoral ridge when the latter is present ; these pits are, however, usually absent. The fosterior tentorial pits (ptp) are found posterolateral to the mouth opening, immediately median to the posterior ends of the postocular ridges ; they are always present. [rom each posterior pit a thread-like posterior tentorial arm (Text-figs. 24, 39 ; IT; pta) extends towards the heavily sclerotized tentorial bridge (Text-figs. 24, 29, T ; tb). From the bridge the somewhat stouter anterior tentorial arms (Text-figs. 24, 39 ; T; ata) extend towards the cranial apophysis. It is difficult to make out exactly how these arms are associated with the cranial apophysis. When the anterior tentorial pits are present, a thread-like anterior extension from each tentorial arm, possibly representing the dorsal tentorial arm links the anterior arms with the edges of the cranial apophysis (Text-fig. 24, T). When the anterior pits are absent, the arms are intimately associated with the cranial apophysis, as shown in Text-fig. 39, T. The latter condition appears to be the result of a process in which the anterior pits have drifted forwards until they
18 MORPHOLOGY AND TAXONOMY OF ADULT MALES
reached the cranial apophysis and apparently the anterior arms have completely fused with the “dorsal” ones. No sclerotized ventral cavity as illustrated by Theron (1958, fig. 22) is apparent and the illustration does not seem to agree with his description and fig. 13, where the ventral cavity is shown to be situated behind the preoral ridge. The tentorium has not been described by other workers, but from an examination of specimens in which the head is distorted, it is clear that what Borchsenius (1957) called the “‘ cephalic sclerotized arch”’ and Bustshik & Saakjan-Baranova (1962) regarded as the occipital ridge, is in fact the tentorial bridge.
Chaetotaxy
Two distinct types of setae, similar to those found in Pseudococcus (Giliomee, 1961), are present on the head of male Coccidae. They are:
(i) a thick-set, fleshy type (fs), which has a rather blunt apex and the setal mem- brane not surrounded by a distinct basal ring, and
(i1) a slender, hair-like type (hs) which has an acute apex and the setal membrane surrounded by a strong basal ring.
Generally the hair-like setae occur in all the species in comparatively small numbers and when only these setae are present the head appears to be rather bare. In those species where fleshy setae are also present the head has a “ hairy ”’ appear- ance, the number of these additional setae being generally much larger.
The setae on the head are arranged in the following groups:
(i) Dorsal head setae (dhs), which are situated on the median crest, but may also occur on the surrounding membrane in front of the dorsal simple eyes. This group consists either of hair-like setae only (the EFULECANIUM and ERIOPELTIS groups) or of both fleshy and hair-like unes (the JNGLISIA and COCCUS groups). The number of hair-like setae, which are present in all the species, varies from I—7 (average 5) in P. acericola to 16-19 (average 17) in P. cormi ; the number of the additional fleshy setae, in the species in which they occur, varies from 4—I1 (average 7) in I. theobromae to 26—42 (average 35) in C. hesperidum.
(ii) Dorsal ocular setae (dos), which are found on each side on the dorsal part of the ocular sclerite between the dorsal simple eye and the postocular ridge, and consist of both fleshy and hair-like setae. The total numbers of these setae are small and variable and the proportion of the two types of setae is also variable within the species.
(11) Ventral head setae (vhs) : This group occurs on the ventral and lateral parts of the ocular sclerite and may extend up to the lateral arms of the midcranial ridge. In the COCCUS and INGLISIA groups, but not in the other species examined, the setae also occur between and behind the ventral eyes. In most species of the EULECANIUM group the setae are situated only on or anterior to the level of the preocular ridge. In a few species (Genus A, S. prunasin, L. luzulae; Text-figs. 20, 22, 27 ; C) one pair of median hair-like setae is distinctly longer than the other setae. The hair-like setae occur in all the species in small numbers, varying from 1-2 (average 1-3) in N. abietis to 7-17 (average 12) in E. ? festucae, but in R. spiraeae
OF THE FAMILY COCCIDAE 19
they may be entirely absent in some individuals. The additional fleshy setae occur only in the ERIOPELTIS, COCCUS and INGLISIA groups (Text-figs. 24-43) and here in rather large numbers, varying from g—16 (average 12) in Eviopeltis sp. to 54-108 (average 84) in Pulvinaria ?ribesiae.
(iv) Genal setae (gs), which are found on the genae and are only present in the COCCUS and INGLISIA groups (Text-figs. 29-43). They consist of both fleshy and hair-like setae. The number of hair-like setae is usually small, varying from 0-4 (average 1-2) in J. theobromae to 7-13 (average 9) in P. cornt, and the number of fleshy setae large, varying from 5—I1 (average 7) in J. theobromae to 17—30 (average 23) in P. cornt.
No other dermal structures, like disc pores or specialized sensilla are present on the head.
Antennae
All the workers that have studied the head of male Coccidae mention the antennae, but their descriptions are very brief and usually cover little more than the number of segments and the general shape of the antennae.
The antennae are inserted fairly low down on the anterolateral margin of the head with the diverging lateral arms of the midcranial ridge between them. They are generally filiform in shape and normally comprise ten segments, but sometimes two or more of the segments between the 4th and gth are intimately fused. The relative length of the antennae varies considerably within the family. They are very long in the ERIOPELTIS group, being about 3-2 as long as the body in Eniopeltis sp., and short in the COCCUS group, being usually less than half as long as the body. The antennae with the shortest relative length are found in EF. tiliae (EULECANIUM group) where they are }-? as long as the body. The average absolute length varies from 622 y in C. berliniae to 1922 uw in E. pela.
’
The antennae always carry a large number of setae, which give them a “ hairy ’ appearance. The setae consist of both fleshy and hair-like ones, similar to those occurring on the head. The fleshy setae (fs) appear usually only on the 2nd to roth segments, but in FE. pela they occur regularly on the Ist segment as well. They are usually slightly longer than the width of the antennal segments, although a few on the distal part of the 4th to 9th segments may be considerably shorter. In E. pela, however, the fleshy setae are exceptionally long, being about 4-5 times as long as the width of the 3rd segment, and in Genus A they are very short and stout, and only about half as long as the width of the 3rd segment. The hair-like setae (hs) are always present on the first two segments and in a number of species (e.g. F. tiliae, F. viburm, Genus A, Text-figs. 2, 14, 20 ; F) also on the 3rd, but only in F. viburni and Genus A do they occur regularly on the 4th to 1oth segments. In addition to the fleshy and hair-like setae two different types of setae are found on the distal segments. The setae of one of them somewhat resemble the ordinary fleshy setae, but are usually much larger, bristle-like and have a large setal membrane ; they can be called antennal bristles (ab). On the roth segment there also occur the long,
20 MORPHOLOGY AND TAXONOMY OF ADULT MALES
capitate, subapical setae (set. scla) found in the Pseudococcidae (Sule 1943 ; Giliomee, 1961) and Diaspididae (Bustshik, 1958 ; Ghauri, 1962). Sule called them setae semi-claviformes. Both types probably have a sensory function. Specialized sensilla are present on the 2nd, 3rd and roth segments.
The scape (scp) is short, wide and subrectangular in shape, with the basal part sclerotized and the distal part mostly membranous. The dorsal margin is longer than the ventral one. The scape articulates with the pedicel by means of a ventral projection, which is situated opposite a corresponding projection on the basal ridge of the pedicel. Posterolaterally it articulates with the preocular ridge by means of a process on its basal ridge. The scape usually carries 3 hair-like setae and in E. pela 2-4 (average 2:6) fleshy setae also occur on this segment.
The pedicel (pdc) is short, broad and subglobular in shape. It is generally well sclerotized with the distal part, especially dorsally, distinctly polygonally reticulated. In P. myrtilli (Text-fig. 12, B), the reticulation is represented by a few wavy lines and in P. ?betulae (Text-fig. 35, B) and J. theobromae (Text-fig. 29, B) the reticulation is usually entirely absent. The base of the pedicel is partly surrounded by a strong basal ridge which is well developed ventrally, but weaker or absent dorsally. At the dorsal end there is a shallow depression in which the constricted basal part of the 3rd segment is received. Both hair-like and fleshy setae are present ; they are mostly located on the ventral and lateral surfaces. A small circular sensillum, probably a sensillum placodeum, is situated dorsally or dorsolaterally. It is also present in the Pseudococcidae (Giliomee, 1961) and Diaspididae (Ghauri, 1962).
The flagellum (Fiii_x) 1s composed of the 3rd to roth segments. The 3rd segment (Text-figs., F) varies considerably in size and shape ; in Eviopeltis spp. (Text- figs. 24, 26; F) it is long and club-shaped, while it is short and barrel-shaped in P. ?betulae (Text-fig. 35, F). This segment carries ventrally a number of small sensilla (bs), which are probably sensilla basiconica. Their number varies indi- vidually and may do so on the two antennae of the same specimen, but it rarely exceeds a total of four. Sensilla were also found in the same area in the Pseudo- coccidae (Giliomee, 1961) and Diaspididae (Ghauri, 1962). The 8th and gth seg- ment each has an antennal bristle, which is sometimes difficult to distinguish from the fleshy setae ; the bristle is situated ventrally near the apex.
In most species the roth (terminal) segment is broadly rounded at its apex, but in some (e.g. C. hesperidum ; Text-fig. 31, G) the distal part of the segment is tubularly constricted. In addition to the fleshy and occasional hair-like setae this segment also bears a number of antennal bristles (ab) and capitate subapical setae (set. scla). The former consist of 3 long and 2 shorter setae. The subapical setae are usually 3 in number, but in J. theobromae there are only 2 and in Genus A there are 4-6 (average 5). On the ventral surface of the roth segment 2 small sensilla (bs) are found, one near the apex and the other somewhat more proximal. They are probably sensilla basiconica and correspond to sensilla on this segment in the Pseudococcidae (Giliomee, 1961) and Diaspididae (Ghauri, 1962).
OF THE FAMILY COCCIDAE 21
The Thorax Prothorax
The prothorax is largely membranous, with only a few sclerites and ridge-like structures present. It is distinctly separated from the head by a deep cervical constriction. In this respect the Coccidae resemble the Pseudococcidae (Theron, 1958 ; Giliomee, 1961).
Dorsally, immediately behind the neck region, the collar-like pronotal ridge (prnr) runs continuously from one side to the other, extending ventrally and closely approxi- mating the proepisternum + cervical sclerite. It is usually interrupted by weak sclerotization dorsomedially, but in E. pela the ridge is apparently uninterrupted, although very narrow medially. This structure has been called the “ protergal sclerite ’’ (Habib, 1956), the “‘ prothoracic suture’”’ (Ezzat, 1956) and the “ pro- thoracic arch ’’ (Borchsenius, 1957). Dorsolaterally, behind the pronotal ridge, a small sclerite is closely associated with it ; this sclerite appears to be homologous with the lateral pronotal sclerite (prn) described by Giliomee (1961) in the Pseudo- coccidae and Ghauri (1962) in the Diaspididae. The sclerite was called the “prothoracic sclerotized plate’’ by Borchsenius (1957). Theron (1958) did not mention them.
In the posterolateral part of the prothorax a small sclerite is situated ; it con- stitutes the so-called post-tergite (pt). It sometimes shows irregular wavy striations (e.g. most species of the COCCUS group). In Eriopeltis sp. the sclerite is apparently absent and only represented by striations of the derm. The post-tergites have not been observed in this family before.
In the pleural region the pleurites and neck sclerites are reduced to a single ridge- like structure, called the proepisternum + cervical sclerite (pepcv) (Ghauri, 1962). Anteriorly it passes just below the ventral end of the pronotal ridge and appears to be joined by weak sclerotization to the postocular ridge. For a short distance behind the level of the pronotal ridge this sclerite is less strongly sclerotized. This phenomenon was also observed in Pseudococcus (Giliomee, 1961). Posteriorly the sclerite is delimited by a short pleural ridge (plr,), which articulates ventrally with the basal process of the coxa ; dorsally it is invaginated to form a small pleural apophysis (pla,). Crampton (1926) called the structure a “neck plate” or “ Jaterocervicale ’’ in Coccus. He also distinguished an episternum and epimeron, but no structure corresponding to the latter was observed in the species studied here. From his illustration (fig. 55) it appears that he misinterpreted the position of the pleural ridge. The proepisternum + cervical sclerite has also been referred to as the “ pleural sclerite of the prothorax ”’ (Ezzat, 1956), the “‘ propleural sclerite ”’ (Habib, 1956) and the “ sclerotized plate of the anterior leg’ (Borchsenius, 1957).
The prosternum (stn,) is represented by a single sclerite, of which the degree of development shows considerable interspecific and also some intraspecific variation. In its most complete form it consists of a triangular, sometimes oval sclerite, which is bounded posteriorly by a strong transverse ridge and traversed by a longitudinal median ridge. Sometimes the prosternal sclerite is more or less reduced (C. hesperidum, Text-fig. 31), while the median ridge may be complete (e.g. P. myrtilli,
22 MORPHOLOGY AND TAXONOMY OF ADULT MALES
C. hesperndum ; Text-figs. 12, 31), interrupted in the middle (e.g. E. tihae, E. Pfestucae, P. cornt ; Text-figs. 2, 26, 38), developed anteriorly only (e.g. Eviopeltis sp., P. pomeramcum ; Text-figs. 24, 39), or represented by a basal stalk (R. spiraeae, L. luzulae ; Text-figs. 8,27). In some species the degree of reduction of the median ridge varies individually, e.g. in F’. viburni and P. ?betulae the ridge may be complete or developed anteriorly only, while in P. bituberculatum and S. prunastni it may be absent or represented by a short basal stalk only. On each side of the transverse ridge a shallow depression probably represents the sternal apophysis. Its position corresponds to that of the sternal apophyses in the more primitive Phenacolea- chiidae (Theron, 1962). In some individuals of most species a small apopysis is situated medially anterior to the prosternum. It probably serves for the attachment of muscles, as four muscles originate in the corresponding area in the Pseudococcidae (Makel, 1942). Theron (1962) describes a “‘ mammillate organ’’ from the same region in the Phenacoleachiidae, saying that it is probably homologous with the so- called salivary gland of Pseudaspidoproctus. The derm of the prosternum is occasion- ally polygonally reticulated (C. hesperidum, Text-fig. 31) or covered with numerous small spines (I. theobromae, Text-fig. 29). The prosternum was overlooked by most earlier workers. Crampton (1926) figures a linear basisternum and sternellum, while Ezzat (1956) called this region a basisternum ; Borchsenius (1957) regarded it as part of the mesosternum.
Dermal Structures. Both fleshy and hair-like setae are present in various regions of the prothorax. They occur in the following groups :
(i) Lateral pronotal setae (Ipns), which occur on or immediately posterior to the lateral pronotal sclerites on each side, and may consist of up to 3 fleshy or hair-like setae. They are of very little taxonomic significance as they are only present in some individuals of certain species (e.g. P. pomeranicum, Text-fig. 39).
(ii) Medial pronotal setae (mpns), which are situated between the pronotal ridge and the post-tergites and usually consist of two widely separated hair-like setae (e.g. P. myrtilli, S. prunastni, C. hesperidum, Text-figs. 12, 22, 31) ; in P. bituber- culatum (Text-fig. 10) the two setae are situated close together on the median line. In other species (e.g. N. abietis, I. theobvomae) one or both setae may be absent, while medial pronotal setae are absent altogether in some species (e.g. EF. tiliae, L. luzulae). One or two fleshy setae are occasionally associated with these setae. Medial pronotal setae are also found in the Pseudococcidae (Giliomee, 1961).
(ii1) Post-tergital setae (pts) occurring on, or behind and below the post-tergites. They consist of fleshy setae only (up to 13) and were found in the closely related genera Pulvinaria, Coccus and Genus B (Text-figs. 31, 33, 35, 37), but not in the other species.
(iv) Anteprosternal setae (astn,s) consisting of a number of fleshy setae (up to 7) which occur immediately ventral to the anterior part of the prosternum + cervical sclerite (pepcv). They are present in the COCCUS group (Text-figs. 31-43) and sometimes in L. luzulae.
(v) Prosternal setae (stn,s), which are found on and around the prosternum, anterior to the level of the transverse ridge ; in the COCCUS group they extend
OF THE FAMILY COCCIDAE 23
laterally to occur anterior to the mesothoracic spiracle. This group therefore corresponds to the prosternal and antespiracular ventral setae of the Pseudococcidae (see Giliomee, 1961). They include both fleshy and hair-like types. The number of these setae varies considerably individually and within the family. The fleshy setae are usually numerous (up to 54) in the COCCUS group and less numerous (up to 25) in the ERIOPELTIS and INGLISIA groups; they are absent in the EULECANIUM group with the exception of S. prunastri, which has 7-16 setae. The hair-like setae are never more than 4 in number and are often entirely absent.
In Ctenochiton sp., C. hesperidum and Genus 6 a number of circular pores, some- what reminiscent of vacant hair sockets, are situated on each side dorsally, posterior to the pronotal ridge. They number 3-7 (average 6-1), 0-1 (average 0-4) and 2-5 (average 3:6) respectively.
Mesothorax
The mesothorax, as the principal wing-bearing segment, is well developed and sclerite degeneration is much less pronounced than in the other thoracic segments ; in addition, some of the sclerites are bounded by strong ridges. The shape of the sclerotized areas varies comparatively little within the family and consequently provides only a few characters of taxonomic importance.
Mesotergum. The usual subdivisions of the mesotergum can easily be discerned. Thus the notum (or alinotum) is widely separated from the postnotum, the former being distinctly subdivided into a prescutum, scutum and scutellum ; this was already recognized by Sulc (1932), Pesson (1941) and Jancke (1955).
The prescutum (prsc) is situated anteromedially and is surrounded laterally and posteriorly by the scutum. It has the shape of a large subrectangular bulge. Anteriorly it curves sharply downwards and forms the mesoprephragma (Text-figs. D) ; the latter has the shape of a simple lamina with the inner margin slightly emarginated in the middle. This emargination varies somewhat individually, but it is inconspicuous or absent in some species (e.g. FE. 2festucae, Text-fig. 26, D) and pronounced in others (e.g. C. hesperidum and Genus B ; Text-figs. 31, 33; D). The phragma was regarded as the prescutum by Ezzat (1956). Laterally the prescutum is separated from the scutum by strong prescutal ridges (pscr), which are fused anteriorly with the mesoprephragma, and extend posteriorly for some distance along the sides of the membranous area of the scutum. The posterior margin is bounded by the prescutal suture (pscs) with its corresponding internal ridge. The median part of the prescutum is often more heavily sclerotized and sometimes a median, ridge-like structure occurs at the posterior (P. bituberculatum, Text-fig. 10) or near the anterior margin (Ceroplastes spp.). In some species the cuticle of the prescutum shows reticulation, which may be regularly polygonal (e.g. P. piceae, L. luzulae, Parthenolecanium spp. ; Text-figs. 6, 27, 38 and 39) or irregular (C. hesperrdum, Genus B ; Text-figs. 31, 33). The prescutum has been called the “ scutum of the prothorax ”’ (Putnam, 1879), the “ proscutum ”’ (Jancke, 1955) and the “ scutum ”’ (Ezzat, 1956).
24 MORPHOLOGY AND TAXONOMY OF ADULT MALES
The scutwm (sct) has a rather curious shape. The median membranous area, which is comparatively small in the margaroid Pseudaspidoproctus and Steingelia (Theron, 1958), has in this family become so expanded that it completely and widely separates the two lateral sclerotized parts. These extend anteriorly along the sides of the prescutum and posteriorly along the sides of the scutellum. In the anterolateral region the scutum is produced into a prealare (pra), which is separated from the former by an internal lamina. The prealare is semitubular in shape, with the more heavily sclerotized and infolded anterior margin continuous with the mesoprephragma. The distal part of the prealare is differentiated into a heavily sclerotized, convex tviangular plate (tp), which extends to the episternum. Behind the prealare the lateral margin of the scutum is infolded and somewhat more heavily sclerotized, the infolded section of the margin terminating in a small rounded projection which constitutes the anterior notal wing process (anp). From this level the posterior extension of the scutum is depressed, laterally emarginated and with a rounded posterior lobe which can be regarded as the posterior notal wing process (pnp). The posterior margin of the scutum probably incorporates the lateral part of the so-called marginal fold of the notum ; posterolaterally it is attached to the postalare by means of a sclerotized band. Part of the scutum adjacent to the scutellum is more heavily sclerotized and usually reticulated. This probably led Theron (1958) to misinterpret it as part of the scutellum, as indicated in his illus- tration of P. pomeramicum. The anterior part of the scutum may also show reticula- tion (e.g. Ceroplastes spp., Text-figs. 41, 43) and even the median membranous area may be weakly reticulated (Eviopeltis sp., L. luzulae ; Text-figs. 24, 27).
The scutellum (scl) in dorsal view has the shape of a transverse rectangle. The anterior and posterior edges, constituting the scutoscutellar ridge and the posterior marginal fold of the notum respectively, curve sharply inwards, are deflected under the scutellum and extended internally. The inner edges usually have become intimately fused, leaving only an oval median foramen (sclf) (F. viburni, Ctenochiton sp., S. prunastnn, ERIOPELTIS group, COCCUS group). This gives the scutellum the appearance of a dorsoventrally flattened tube. In some species, however, the inner edges do not unite with each other (J. theobromae and most species of the EULECANIUM group). In the species where the scutellum is tubular it is usually shorter and wider than in those where the scutellum is not tubular. The scutellum was called the “ apodema’”’ by Putnam (1879), Green (1904-1909) and Durr (1954) while Habib (1956), Borchsenius (1957) and Bustshik & Saakjan-Baranova (1962) regarded it as part of the scutum. The scutellar foramen was referred to as a “membranous area ”’ by Pesson (1941), Jancke (1955), Ezzat (1956) and Borchsenius (1957). Bustshik & Saakjan-Baranova (1962) state that it is absent in C. hesperidum.
The scutellum is followed by a large, subtriangular membranous area which separates it from the postnotum. This membranous area was regarded as the scutellum by Putnam (1879), Green (1904-1909), Habib (1956) and Borchsenius (1957), and as the postnotum by Sulc (1932), Pesson (1941) and Jancke (1955). The postnotum (png) is a curved structure which extends deeply into the metathoracic cavity and is overlapped by the similarly inflected metanotum. The anterior margin
OF THE FAMILY COCCIDAE 25
of the postnotum is usually weakly sclerotized and irregular, and may be exposed (e.g. P. myrtilli, Ctenochiton sp., Eriopeltis spp, Ceroplastes spp. ; Text-figs. 12, 16, 24 & 26, 41 & 43) or medially overlapped by the metathoracic fold (e.g. E. tilzae, Genus A, I. theobromae ; Text-figs. 2, 20, 29), but this varies somewhat individually. Anterolaterally the postnotum bears a deep finger-like apophysis (pna). Some- times the whole postnotum is polygonally reticulated (Ceroplastes spp.) or reticulation occurs near the anterior margin only (P. corni). At the posterior margin of the post- notum a mesopostphragma is formed which is usually deeply emarginated medially. On each side the postnotum is produced into a strong postalare (pa), which extends anterolaterally to articulate with the mesopleural ridge. The anterior postalar ridge (apar) is well developed, while the posterior postalar ridge (ppar) is weak. The anteroventral part of the postalare is densely reticulated. Dorsally the postalare is produced into two small processes ; the hind margin of the wing is attached to the anterior one, and the posterior marginal fold of the notum to the pos- terior.
Mesopleuron. A striking feature of the mesopleural region is the strong meso- pleural ridge (plr,)._ The ridge winds obliquely across the pleuron, with a sharp bend which separates the strongly developed vertical part from the weaker ventral section, the latter extending obliquely towards the coxa. The dorsal part of the ridge gives firm support to the pleural wing process, the ventral extremity articulates with the coxa. The ventral part is partly overlapped by the postalare and at this point a pleural apophysis (plag) is invaginated. In some species (e.g. F°. viburni, Ctenochiton sp., E. pela ; Text-figs. 15, 17, 19) the ridge fades away into the pleural sclerite, as is also the case in the margaroid Steingelia (Theron, 1958) and some Pseudococcidae (Giliomee, 1961). The pleural wing process (pwp,.) is a large rounded structure. On its lower anterior margin there is a small tendon-like apodeme (t) from which a muscle extends anterodorsally to the tegula. Posterodorsal to the pleural wing process a small, meniscate swbalare (Text-figs. 1, A,, sa; 18, N, sa ; 18, P) is found. Dorsally it is produced into a finger-like process, which apparently articulates with the second axillary sclerite (Text-fig. 18, N). The basalare (bas) isa narrow but distinct sclerite (in the EFULECANIUM, ERIOPELTIS and INGLISIA groups) which connects the anterior margin of the wing process with the episternum, or it is vestigial and incorporated into the pleural wing process (in the COCCUS group). The vertical part of the pleural ridge is separated from the episternum by a strip of membrane which corresponds to the basalar cleft of Matsuda (1960). The episternum (epsz) is large and well sclerotized ; a membranous cleft, extending anteriorly from the region of the pleural apophysis completely divides it into dorsal and ventral parts. The dorsal part is strongly convex and sometimes reticulated (e.g. C. hesperidum, Ceroplastes spp ; Text-figs. 17, 40 & 42) ; the ventral part is a narrow elongated sclerite which joins the lateropleurite anteriorly. The dorsal part is bounded anteriorly by a well developed subepisternal ridge (ser). This ridge extends dorsally from the triangular plate of the prealare towards the marginal ridge of the basisternum. Below the membranous cleft, however, it is reduced and only marked
20 MORPHOLOGY AND TAXONOMY OF ADULT MALES
by a band of darker sclerotization. The latevopleurite (Ipl) is always well developed and the anterior margin often bounded by an extension from the anterior part of the marginal ridge (e.g. EF. tuliae, L. luzulae, I. theobromae ; Text-figs. 2,27, 29). The epimeron (epm,) is represented by a small sclerite posterodorsal to the coxal articula- tion. The mesothoracic spiracle (spy), with its supporting peritreme (ptr,), is situated in the membrane anterior to the subepisternal ridge. Except for Theron (1958), none of the earlier workers on the Coccidae studied the mesopleuron in any detail. Ezzat (1956) referred to the subepisternal ridge as the “ pleural bridge ”’ and to the pleural ridge as the “ pleural sclerite’’. It is difficult to determine with certainty the homologies of the structures described above with those of the basic pleurosternal region, as proposed by Matsuda (1960). The membranous cleft probably represents or incorporates his anapleural cleft. According to his defini- tions, the dorsal part of the episternum then represents the anepisternum (called pre- episternum by Weber, 1928) and the ventral part of the pre-episternum, with the katepisternum either absent or incorporated into the latter. Theron (1958), following Weber (1928), referred only to the area anterior to the ventral part of the subepisternal ridge as a lateropleurite (pre-episternum). Roberti (1946) called the same area a laterosternite, a term used by Weber (1928) to describe a more ventral part of the precoxal region.
Mesosternum. The mesosternum is almost entirely represented by the large hexagonally shaped basisternum (stn,). At the junction of the basisternum and episternum there is a strong marginal ridge (mr), which extends medially to delimit the basisternum anteriorly. Posteriorly it unites with the precoxal ridge and further posteriorly the ridge fuses with the pleural ridge immediately above the coxal artic- ulation. The strong precoxal ridge (pcre) curves round the posterolateral edge of the basisternum, but fades away before reaching the median line. A strong longi- tudinal median ridge (mdr) completely divides the basisternum into two halves ; sometimes, the ridge is more or less reduced (Eviopeltis spp. ; Text-figs. 24, 26) or vestigial (I. theobromae ; Text-fig. 29). The posterior margin of the basisternum is invaginated to form a transverse furcal pit (fp) from which a well developed furca (f) originates. The furca consists of a broad basal stalk and two strong furcal arms. No separate sternellum is found ; Makel (1942), from a study of some Pseudococcidae suggested that it is represented by the base of the furca. The sclerite which Ghauri (1962) regards as the sternellum in the Diaspididae is probably part of the meta- thorax (as discussed later). Ezzat (1956) referred to the basisternum as the “ furca- sternum ”’ and called the transverse part of the marginal ridge a “ sternacostal
,
suture’; Borchsenius (1957) called the basisternum a “‘ mesosternal frame ’’.
Articulation of the wings. The articulation of the wing is facilitated by a number of minute alary sclerites or pteralia which lie embedded in the basal articular membrane of the wing (Text-figs. 1 ; 18, N). They consist of the tegula, the first, second and third axillary sclerites, and the additional sclerite. Other structures involved in the wing articulation are the anterior notal wing process, the pleural wing process, the epipleurites and the costal complex of wing veins.
OF THE FAMILY COCCIDAE 27
The small, meniscate tegula (teg) is situated far anterior to the wing base, from which it is separated by a large membranous bulge. This bulge carries a small, weak sclerite posteriorly and in some cases (e.g. the ERIOPELTIS group) it is weakly reticulated. The first axillary sclerite (Text-figs. 1 ; 18, N ; ax,) is triangular in shape and its mesal edge lies against the lateral margin of the scutum just behind the anterior notal wing process. The anterior part of the sclerite is drawn out into a slender arm, which curves round the anterior apex of the second axillary sclerite and articulates with the costal complex of wing veins. The posterolateral part of the sclerite articulates with the second axillary sclerite. The first axillary sclerite rather closely resembles those of the more primitive Coccoidea described by Theron (1958). The second axillary sclerite (Text-figs. I ; 18, N ; axg) is elongate, slightly curved, with both the anterior and posterior apices acute. The anterior part articu- lates with the first axillary and the posterior apex with the third axillary sclerite. Apparently it also articulates with the subalare which hes directly below it. The structure of the third axillary sclerite (Text-figs. 1 ; 18, N ; ax,) is more complex and it shows some variation in the species studied. The distal, somewhat triangular part articulates anteriorly with the posterior apex of the second axillary while its posterior margin is confluent with the hind margin of the wing. Mesally this plate extends into a narrow arm which is twisted in such a way that the plane changes from horizontal to vertical (Text-fig. 18, O) ; this arm has an anterior, scoop-like extension which is attached to a process on the postalare by means of a tiny axillary cord. In some species (e.g. ERIOPELTIS group, C. hesperidum, Parthenolecanium spp.; Text-figs. 24, 26, 27 and 31, 38, 29 ; I) this anterior extension is very small or absent. The additional sclerite (asc) is situated at the base of the wing immediately distal to the second and third axillary sclerites, but does not articulate with them. It is weakly sclerotized and irregularly elongate. As suggested earlier (Giliomee, 1961) this structure may represent the second median sclerite, defined by Snodgrass (1935).
The costal complex of wing veins will be discussed later.
Chaetotaxy. Both fleshy and hair-like setae may occur on the mesothorax. Setae are found on the scutum, scutellum, tegular bulge, the membrane anterior to the basisternum and episternum, and on the basisternum itself. No setae were found on the prescutum.
(i) The scutal setae (sctse) are scattered over the median membranous area of the scutum. In some species (Ceroplastodes chiton Green, and sometimes in E. ?festucae) they also extend beyond the posterolateral corner of the membranous area to occur on the sclerotized parts. Inanumber of species (I NGLIS/A group and most species of the COCCUS group) the scutal setae comprise both fleshy and hair-like setae in various proportions, but usually in fair numbers (e.g. 10-32, average 24 fleshy setae in C. hesperidum and 14-22, average 18 hair-like setae in Genus 5). In the other species the scutal setae are either absent (F. tiliae, N. abietis, P. piceae, R. spiraeae and E. pela) or consist of hair-like setae only ; the hair-like setae may be few (up to 4) in some species (P. bituberculatum, P. myrtilli, S. prunastri) and numerous (up to 30) in others (e.g. Pulvinania spp.).
28 MORPHOLOGY AND TAXONOMY OF ADULT MALES
(1) Scutellar setae (scls). One or two hair-like setae are sometimes present on the scutellum, but their occurrence is very irregular and variable even within a species.
(iii) The ¢egular setae (tegs) are carried on the anterior part of the tegular bulge and consist usually of a small number (up to 11) of hair-like setae. Fleshy tegular setae were only observed in Ceroplastodes chiton Green and occasionally in J. theobromae.
(iv) The postmesospiracular setae (pms) are arranged in a broad band on the membane posterior to the prosternum and mesothoracic spiracles. When they are numerous, some setae may also occur on the episterna. They consist almost entirely of fleshy setae and are only present in the COCCUS and INGLISIA groups. Their numbers vary from 14—29 (average 21) in Ceroplastes to 71-97 (average 89) in P. cornt.
(v) The basisternal setae (stn,s) are situated on or near the median ridge in the posterior part of the basisternum. They were found in only 3 species, consisting of 1 or 2 hair-like setae in N. abietis and P. piceae (Text-figs. 4, 6) and 2—9 (average 5) fleshy setae in Genus B (Text-fig. 33).
Metathorax
The metathorax is very weakly sclerotized and the sclerites have to a large extent been replaced by membrane ; this is due to the reduction of the hind wings. The metanotum, however, is relatively well developed. It consists of a large plate which closely overlaps the invaginated mesopostnotum. The dorsal edge (morphologically posterior margin) of the sclerite is heavily sclerotized, forming a ridge-like structure which usually extends continuously from one side to the other, but in a number of species (e.g. EF. tiliae, F. viburm, Ceroplastes spp.) its median part is somewhat desclerotized. Externally the metanotum is represented by a small, lateral suspen- sorial sclerite, which is connected to the haltere by means of a sclerotized band. The suspensorial sclerites are absent when the halteres are lacking. Somewhat more posteriorly an additional, small, weak sclerite is sometimes present (e.g. in E. tiliae, L. luzulae, P. corni, Text-figs. 2, 27, 38), but 1t may be absent or present within the same species. In the intersegmental region between the metathorax and Ist ab- dominal segment there is an irregular, transverse, lateral sclerite which corresponds to the acrotergite or postnotum (pns) of the more primitive Coccoidea (see Theron, 1958). In one of the species (L. luzulae, Text-fig. 27) the sclerites of the opposite sides meet or closely approximate each other, and in another species (J. theobromae, Text-fig. 29) they are divided by the intersegmental line.
The degree of development of the pleural region depends to a considerable extent on the absence or presence of halteres. The pleural ridge (plr,) extends from the coxal articulation in an anterodorsal direction across the pleuron. When the haltere is absent (e.g. ERIOPELTIS, INGLISIA and COCCUS groups) the ridge only extends for a short distance above the coxal articulation. When the haltere is present, however, (most of the EFULECANIUM group), it extends towards the base of the haltere where it is slightly expanded to form a small metapleural wing process (pwps;). In this condition the ridge becomes weaker or is interrupted at about half- way from the coxal articulation ; a shallow depression in this area appears to represent a reduced metapleural apophysts (plas).
OF THE FAMILY COCCIDAE 29
The episternum (epss) is small and subtriangular in shape, but when the haltere is present it expands in a ventral direction ; in some species (e.g. E. tiliae and R. spiraeae ; Text-figs. 3, 9) its anterior margin is partly bounded by a more or less developed ridge, resembling the subepisternal ridge of the mesothorax. The epimeron (epm,) is represented by an irregular, posteriorly produced sclerite. In most species a vestigial precoxal ridge (pcr,) extends anteriorly for a short distance along the ventral margin of the episternum, but the absence or presence of this ridge varies individually. The metathoracic spiracle (sp), supported by a peritreme (ptrs), is situated in the membrane anterior to the episternum.
The metasternum (stn) is usually represented by a fairly large, irregular, median plate, which is generally more heavily sclerotized anteriorly and weaker posteriorly, but in some species (e.g. S. prunastn’, Text-fig. 22 ; Ceroplastes spp., Text-figs. 41, 43) its posterior part is entirely membranous and only a narrow strip of it remains anteriorly. In Eviopeltis sp. (Text-fig. 13) small and irregular sclerotized areas are situated anterior to this plate. The sternal apophyses are absent and this makes it difficult to establish the homologies of the metasternal structures. The rather similar topographical conditions in the Pseudococcidae, in which the metasternal apophyses are present (Giliomee, 1961), indicate that the large metasternal plate of the Coccidae represents a sternellum and that the small sclerites found in Eriopeltis sp. corresponds to a basisternum. This conclusion is supported by the position of the sternal apophyses and the large sternellum in Margarodes (Theron, 1958), and the general structure of the meso- and metasterna in Aphis (Weber, 1928). Ghauri's (1962) interpretation of corresponding sclerites in the Diaspididae as a metabasi- sternum and mesosternellum respectively, consequently appears to be incorrect.
Habib (1956) recognized the inverted nature of the metanotum ; Ezzat (1956) and Borchsenius (1957) illustrated the pleural sclerotization but do not discuss it in any detail ; Theron (1958) overlooked the postnotal and sternal sclerites in P. pomer- anicum.
Dermal structures. The fleshy and hair-like setae are arranged in the following groups:
(i) Metatergal setae (mts), occurring laterodorsally, anterior to the postnotal sclerite. They usually consist of a single hair-like seta on each side, but in the INGLISIA and COCCUS groups (Text-figs. 29-43) up to 10 fleshy setae may also be present in this region. In Ceroplastes spp. only fleshy setae are present and in E. tiliae, P. piceae, E. pela and Genus A no metatergal setae were observed. The setae of this region are sometimes difficult to observe because of the heavy sclerotiza- tion of the invaginated structures which lie directly underneath.
(u) Dorsospiracular setae (dss), which are situated pleurally, dorsal to the meta- spiracle and in line with the pleural setae of the abdomen. When they are numerous they are sometimes difficult to separate from the metatergal setae. They are only present in the JNGLISIA and COCCUS groups. The number of fleshy setae varies from 1-8 (average 3:6) in J. theobromae to 10-23 (average 15) in P. acericola ; hair- like setae are rarely present and never total more than 3.
30 MORPHOLOGY AND TAXONOMY OF ADULT MALES
(ii) Antemetaspiracular setae (ams), which are found immediately anterior to the metaspiracle and consist entirely of fleshy setae (up to 12). They are present only in the JNGLISIA and COCCUS groups.
(iv) Postmetaspiracular setae (eps,s), which occur in the pleural region posterior to the metaspiracle, partly on the metepisternum, and may extend ventrally towards the metasternum without reaching it. The fleshy setae are generally numerous in the COCCUS group (up to 35 in P. cornz), but few in the other groups and absent in some species of the EULECANIUM group (e.g. FE. téliae). The hair-like setae occur irregularly and in small numbers (0-3), but in E. pela they are always present (4-8, average 5°6).
(v) Anterior metasternal setae (amss), situated in the membranous area between the meso- and metasternal plates. Fleshy setae are present in the ERJOPELTIS, INGLISIA and COCCUS groups (Text-figs. 24, 43) and in S. prunastri (Text-fig. 22) of the EULECANIUM group. Their number varies from 8-17 (average 12) in E. ?festucae to 74-94 (average 86) in P. corni. A few hair-like setae are present in some individuals of most species.
(vi) Posterior metasternal setae (pmss), occurring on the metasternal plate, or in the area normally occupied by this sclerite. Fleshy setae are found in the ERJO- PELTIS, INGLISIA and COCCUS groups (Text-figs. 24-43) where their number varies from 5-16 (average 8) in J. theobromae to 31-60 (average 42) in P. corni. In S. prunastri (EULECANIUM group) up to 3 fleshy setae may be present in some individuals, but are absent in others. A few hair-like setae (up to 3) may be present in some individuals of a number of species.
In Ctenochiton sp. 3-11 (average 6-3) circular pores, somewhat reminiscent of vacant hair sockets, occur near the metatergal seta on each side.
Wings and Halteres
The fore wings are large, with a narrow basal part and a broadly rounded apex. They may be relatively short and broad, i.e. the length 1-g-2-3 times the width (e.g. the COCCUS group), or long and narrow, i.e. the length 2-8-3-3 times the width (e.g. the ERIOPELTIS group, Text-figs. 24, 26, 27). Posteriorly, near the base, a small pouch or alar lobe (al) (Stickney, 1934b) is formed by the dilation of the margin of the wing. It is ventrally invaginated and provides a receptacle for the hooked distal ends of the apical setae of the haltere. When the haltere is lacking the alar lobe is absent.
The wing is semitransparent, although in some species, e.g. P. bituberculatum, the area between the anterior margin and the first wing vein has a purplish tinge. The surface of the wing is covered with minute hairs (microtrichia), with those on the margins somewhat longer than elsewhere. Only two distinct veins are present. Patch (1909) suggested that corresponding veins in the Pseudococcidae represent the vadius (rad) and media (med) ; these veins were also called radius and “‘ medius + subcosta’”’ by Diirr (1954), but the last mentioned designation is obviously in- correct. The radius runs parallel to the anterior margin of the wing, the media deflects towards the hind margin. As in the Pseudococcidae (Patch, 1909) the two
OF THE FAMILY COCCIDAE 31
veins are not visibly connected. At the base of the wing, near the anterior margin, an elongate sclerite forms the costal complex of veins (Text-fig. 1 ; 18, N ; cex). The proximal part of the sclerite is pointed and articulates with the anterior notal wing process. Near the base an anterior extension is found which curves ventrally (see Text-fig. 18, N) to articulate with the pleural wing process. A small number of hair-like alar setae (as) are found in the anterior part of the base of the wing in E. tiliae, N. abietis, R. spiraeae and Ctenochiton sp. (Text-figs. 2, 4, 8, 16). Their number does not exceed 3 ; they are sometimes absent on one of the two wings.
The hind wings are either absent (P. myrtilli, S. prunasinn, the ERIOPELTIS, INGLISIA and COCCUS groups ; Text-figs. 12, 22-43) or reduced to halteres (h) (most of the EULECANIUM group). The anterior half of the haltere is weakly sclerotized and near the base the anterior margin is strengthened by a ridge, which resembles a wing vein. At the apex each haltere carries at least one long seta, but in E. tiliae, N. abietis, R. spivaeae, P. piceae and E. pela (Text-figs. 2-9) three or four may be present. These setae are curved apically and hook on to the alar lobe. It is worthy of note that the halteres of Margarodes (see Theron, 1958) resemble the hind wings of certain Aphididae, e.g. Anomalaphis compert and Miucroparsus variabilis (see Baker, 1920) to a considerable degree, presumably through con- vergence. The halteres were called ‘“‘ pseudohalteres’’’ by Kawecki (1958b), and recently (1964) he suggested the term “‘ hamulohalterae ’’ for these structures.
Legs
The three pairs of legs are very similar, long and slender, and composed of the usual segments, with a one-segmented tarsus and a single claw. The fore legs are usually the shortest and the hind legs the longest, but conditions vary and there are species in which the hind or the middle legs are the shortest. All the segments of the leg are well sclerotized and all except the claw are covered with numerous fleshy (fs) and hair-like (hs) setae, although the fleshy ones are sometimes absent on the tarsus (Genus A and E. P?festucae). They are not arranged into groups, but scattered over the whole surface of the segment. These setae are similar to those occurring elsewhere on the body. Conforming with the conditions on the antennae, the fleshy setae of FE. pela are very long, from 3 to 5 times as long as the width of the tibia, and in Genus A they are very short, about ? as long as the width of the tibia ; in the other species they are slightly longer than the width of the tibia. In E. tiliae the fleshy setae are very thin and it is difficult to distinguish them with certainty from the hair-like setae. The hair-like setae are very similar in all the species studied and are usually a little longer than the width of the tibia. Dis- tinctly different setae occur on the inner margin of the anterior coxae of some species. They are large, rigid, sometimes capitate, with the setal membrane sur- rounded by a distinct basal ring ; they are here called coxal bristles (cb), and are probably sensory in nature. A pair of tarsal digitules (tdgt), i.e. long, capitate setae, is present near the dorsal apex of each tarsus and two smaller wngual digitules (udgt) occur on each claw.
32 MORPHOLOGY AND TAXONOMY OF ADULT MALES
The coxa (cx) is short and broad. Its base is strengthened by a well sclerotized basal ridge which articulates dorsally with the pleural ridge by means of a short basal process. The apical margin is also ridge-like and bears an anterior and a posterior process, which articulate with corresponding processes on the trochanter. The hair-like setae on the coxae vary considerably in length, those near the basal process being very short and those near the apex being longer. The longest seta on the inner terminal part, called the apical seta (ase) may be short, i.e. length about half that of the trochanter (e.g. E. tuhiae, F. viburni, Pulvinaria spp. ; Text-figs. 2, 14, 35 & 37) or long, i.e. as long as the trochanter (ERIOPELTIS group ; Text-figs. 24, 26, 27). Coxal bristles were found in some species of the EULECANIUM group (e.g. E. tiliae, N. abtetis ; Text-figs. 2, 3 ; M ; cb) and all the species of the COCCUS group (Text-figs. 31, 33735, 37, 38,.42 } MM; cb), except C. bere They appear to be capitate in all the species of the COCCUS group except P. pomeranicum and pointed in most species of the EULECANIUM group ; in some specimens of E. tiliae both capitate and pointed bristles may occur. The number of coxal bristles varies from 1-2 (average 1-4) in P. bituberculatum to 5-8 (average 6-2) in E. tilvae.
The trochanter (tr) is elongate, narrow basally and broad distally. The strong basal ridge bears an anterior and posterior articular process and continues for some distance along the outer margin. The trochanter is separated from the femur by a narrow membrane. A minute hair-like seta occurs both anteriorly and posteriorly in the membrane near the basal ridge and a small rigid seta is always present on the outer margin. These setae appear to be proprioceptors. Ventrally near the apex there is usually one, but in some species (e.g. E. tiliae, P. piceae, R. spiraeae) two long hair-like setae. The longest seta, the apical seta (ase), may be comparatively short, 1.e. less than 1} times as long as the width of the trochanter (Genus A, C. hesperidum ; Text-figs. 20, 31) or long, i.e. more than 3 times as long as the width of the trochanter (E. ttliae, N. abietis, Eriopeltis spp., Text-figs. 2, 4, 24 & 26). A ring of oval campaniform sensilla are found in the basal half of the trochanter. They are usually 6 in number, but in E£. pela up to 8 may be present.
The femur (fm) varies in shape from being long and narrow, i.e. 6 times longer than wide (C. hesperidum, Genus 6b ; Text-figs. 31, 33) to short and broad, 1.e. 33 times longer than wide (Genus A, S. prunastn ; Text-figs. 20, 22). The distal ridge is well developed and bears an anterior and posterior process which articulate with corresponding processes on the tibia. All the setae are of the ordinary fleshy and hair-like types.
The tzbia (tib) is long and slender. The width/length ratio varies from about 1: 1rinS. prunastn and L. luzulae (Text-figs. 22, 27) to about I : 21 in C. hesperidum (Text-fig. 31). Basally it articulates with the femur by means of two processes and distally it is connected to the tarsus by means of a narrow, articular membrane, with- out a sclerotized joint being formed. The relative numbers of fleshy and hair-like setae vary ; in the COCCUS and INGLISIA groups and some species of the EULECANIUM group the fleshy setae are more numerous and in the ERIOPELTIS group and some species of the EULECANIUM group the hair-like setae are more
OF THE FAMILY COCCIDAE 33
numerous. On the inner margin, near the apex, an apical spur (tibs) is present in all the species studied. In some species, e.g. E. pela, some of the hair-like setae near the apex also have a spur-like appearance. In J. theobromae the apical spur on the front leg is short, about half as long as on the other tibiae.
The tarsus (tar) is elongate; the length varies from being about 3 times longer than wide (P. piceae, Text-fig. 6) to about g times that (Genus B, Text-fig. 33). The tarsus is broadest near the base or in the middle and tapers distally. Distally it articulates with the claw by means of a small dorsal process. As is the case with the tibia, there are more fleshy than hair-like setae in some species and more hair- like than fleshy setae in others. In Genus A the fleshy setae are completely absent on the middle, hind and sometimes front tarsi, while they are sometimes absent on the tarsi of FE. ?festucae. Two long, subequal, capitate tarsal digitules (tdgt) are found dorsally near the apex.
The claw (cl) is well developed, curved, pointed, with a small denticle ventrally near the apex. In some species, e.g. P. myrtilli and C. hesperidum the denticle is minute. Each side of the base of the claw bears a capitate wngual digitule (udgt), which is usually about as long as the claw. The claws on all three legs are sub- equal and they show little variation within the family.
Most of the earlier workers give brief descriptions of the leg. The tarsal and ungual digitules were already observed by Putnam (1879).
The Abdomen
The abdomen is elongated, more or less parallel sided, with the posterior end tapering and carrying the narrow and sclerotized genital segment. In cross section it is strongly convex ventrally and only moderately so dorsally. In most of the species the pregenital segments are almost completely membranous. The seg- mentation is not very distinct, but it is indicated by shallow intersegmental grooves, and the segmental arrangement of the setae and transverse bands of minute dermal denticulations. These denticulations, which are also present in male Diaspididae (Ghauri, 1962), Pseudococcidae and some female Coccoidea and Aphididae, occur on the dorsal and ventral surfaces of the median part of each segment. The abdomen is composed of eight pregenital segments and the oth or genital segment; this was recognized by Putnam (1879), Silvestri (1919a, 1g1g9b, 1920), Sule (1932), Pesson (1941), Borchsenius (1957), Theron (1958) and Bustshik & Saakjan-Baranova (1962).
Pregemital Segments
The 1st segment is developed dorsally and pleurally, but not ventrally ; the other segments are complete. The sclerotization of the abdomen varies considerably within the family. Where it is most fully developed, as in E. tiliae, N. abietis, P. piceae, P. myrtilli and L. luzulae (Text-figs. 2, 4, 6, 12, 27) tergal and sternal plates are found on all the abdominal segments. In other species (P. bituberculatum, F. viburm, Ctenochiton sp., Eriopeltis spp., I. theobromae and P. ?betulae ; Text-figs. 10, 14, 16, 24 & 26, 29, 35) sternites are present on all the segments, but tergites are absent on one or more of the middle segments and in the remaining species both tergites and sternites are absent on these segments.
34 MORPHOLOGY AND TAXONOMY OF ADULT MALES
The tergites (at) of segments I-III are situated in the intersegmental or ante- costal region and they usually consist of a small transverse sclerite on each side. Those in front of segment I are enlarged and can perhaps best be regarded as the postnotum of the metathorax ; in the other segments they can be regarded as belonging to the segments posterior to them (similar tergites in Planococcus (Theron, 1958) and Pseudococcus (Giliomee, 1961) are probably also intersegmental and do not belong to the preceding segments as has been indicated). In some of the species, e.g. Genus B, Pulvinaria spp. (Text-figs. 33, 35 & 37) the tergites of segments II-III consist of a small sclerite on each side and a separate additional median sclerite, but in Genus A and sometimes in P. myrtilli there is a continuous transverse median sclerite. In segments IV—VIII the tergal plates are large, transverse and situated in the middle of each segment.
The sternites (as) also consist of large, transverse, segmental plates. In the more anterior and posterior segments they are usually complete, but in the intermediate segments they are either interrupted or completely absent. In some of the inter- mediate segments the membrane near the segmental boundary is bulging and irregularly folded.
Pleural sclerotization is only found on the caudal extensions of segment VII of the ERIOPELTIS and COCCUS groups (Text-figs. 24-43), on the caudal extensions of VIII in the COCCUS group, and in the pleural region of the INGLISIA group (Text-fig. 30). In J. theobromae a continuous band of sclerotization extends along the pleural region of segments IV—VII, with a small sclerotized area situated some- what more ventrally on each of segments V-VI. This bears some resemblance to the condition in the winged female of Aphis fabae, where the lateral plates occur on abdominal segments II and VI (Weber, 1928) and, according to Weber, are morpho- logically part of the terga. This resemblance is probably merely superficial and devoid of any phylogenetic significance as no lateral sclerotization is present in any of the other Coccoidea studied so far.
In the COCCUS group, segment VII laterally bears a very prominent, tapering, caudal extension (ce). In the other groups this extension is small and broadly rounded or somewhat pointed. In the COCCUS and ERIOPELTIS groups they are weakly sclerotized lateroventrally. Theron (1958) incorrectly describes them as belonging to abdominal segment VIII and Sule (1932) correctly illustrates the segmental position in one of his figures (fig. 23), but incorrectly in another (fig. 25). The caudal extension (ce) of segment VIII is also shaped in a variety of ways. It may form a small, simple lobe (the ERIOPELTIS group, the INGLISIA group and most of the EULECANIUM group), a papilla-shaped lobe (Parthenolecanium spp. ; Text-figs. 38, 39 ; Q), a large, straight, cylindrical lobe (S. prunastri, Text-fig. 22 ; C. hesperidum ; Text-fig. 31, Q) or a somewhat geniculate lobe (Genus B ; Text- fig. 33, Q), a mammillate lobe (Pulvinaria spp. ; Text-figs. 35, 37 ; Q) or a prom- inent and semicircular lobe (Ceroplastes spp. ; Text-figs. 41, 43; Q). In the COCCUS group the distal part of the lobe is weakly sclerotized and it bears a structure which is usually membranous but weakly reticulated; sometimes (Cero-
OF THE FAMILY COCCIDAE 35
plastes spp.) it is weakly sclerotized in the middle. The structure, which can conven- iently be called a cicatrix (c), varies in shape and relative size ; in Ceroplastes spp. it is large, circular and occurs dorsally ; in C. hesperidum and Genus B it is also large but occupies the posterior surface of the lobe and in Parthenolecanium spp. it is small and occurs apically. The caudal extensions resemble the fleshy tassels of some Monophlebidae (Morrison, 1928), presumably through convergence. Newstead (1916), in describing the male of C. hesperidum, referred to the caudal extensions of the 7th segment as “ long slender hairy tubercles ’’ and those of the 8th as “ pro- truding gland-like processes ”’
Dermal structures. Both fleshy and hair-like setae are present on the abdomen. They are segmentally arranged and occur in distinct groups on the dorsal, pleural and ventral surfaces and are referred to as abdominal dorsal (ads), abdominal pleural (dps and vps), and abdominal ventral (avs) setae.
The dorsal setae (ads) normally consist of 2 hair-like setae (one on each side) on all but segments II and III, but in P. bituberculatum and Genus B (Text-figs. 10, 33) they are regularly present on allsegments. In addition to these a small but variable number of fleshy dorsal setae occur in the ERIOPELTIS and COCCUS groups (Text-figs. 24-43).
The pleural setae can be subdivided into a dorsopleural group (dps) and a ventro- pleural group (vps). These two groups are not in line with each other, the former being situated nearer to the posterior margin of the segment. When numerous the two groups coalesce to a certain degree ; on the 7th segment the two groups are not differentiated. The dorsopleural group consists of both fleshy and hair-like setae, occurring in different proportions and numbers in different species. In some species (most of the EULECANIUM group, Text-figs. 2-21) all dorsopleural setae are hair-like ; except for the COCCUS group, there are no fleshy setae on segments I-III, and frequently the anterior segments have no dorsopleural setae at all. The ventropleural setae usually consist of a single hair-like seta and none or but a few fleshy setae ; they are never found on I and rarely on segments II and III.
The ventral setae are arranged in a median group in the middle of each segment. They often consist of both fleshy and hair-like setae. The hair-like setae are always present, arranged on each side of the body into two longitudinal series, one median and one lateral, each series usually with one seta per segment. Both series are usually present on segments V—VII, but on segments II-IV either median or lateral setae are frequently absent, and on segment II hair-like ventral setae are often absent altogether (e.g. L. luzulae, C. hesperidum, Ceroplastes spp.). In most species (S. prunasini, ERIOPELTIS, INGLISIA and COCCUS groups ; Text-figs. 22-43) fleshy ventral setae are also present. If present, they are usually numerous, except on segment VIII, but a comparatively large number (more than 5) on this segment is characteristic of the JNGLISIA group. In S. prunastri fleshy setae occur only on segments II and sometimes III.
In addition to the above-mentioned setae, a number of setae are present on the posterior margin of segment VIII. Lateral to the glandular pouch this segment always carries 3 hair-like setae ; occasionally a fleshy seta may also be present,
36 MORPHOLOGY AND TAXONOMY OF ADULT MALES
but a comparatively large number (2~7) is characteristic of Ceroplastes spp. (Text-figs. 41, 43). In many species setae are present in the region anterior to the anus where, in some species, the small tergal plate of segment IX (mentioned later) is found ; these setae are called ante-anal setae (aas). In most of the species they consist of two long pointed hair-like setae, but in Parthenolecanium spp. one or both are some- times bifurcate. In some genera (Eviopeltis, Pulvinaria and Ceroplastes ; Text- figs. 24 & 26, 35 & 37, 41 & 43) a number of fleshy setae are also present in this region, and in most species one or two small hair-like setae may occur in some individuals.
A group of small circular pores, reminiscent of vacant hair sockets, is found dorsally on each side of abdominal segment I of Ctenochiton sp. (Text-fig. 16) and a small number of these pores are also found in the ante-anal region in Ctenochiton sp., L. luzulae and Genus B (Text-figs. 16, 27, 33). On each side of the base of the penial sheath there is a funnel-shaped pouch (gp), which contains 2 long setae arising from its bottom. From about halfway up to the rim, the pouch is lined with numerous quadrilocular, but also with a few tri- and quinquelocular pores. According to Sulc (1931) the basal half of the pouch is lined with tubular pores. The pores secrete a waxy substance which slides along the setae and constitutes the conspicuous long waxy filament of the living male. The structure of the filament and pores was studied in detail by Sulc. In Ctenochiton sp., E. pela, Genus A and, as reported by Durr (1954) in Lecanium pumilum Brain (= Saissetia oleae (Bern.) according to De Lotto (1959)) the setae are knobbed apically. The setae vary in length from short, i.e. the length of the protruding part only twice that of the part concealed within the pouch (e.g. E. tiliae, Text-fig. 2) to long, where the length of the protruding part is 4-6 times longer than the concealed section (e.g. C. hesperidum, Text-fig. 31). In L. luzulae the pouch is absent and replaced by a shallow depression with one long seta, but no pores at all. From available information it is known that the pouch is also reduced in Vinsonza stellifera (Newstead, 1903) and Ceroplastes jyaponicus (Borchsenius, 1957). The glandular pouch corresponds to what is called the “glandular plate” (Pflugfelder, 1939 ; Giliomee, 1961) in the Pseudoccidae.
Gemtal Segment and External Genitalia
The genital segment has become elongated to form a long tubular style which tapers posteriorly. The anus (an) is situated dorsally in the membrane at the basal part of the segment. The penzal sheath (ps), which is composed of sternum IX (Theron, 1958), is well sclerotized laterally and membranous ventromedially. The lateral sclerotizations fuse dorsally with each other at some distance posterior to the anus. In some species (e.g. most of the COCCUS group) they are also narrowly joined anterior to the anus, a condition which obtains in the Diaspididae (Theron, 1958 ; Ghauri, 1962). The apex is sometimes produced into a small membranous extension, which is best developed and finger-like in Ceroplastes spp. (Text-figs. 41, 43; R). The ventral membrane widens anteriorly to form a triangular area which, for descriptive purposes, can be called the basal membranous area (bma). Posterior to this area a narrow ridge is formed on the median line, which appears
OF THE FAMILY COCCIDAE 37
to be homologous to the basal rod (bra) found in other Coccoidea (see Theron, 1958, 1962), but was overlooked by Theron (1958) in P. pomeranicum. According to Theron the basal rod may incorporate the basal plate. Parameres are absent.
Posteriorly the basal rod is connected to the base of the aedeagus (aed), which is accommodated in a slit in the ventral wall of the penial sheath. The aedeagus consists of a straight tube, which does not narrow appreciably towards the apex. It ends bluntly before reaching the apex of the penial sheath. The ductus ejaculatorius can be seen to run along the ventral wall of the penial sheath, but the position of the gonopore is impossible to observe in mounted specimens. The aedeagus appears to contain an eversible endophallus, as is indicated by occasional specimens in which the everted condition has been observed. The genitalia with everted aedeagus, but considered to represent the normal condition, were illustrated by Sulc (1932). The relative length of the penial sheath, aedeagus and basal rod show considerable variation within the family. Silvestri (1g19a, 1g19b, 1920) and Jancke (1955) correctly illustrated and interpreted the aedeagus, while the illus- trations given by Leonardi (1920) and Theron (1958) are inaccurate in some details ; Putnam (1879), Diirr (1954) and Husseiny & Madsen (1962) called the entire gth segment either a penis or aedeagus.
In well stained specimens of some species (e.g. the COCCUS group ; Text-figs. 31-43) a small gth tergite (atg) can be seen in the membrane anterior to the anus, but the roth and 11th tergites described by Sulc (1932) were not observed.
Dermal structures. A number of small setae (gts), which are possibly tactile sensilla, are scattered over the genital segment. Distally the setae become con- siderably smaller and at the apex only small, circular discs can be discerned ; the latter may be campaniform sensilla.
DESCRIPTION OF THE SPECIES
In the descriptions of the individual species considerable detail has been included and they may well appear to be unduly long and repetitive. However, detailed descriptions are considered necessary since the taxonomic significance of the characters has, as yet, not been properly evaluated.
For the sake of brevity the usual telegraphic style of describing species has been adopted and the following abbreviations are used : h.s. = hair-like seta(e), f.s. = fleshy seta(e) ; the figures in brackets signify averages.
THE EULECANIUM Group EULECANIUM
Eulecanium tiliae (Linnaeus) (Text-figs. 2 and 3)
Living specimens reddish, with sclerotized areas dark brown and the appendages light yellow, wings with a purplish tinge between anterior margin and first wing vein ; very long and moderately robust, with comparatively short antennae and legs which carry many setae.
38 MORPHOLOGY AND TAXONOMY OF ADULT MALES
When mounted, total body length 2440-2700 (average 2569) u ; width at mesothorax 570-660 (average 601) uw. Wing expanse 3950-4550 (average 4296) uw.
Head subconical in dorsal view ; in lateral view obliquely dorsoventrally elongated, with the anterodorsal bulge not pronounced ; length from apex to pronotal ridge 274-319 (average 303) uw, width across genae 296-338 (average 315) up. Median crest sclerotized and distinctly polygonally reticulated ; with 3-7 (average 5-3) hair-like dorsal head setae, arranged in two groups : one, with 2-4 (average 3-3) short setae, posterior to the level of the dorsal eyes, and the other with o—4 (average 2) longer setae, anterior to the eyes. Midcranial vidge dorsally represented by a short, weak ridge anterior to the level of the eyes ; ventrally narrow but well defined, reaching ocular sclerite posteriorly, with surrounding area showing weak, polygonal reticulation posteriorly. Genae large, sclerotized, weakly polygonally reticulated, without setae. Eyes: five pairs ; dorsal and ventral pairs large, subequal, lateral pairs smaller, sub- equal ; corneae of dorsal eyes 34-42 (average 38) u in diameter and 2-2—2-9 (average 2-5) times as much apart ; those of the ventral eyes 32—42 (average 35) u in diameter and 1-1-1-6 (average 1-3) times as much apart. Ocellus small. Ocular sclevite well sclerotized except between the ventral eyes where the cuticle is produced into a keel ; polygonally reticulated throughout. Preoculay ridge extending only a short distance below articular process. Postocular vidge very weak dorsally, sometimes missing posterior to ocellus ; well developed lateroventrally, but weak posteromedially ; below the ocellus the ridge splits up, with the anterior branch partly surrounding ocellus. Jntevoculay vidge absent. Dorsal ocular setae absent. Ventral head setae consisting of 1-5 (average 3:2) h.s., situated anterior to the ocular sclerite on each side of the midcranial ridge. Pveoval ridge present. Tendon-like apodeme long. Cvranial apophysis short ; apex bifurcate or occasionally truncate, not quite reaching level of anterior margin of ventral eyes. Mouth opening irregular. Anterior tentorial pits absent.
Antennae to-segmented ; filiform ; 895-1075 (average 988) uw long, i.e. shorter than half body length (ratio 1 : 2:46-2:92, average 2-62) ; shorter than posterior leg (ratio I : 0-73—-0-87, average o-80) and longer than penial sheath (ratio 1 : 1-:16-1-50, average 1-31). Scape 57-68 (average 61) uw long and 57-67 (average 62) uw wide, with 3 h.s. Pedicel with distinct, polygonal, dorsal reticulation ; 49-57 (average 54) u long and 49-57 (average 53) uw wide, with o-3 (average 1) f.s., 2-4 (average 2-9) h.s. and a sensillum placodeum. Segment III club- shaped, 1-8-2:2 (average 2) times longer than wide (84-106, average 90 uw long and 42-49, average 45 uw wide) ; with 4-12 (average 8-5) h.s. and 4-12 (average 5-8) f.s., the latter of medium length, 0-7-1-o0 (average o-9) times as long as width of segment ; with 4-9 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths of these segments (in w) 114-171 (average 152), 137-160 (average 149), 106-148 (average 131), 95-122 (average 112), 80-95 (average 90) and 68-91 (average 78) respectively, widths varying from 30 to 42 yp, with distal segments wider than proximal ones ; with 18-53 (average 33), 33-55 (average 44), 26-46 (average 37), 28-43 (average 36), 22-32 (average 27), 18-34 (average 25) f.s. respectively, but no h.s. ; antennal bristles on segments VIII-IX thicker than f.s. Segment X : terminal part not constricted ; 57-84 (average 72) wu long and 30-38 (average 33) uw wide ; carrying 8-14 (average 12) f.s., 3 capitate subapical setae and 5 antennal bristles of which the 3 long ones are about half as long as the segment and the 2 shorter ones about as long as the f.s., though thicker ; with 2 sensilla basiconica ventrally, one near apex and the other more proximal.
Thorax 806-901 (average 864) u long.
Prothovax. Pyronotal ridge strong, but medially interrupted by weak sclerotization. Lateral pronotal sclervites large, without setae. Medial pronotal setae absent. Post-tergites medium- sized, without striations and without setae. Pleural stvuctures typical of the family. Steynum with strong transverse ridge, interrupted median ridge and a triangular sclerite. Anteprosternal setae absent ; prosternal setae o—3 (average 0-8) h:s.
Mesothovax. Mesoprephvagma with shallow emargination. Prescutum about twice as wide as long (average 268 and 138 uw respectively) ; anterior margin slightly curved ; laterally bounded by the prescutal ridges and posteriorly by the prescutal suture ; sometimes with very weak polygonal reticulation. Scutum. Median membranous area transverse ; 80-106 (average
39
THE FAMILY ‘COCCIDAE
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40 MORPHOLOGY AND TAXONOMY OF ADULT MALES
95) w long and 2-06—3-00 (average 2:36) times as wide (width 203-251, average 223 uw) ; without setae. Scutellum 93-114 (average 107) uw long and 217-266 (average 236) uw wide, the ratio being I : 2-0-2:5 (average 2-2) ; not tubular and without setae. Postnotum with anterior margin usually irregular and partly overlapped by metathoracic fold ; postnotal apophysis and postalave well developed, the latter densely reticulated distally. Mesopostphvagma with moderately deep emargination. Mesopleuron. Mesopleural ridge strong, but interrupted above coxal articulation ; pleural apophysis and pleural wing process well developed, the latter connected with episternum by a narrow basalave. Subalave small. Episterynwm not reticulated; subepisteynal ridge well developed, becoming appreciably broader ventrally, but below mem- branous cleft indistinct and only marked by a band of dark sclerotization. Epimeron small. Lateropleurite partly bounded anteriorly by an extension from marginal ridge. Basisternum large, about 383 uw wide and 287 y long, i.e. 2:70-3:62 (average 3-04) times longer than mem- branous area of scutum ; with strong median ridge and bounded by strong marginal and precoxal vidges. Furca well developed. Mesothoracic spivacle with well developed peritreme ; postmesospivaculay setae absent. Tegula comparatively large ; membranous bulge with a small weak sclerite posteriorly, with 6-10 (average 7:7) h.s. and sometimes showing wavy striations. Third axillary wing sclevite with a pronounced ventral projection at its base. Additional sclerite well defined. Antemetaspivacular setae absent.
Metathorvax. Metanotum with thickening of posterior margin sometimes desclerotized medially ; suspensorial sclerites small, irregular ; a small additional sclerite always present anterior to postnotum. Postnotum consisting of a transverse sclerite on each side. Metatergal setae absent. Pleural ridge well developed, though interrupted near the middle ; with a small wing process. Episternum with anterior margin ridge-like in parts ; epimevon produced posteriorly. Metathoracic spivacle similar to mesothoracic one. Dorsospivacular setae absent. Postmetaspivaculay setae: o-2 (average 0-9) h.s. Metasternal plate weak and irregular. Anterior metasternal setae consisting of o—2 (average 1-4) medial h.s. and occasionally with one or two lateral to the posterior part of the basisternum ; postevioy metasternal setae usually 2 (range 1-3) h.s. medially.
Wings hyaline ; of medium length (1750-2000, average 1879 uw) but comparatively broad (width 780-870, average 840 uw), ratio width to length being 1 : 2:13-2:35 (average 2-24) ; alar lobe present ; alay setae : 1-3 (average 1-7) h.s. on each wing. Halteves well developed, 163-201 (average 178) uw long and 42-61 (average 53) uw wide, each with 2-4 (average 2-6) apically hooked setae which are about 70 yu long.
Legs short and slender, with fore pair shortest and hind pair longest ; ratio length of hind leg to body length is i : 2:07-2:15 (average 2-10). Length of segments (in y) :
Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 91-103 118-148 274-346 35355437 93-103 29-34 973-1094 (95) (128) (301) (391) (98) (30) (1043)
Il 99-118 120-133 251-308 433-479 114-133 29-34 1058-1203 (110) (129) (285) (45°) (124) (32) (1130)
Ill 106-141 125-143 281-315 450-532 125-148 30-34 1132-1303 (124) (134) (295) (499) (134) (32) (1217)
Fic. 3. Eulecanium tiliae (L.), lateral view.
OF THE FAMILY COCCIDAE 41
F.s. on the legs very slender, making it impossible to distinguish with accuracy between f.s. and hs.
Coxae each with 24-35 setae ; fore coxa with 5-8 (average 6-2) coxal bristles, of which some are capitate ; apical seta about half as long as trochanter. Tvochanters 34-44 u wide ; with 6 oval sensilla ; with 16—25 setae, including 2 or 3 minute setae near basal ridge, one small seta on the outer margin and 2 long setae, of which the longest (apical) on the fore trochanter is 2°9-3°4 (average 3-1) times as long as width of trochanter. Femova of medium width (53-67 uw), ratio width to length of hind femur being 1 : 4:5—5-1 (average 4-9) ; each with 37-65 setae. Tibiae 29-38 pw wide, ratio width to length of hind tibia being 1 : 14-1~-16-3 (average 15-1) ; each with 74-121 setae which are about as long as width of tibia ; apical spur of about the same size on all tibiae. Yavsi 28-34 » wide, hind tarsus 4-1—4-9 (average 4-4) times longer than wide ; each with 18-37 setae ; tarsal digitules subequal, longer than claw. Claws of medium length, about as long as width of tarsus ; slightly curved, with small denticle near tip ; anterior ungual digitule with larger apical knob than posterior one, digitules longer than claw.
Abdomen 670-810 (average 713) uw long and 450-600 (average 521) wp wide.
Segments I-VII : tergites and sternites present on all segments ; tergites on segments II and Ill represented by a small sclerite on each side on the anterior margin, and on the IV—VII by a transverse plate ; sternites represented by a weak plate on the anterior and posterior segments and a small sclerite on each side on the intermediate segments. Caudal extension of segment VII small, rounded, not sclerotized. Dorsal setae: f{.s. absent ; h.s. absent on segment I, sometimes one or two present on each of the segments II-VII. Pleural setae consisting of h.s. only, which include dorsopleural setae : o—3 (average 1-3), 2-8 (average 3-8), 1-6 (average 3:5) and 2-6 (average 3-8) on segments III—VI respectively, and ventyropleural setae : occasionally one on each of III and IV, and usually one on each of segments V and VI. Segment VII with 6-9 (average 7°4) h.s. Ventral setae : h.s. only, usually 2 medially on each of II and III, and 4 on each of segments [V—VII.
Segment VIII with transverse tergite and sternite ; caudal extension forming a small, simple lobe ; glandular pouch with 2 long setae which are sometimes capitate and whose protruding part is about 1} times as long as the section within the pouch. No IX tergite observed. A nte- anal setae : 2longh.s. Posterior margin with 2—4 (average 2-6) h.s. on each side.
Genital segment. Penial sheath long, about ? total body length (ratio 1 : 3-3-3-7, average 3°4), 654-809 (average 756) uw long and 49-59 (average 55) uw wide ; lateral sclerotizations not joined anterior to anus ; length of basal vod 2-% that of aedeagus, the rod extending anteriorly from base of aedeagus for 3—} of the distance to apex of basal membranous area ; apex of sheath without membranous extension. The area from base of sheath to tip of aedeagus with 27-40 (average 35) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath. Aedeagus long (300-376, average 349 uv), penial sheath longer and basisternum shorter, the ratios being 1 : 1-9-2-4 (average 2-2) and 1 : 0:71-0:96 (average 0-82) respectively.
Material examined : 10 specimens, bred in the laboratory from material collected by myself on horse-chestnut (Aesculus hippocastanum L.) at the Imperial College Field Station, Silwood Park, Sunninghill, Berks. ; males emerged during May, 1962. Five specimens collected by J. Rehdéek in Bratislava, Czechoslovakia on I4.1v.53 (remounted from Swann’s mountant) agreed well with the above description.
NEMOLECANIUM Nemolecanium abietis Borchsenius (Text-figs. 4 and 5)
A long, slender species with comparatively short antennae and legs ; with numerous setae on the appendages, but few on the body itself. When mounted, total body length 1930-2270
MALES
MORPHOLOGY AND TAXONOMY OF ADULT
“MOIA [LIJUBA pue [eSIOp “SyoIOg szarqv umnimozajoma Ny
BEES |
NAGY
v OWT
OF THE FAMILY COCCIDAE 43
(average 2136) uw ; width at mesothorax 415-480 (average 445) u. Wing expanse 3870-4190 (average 4072) uw.
Head subconical in dorsal view ; in lateral view dorsoventrally elongated, with the antero- dorsal bulge not pronounced ; length from apex to pronotal ridge 251-293 (average 269) u, width across genae 251-289 (average 274) u. Median crest sclerotized, with a small area near posterior margin more heavily sclerotized ; weakly polygonally reticulated ; with 8-12 (average 10) hair-like dorsal head setae, of which 3—5 are situated posterior and 4—8 anterior to the level of the dorsal eyes. Midcranial ridge dorsally represented by a weak ridge which usually extends posteriorly to the posterior level of the eyes ; ventrally narrow but well defined, reaching ocular sclerite posteriorly, with surrounding area showing weak polygonal reticulation posteriorly. Genae large, sclerotized, not reticulated, without setae. Eyes: three pairs ; dorsal and ventral pairs subequal, lateral pair smaller ; corneae of dorsal eyes 22-30 (average 27) uw in diameter and 2-3-3-8 (average 2-9) times as much apart ; those of the ventral eyes 21-30 (average 26) uw in diameter and 1-1—2-2 (average 1-5) times as much apart. Ocellus small. Ocular sclerite well sclerotized except between the ventral eyes, where the cuticle is produced into a keel ; polygonally reticulated throughout. Pyreocular ridge long, ventrally extending half-way or more of the distance between the articular process and the midcranial ridge. Postoculay ridge very weak dorsally, well developed latero-ventrally, and tapering but well defined posteromedially ; below ocellus the ridge usually splits up, with the anterior branch partly surrounding ocellus. IJnteroculay ridge absent. Dorsal ocular setae absent ; ventral head setae: 2-4 h.s., situated immediately anterior to ocular sclerite, on each side of midcranial ridge. Preoval ridge present. Tendon-like apodeme long. Cranial apophysis medium-sized ; apex bifurcate, extending to around the level of posterior margin of ventral eyes. Mouth opening irregular. Anterior tentorial pits absent.
Antennae to-segmented, filiform ; 927-1087 (average 1025) u long, i.e. shorter than half body length (ratio 1 : 2:02-2:13, average 2-09), slightly longer than posterior leg (ratio I : 1-04— I-I4, average 1-07) and longer than penial sheath (ratio 1 : 1:55-1:71, average 1°64). Scape 53-68 (average 60) uw long and 46-55 (average 51) uw wide, with 3—4 (average 3-2) h.s., area of sclerotization reduced ventrally. Pedicel with distinct, polygonal, dorsal reticulation ; 55-68 (average 59) uw long and 42-51 (average 47) u wide ; with o—3 (average 2-1) f.s., 1-3 (average 2°5) h.s. anda sensillum placodeum. Segment I11 somewhat club-shaped, 2-5—3-0 (average 2-7) times longer than wide (80-95, average 90 uw long and 30-36, average 34 uw wide) ; with 2-6 (average 4) h.s. and 12-22 (average 15) f.s., the latter of medium length, 1-3—1-8 (average 1-5) times longer than width of segment ; with 1 or 2 usual sensilla basiconica. Segments 1V-IX cylindrical ; lengths of these segments (in uw) 80-95 (average 90), 103-144 (average 124), 133-171 (average 153), 137-178 (average 160), 103-133 (average 124), 87-106 (average 98) and 76-91 (average 84) respectively, all of about the same width, varying from 23 to 30 uw; with 20-34 (average 24), 27-36 (average 32), 27-44 (average 35), 25-41 (average 30), 21-31 (average 25) and 19-30 (average 23) f.s. respectively, but no h.s. ; antennal bristles on segments VIII-IX somewhat larger than f.s. Segment X : terminal part not constricted ; 68-82 (average 74) pv long and 23-29 (average 26) u wide ; carrying 8-13 (average 11) f.s., 3 capitate subapical setae and 5 antennal bristles of which the 3 long ones are about ¢ as long as the segment and the 2 shorter ones about as long as the f.s., though somewhat thicker ; with 2 sensilla basiconica ventrally, one near apex and the other more proximal.
44 MORPHOLOGY AND TAXONOMY OF ADULT MALES
Thovax : 619-752 (average 699) u long.
Prothovax. Pyronotal ridge strong, but medially interrupted by weak sclerotization. Lateral pronotal sclevites large, without setae. Medial pronotal setae absent. Post-tergites medium- sized, without wavy striations, and without setae. Pleural structures typical of the family. Sternum with strong transverse ridge, interrupted median ridge and narrow triangular sclerite. Anteprosternal and prosternal setae absent.
Mesothovax. Mesoprephragma with shallow emargination. Pvescutum less than twice as wide as long (average 197 and 113 uw respectively) ; anterior margin curved ; laterally bounded by the prescutal vidges and posteriorly by the prescutal suture ; medially with more heavy sclerotization which is ridge-like posteriorly ; not reticulated. Scutum. Median membranous area transverse, subrectangular ; 70-95 (average 86) uw long and 1-72—2-27 (average 2) times as wide (width 156-190, average 171 uw) ; without setae. Scutellum 72-91 (average 80) pu long and 148-190 (average 164) uw wide, the ratio being 1 : 2-0—2-3 (average 2-1) ; not tubular ; usually with 2 h.s. Postnotum with anterior margin irregular and partly overlapped by meta- thoracic fold ; postnotal apophysis and postalare well developed, the latter densely reticulated distally. Mesopostphragma with moderately deep emargination. Mesopleuron. Mesopleural vidge strong, but interrupted above coxal articulation ; pleural apophysis and pleural wing process well developed, the latter connected with episternum by a narrow basalare. Subalare small. Episteynum not reticulated ; subepisternal ridge well developed, becoming appreciably broader ventrally, but below membranous cleft indistinct and only marked by a band of dark sclerotization. Epimeron small. Lateropleurite partly bounded anteriorly by an extension from marginal ridge. Basisternum large, about 272 w wide and 230 yu long, 1.e. 2:46-3:03 (average 2-70) times longer than membranous area of scutum ; with strong median ridge and bounded by strong marginal and precoxal ridges ; with o-2 (average 1-4) h.s. on the median ridge. Furca well developed. Mesothovacic spivacle with well developed peritreme ; fost- mesospivaculay setae absent. Tegula small, membranous bulge with a small weak sclerite posteriorly and with 1-5 (average 2:5) h.s. Third axillary wing sclerite with a pronounced ventral projection at its base. Additional sclerite well defined. Antemetaspivacular setae absent.
Metathovax. Metanotum with thickening of posterior margin, sometimes desclerotized medially ; suspensorial sclerites small, spot-like ; a small additional sclerite usually present anterior to postnotum. Postnotum consisting of a transverse sclerite on each side. Metatergal setae : occasionally one h.s. on each side. Pleural ridge well developed, though interrupted near middle ; with a small wing process. Episternum with anterior margin ridge-like in parts ; epimeron produced posteriorly. Metathovacic spivacle similar to mesothoracic one. Dorso- spivacular setae absent. Postmetaspivaculay setae : o-2 (average 0-7) h.s. Metasternal plate weak and irregular. Anterior and posterioy metasternal setae usually consisting of 2 (range 1-3) medial h.s. each ; occasionally a h.s. occurs lateral to the posterior part of the basisternum.
Wings hyaline; long (1760-1910, average 1856 uw) and of medium width (700—780, average 753 wu), the ratio width to length being I : 2:42-2:51 (average 2-46) ; alar lobe present ; alar setae usually one (range o—2) h.s. on each wing. AHalteves well developed, 125-156 (average 142) uw long and 38-46 (average 43) p wide, each with 1-3 (average 1-8) apically hooked setae which are about 73 yu long.
Legs short and moderately slender, with middle pair shortest and hind pair longest ; ratio length of hind leg to body length is 1 : 2:20-2:22 (average 2:21). Length of segments (in yu) :
Leg Coxa ~ Trochanter Femur Tibia Tarsus Claw Total I 76-80 93-118 220-266 344-391 95-103 24-29 857-984 (79) (107) (245) (367) (98) (26) (922) II 76-87 95-110 194-251 308-365 IOI—I114 25-28 798-955 (83) (103) (222) (345) (108) (26) (886) IIl QI—-110 106-125 210-247 331-395 114-125 26-30 878-1034
(100) (114) (235) (371) (118) (27) (964)
OF THE FAMILY COCCIDAE 45
I’.s. slender and sometimes difficult to separate from h.s.
Coxae with 15-21 (average 19) f.s. on the fore, and 19-27 on the middle and hind coxa, and each with 11-19 h.s. ; fore coxa with 5-6 (average 5-6) coxal bristles, each with a small apical knob ; apical seta about half as long as trochanter. Tvochanters 26-38 uw wide, with 6 oval sensilla ; with 11-16 (average 12), 7-14 (average 10) and 5-12 (average 8) f.s. on the fore, middle and hind coxa respectively, and with 6-11 h.s., the latter including 2 minute setae near basal ridge, one small seta on the outer margin and 2 long setae of which the longest (apical), on the fore trochanter, is 3:0-3-9 (average 3-4) times as long as the width of the trochanter. Femora of medium width (42-57 wu), ratio width to length of hind femur being 1 : 4°5—5:2 (average 4°8) ; each with 21-35 f.s. and 12-18h.s. Tibiae 24-30 p wide, ratio width to length of hind tibia being 1 : 12-3~14-6 (average 13-5) ; each with 69-85 setae of which 20-34 are h:s. and 44-59 f.s., the latter about 14—1} times as long as width of tibia ; apical spur about the same size on all tibiae. Tarsi 23-30 uw wide, hind tarsus 3-9—5-0 (average 4:5) times longer than wide ; each with 12-18 f.s. and 11-19 h:s. ; tarsal digitules subequal, longer than claw. Claws of medium length, about as long as width of tarsus ; slightly curved, with small denticle near tip ; anterior wngual digitule with larger apical knob than posterior one, digitules about as long as Claw.
Abdomen 490-650 (average 579) uw long and 360—430 (average 390) w wide.
Segments I-VI1 : tergites and sternites present on all segments ; tergites on segments II, III and sometimes IV represented by a small sclerite on each side on anterior margin, and on IV-VII by a transverse plate ; sternites represented by a weak transverse plate on the anterior and posterior segments and a small sclerite on each side on the intermediate segments. Caudal extension of segment VII small, rounded, not sclerotized. Dorsal setae : h.s. only, segments I-III occasionally with one, and segments [V—VII usually with one seta on each side. Pleural setae absent on segments I and II and on III-VI represented by h.s. only, which usually include 2 (range 1-3) dorsopleural setae and 1 ventropleural seta on each segment. Segment VII with 4-6 (average 4:4) h.s. Ventral setae : h.s. only, usually one on each side on II and 4 on each of segments III-VII.
Segment VIII with a weak tergite and transverse sternite, caudal extension forming a small, simple lobe ; glandular pouch with 2 long, pointed setae, whose protruding part is about twice as long as section within pouch. No IX tergite observed. Ante-anal setae: 2 long, and occasionally one small h.s. Posterior margin with 3 h.s. on each side.
Genital segment. Penial sheath long, about 7 total body length (ratio 1 : 3-3-3+5, average 3-4), 581-695 (average 628) yp long and 46~—53 (average 49) u wide ; lateral sclerotizations not joined anterior to anus ; length of basal rod about % that of aedeagus, the rod extending anteriorly from base of aedeagus for about 3% of the distance to apex of basal membranous area ; apex of sheath without membranous extension. The area from base of sheath to tip of aedeagus with 28-40 (average 32) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath. Aedeagus long (258-315, average 293 uw), penial sheath longer and basisternum shorter, the ratios being I : 2-0—2-3 (average 2-1) and 1 : 0-74—-0-82 (average 0-78) respectively.
Material examined : 10 specimens, collected by N. S. Borchsenius on Abies sp. in the Nikitskii Botanical Gardens, Crimea, USSR on 25.v.54.
PHYSOKERMES Physokermes piceae (Schrank) (Text-figs. 6 and 7)
A medium-sized, robust species with comparatively short antennae and legs ; with many setae on the appendages, but few on the body itself. When mounted, total body length 1550-2140 (average 1803) uw ; width at mesothorax 380-500 (average 429) u. Wing expanse 2950-3200 (average 3063) wu.
46 MORPHOLOGY AND TAXONOMY OF ADULT MALES
Head subconical in dorsal view ; in lateral view obliquely dorsoventrally elongated, with the anterodorsal bulge not pronounced ; length from apex to pronotal ridge 217—270 (average 242) wu, width across genae 171-224 (average 195) u. Median crest sclerotized and distinctly poly- gonally reticulated ; with 5-8 (average 6-5) hair-like dorsal head setae, arranged in a group of 1-4 (average 2-8) on the anterior margin of the head and a group of 3-4 (average 3-8) more posteriorly. Muzidcranial ridge dorsally represented by a short, weak ridge between the eyes ; ventrally narrow but well defined, reaching ocular sclerite posteriorly, surrounding area with weak polygonal reticulation posteriorly. Genae large, sclerotized, not distinctly reticulated, without setae. Eyes : two pairs ; subequal ; corneae of dorsal eyes 20-30 (average 26) wu in diameter and 3-1—4-6 (average 3-9) times as much apart ; those of the ventral eyes with cornea 21-30 (average 26) uw in diameter and 1-o-1-'5 (average 1:2) times as much apart. Ocellus small. Ocular sclerite well sclerotized and polygonally reticulated throughout. Preocular ridge with ventral part reaching or almost reaching midcranial ridge. Postoculay ridge weak and tapering dorsally, well developed lateroventrally, but weak posteromedially ; below ocellus the ridge splits up, with anterior branch partly surrounding ocellus. Intevoculay ridge absent. Dorsal ocular setae absent, ventral head setae consisting of 4-8 (average 5-3) h.s., situated anterior to the ocular sclerite on each side of the midcranial ridge. Pyreoval ridge weak, sometimes interrupted. Tendon-like apodeme long. Cranial apophysis short; apex bifurcate, not reaching level of posterior margin of ventraleyes. /outh opening irregular. Anterior tentorial pits apparently absent.
Fic. 6. Physokermes piceae (Schr.), dorsal and ventral view.
OF THE FAMILY COCCIDAE 47
Antennae 1o-segmented, filiform ; 604-1011 (average 770) u long, i.e. shorter than half body length (ratio 1 : 2:12-2:63, average 2-37), about as long as posterior leg (ratio I : o-gI-1-II, average 0-99) and longer than penial sheath (ratio 1 : 1-32~1-69, average 1°47). Scape 46-72 (average 59) uw long and 42-61 (average 51) uw wide, with 3 h.s. Pedicel with distinct, polygonal dorsal reticulation ; 38-57 (average 46) u long and 42-53 (average 46) w wide ; with 2 hss. and a sensillum placodeum. Segment III bulging in middle, 1-8—2-6 (average 2-1) times longer than wide (61-91, average 73 uv long and 30-42, average 35 uw wide) ; with 1-3 (average 2-1) h.s. and 7-19 (average 11) f.s., the latter of medium length, 0-9-1-3 (average 1-1) times as long as width of segment ; with 1-6 usual sensilla basiconica. Segments 1V-IX cylindrical ; lengths of these segments (in p) 80-160 (average 114), 68-137 (average 98), 76-137 (average Iot), 65-114 (average 86), 57-95 (average 74) and 46-84 (average 61) respectively, widths varying from 25 to 34 pu, with distal segments wider than proximal ones ; with 12-37 (average 25), 14-31 (average 23), 21-30 (average 25), 17-27 (average 24), 15-23 (average 19) and 14-19 (average 17) f.s. respectively, but no h.s. ; antennal bristles on segments VIII-IX slightly thicker than f.s. Segment X : terminal part not constricted ; 46-68 (average 58) u long and 27-31 (average 30) w wide ; carrying 7-11 (average 9) f.s., 3 capitate subapical setae and 5 antennal bristles of which the 3 long ones are about ? as long as the segment and the 2 shorter ones not markedly different from the f.s. ; with 2 sensilla basiconica ventrally, one near apex and the other more proximal.
Thorax 494-673 (average 565) u long.
Prothovax. Pyronotal ridge strong, but medially interrupted by weak sclerotization. Lateral pronotal sclerites large, without setae. Medial setae absent. Post-tergites medium-sized, without striations and without setae. Pleural structures typical of the family. Steynwm with trans- verse ridge strong, median ridge reduced to a basal stalk, and a triangular sclerite. Ante- prosternal setae absent ; occasionally a hair-like prosternal seta present.
Mesothovax. Mesoprephragma with no emargination. Pyrescutum about twice as wide as long (average 163 and 88 u respectively) ; anterior margin slightly curved ; laterally bounded by the prescutal ridges and posteriorly by the prescutal suture ; polygonally reticulated ; slightly more heavily sclerotized medially. Scutum. Median membranous area subrectangular ; 68-91 (average 82) u long and 1-48—2-30 (average 1-79) times as wide, (width 122-179, average 146 w) ; without setae. Scutellum 59-103 (average 78) » long and 133-194 (average 151) pu wide, ratio being 1 : 1-8—2-4 (average 2); not tubular; without setae. Postnotum with anterior margin irregular and partly overlapped by the metathoracic fold ; postnotal apophysis and postalare well developed, the latter densely reticulated distally. Mesopostphragma with shallow emargination. Mesoplewron. Mesopleural ridge strong, but interrupted above coxal articulation ; pleural apophysis and pleural wing process well developed, the latter connected with episternum by a narrow basalave. Subalare small. Episteyrnum not reticulated ; sub- episternal ridge well developed, but below the membranous cleft indistinct and only marked by a band of dark sclerotization. Epimeron small. Lateropleurite bounded anteriorly by an extension from marginal ridge. Basisternum large, about 251 uw wide and 168 yw long, i.e. 1-7-2-4 (average 2-1) times longer than membranous area of scutum ; with strong median ridge and bounded by strong marginal and precoxal ridges ; with 1-2 (average 1-8) h.s. on or near median ridge. Furca well developed. Mesothovacic spivacle with well developed peri-
Fic. 7. Physokermes piceae (Schr.), lateral view.
48 MORPHOLOGY AND TAXONOMY OF ADULT MALES
treme ; postmesospivaculay setae absent. Tegula small, membranous bulge with a small weak sclerite posteriorly and 2-7 (average 4:2) h.s. Third axillary wing sclerite with a pronounced ventral projection at its base. Additional sclerite well defined. Antemetaspivacular setae absent.
Metathovax. Metanotum with posterior margin usually strong and well developed throughout; suspensorial sclervites small, spot-like. Postnotum consisting of a transverse sclerite on each side. Metatergal setae absent. Pleural ridge well developed, though interrupted near middle ; with a small wing process. Episteynum with anterior margin not ridge-like ; vestigial precoxal vidge present ; epimevon produced posteriorly. Metathovacic spivacle similar to mesothoracic one. WDorsospiraculay setae absent. Postmetaspivaculay setae : occasionally one h.s. present. Metasternal plate weak and irregular. Anterior and posterioy metasternal setae : 1-5 (average 2-4) and o-3 (average 1-8) h.s. respectively, arranged medially.
Wings hyaline; medium-sized : 1300-1450 (average 1367) u long and 470-580 (averago 542) u wide, ratio width to length being 1 : 2:44-2:77 (average 2:53) ; alar lobe present ; alar setae absent. Halteves well developed, 103-137 (average 117) uv. long and 27—43 (average 35) u wide ; each usually with 2 (range 1-3) apically hooked setae, which are about 62 yu long.
Legs short and moderately slender, with middle pair usually shortest and hind pair longest ; ratio length of hind leg to body length is 1 : 2:30-2:47 (average 2-41). Length of segments (in y) :
Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 57-76 78-110 165-217 209-300 63-80 23-27 600-805 (66) (94) (187) (248) (72) (25) (691)
II 65-91 78-110 154-205 209-289 74-95 26-28 608-815 (73) (88) (171) (242) (82) (27) (684)
Ill 72-95 82-110 163-220 220-314 76-110 27-30 667-866 (80) (94) (187) (262) (90) (28) (741)
Fs. slender and sometimes difficult to separate from h.s.
Coxae with 1-6 (average 3-4) f.s. on the fore and 5~14 on the middle and hind coxa, and each with 9-16 h.s. ; fore coxa without coxal bristles ; apical seta about }# as long as trochanter. Trochanters 27-38 uw wide ; with 6 oval sensilla, 1-8 f.s. and 5-6 h.s., the latter including 2 minute setae near basal ridge, one small seta on outer margin and 2 long setae, of which the longest (apical), on the fore trochanter, is 2-1-2:6 (average 2-4) times as long as width of trochanter. Jemora of medium width (42-49 yu), ratio width to length of hind femur being I : 3°6—-4-4 (average 3°8) ; with 1-11 (average 5-3), 3-14 (average 7-8) and 4-17 (average 9°8) f.s. on the fore, middle and hind femur respectively and with 6-19 h.s. on each. Tzibiae 23-30 uw wide, ratio width to length of hind tibia being 1 : 8-3—10-0 (average 9-2) ; each with 24-49 setae of which 15-26 are h.s. and 7—28 f.s., the latter about as long as width of tibia ; apical spur about the same size on all tibiae. Tarsi 23-30 u wide, hind tarsus 2-9-3-9 (average 3-3) times longer than wide ; each with 1-6 f.s. and 6-10 his. ; tarsal digitules subequal, longer than claw. Claws of medium length, about as long as width of tarsus ; slightly curved, with small denticle near tip ; ungual digitules subequal, about as long as claw.
Abdomen 380-760 (average 499) » long and 320-440 (average 376) uw wide.
Segments I-VII : tergites and sternites present on all segments ; tergites on segments II-III represented by a small sclerite on each side on the anterior margin, and on segments IV—VII represented by a small sclerite on each side on the anterior margin, and on the IV—VII segments by weak plates ; sternites consisting of weak transverse plates. Caudal extension of segment VII small, rounded, not sclerotized. Dorsal setae : f{.s. absent ; h.s. usually absent on segments I-VI, but segment VII with one on each side. Pleural setae consisting of h.s. only, which include dorsopleural setae : occasionally one on II, o—2 (average 1-1) on III and usually 2 on each of segments IV—VI, and ventvopleural setae : sometimes one on II and III, and one on each of segments IV-VI. Segment VII with 3-4 (average 3-9) h.s. Ventral setae : h.s. only, usually 2 medially on all segments, but occasionally only one or up to 4 present.
OF THE FAMILY COCCIDAE 49
Segment VIII with transverse tergite and a very large transverse sternite ; caudal extension forming a small, simple lobe ; glandular pouch with 2 long, pointed setae, whose protruding part is about twice as long as section within pouch ; with 3 h.s. on each side near pouch. Ante-anal area expanded, with 1-2 (average 1-8) strong h.s. and sometimes with indications of a small IX tergite.
Genital segment. Penial sheath long, about 7 total body length (ratio 1 : 3-2-3-6, average 3°4), 456-597 (average 530) vp long and 43-53 (average 48) uw wide ; the basal area anterior to the aedeagus about twice as wide as rest of sheath ; lateral sclerotizations not joined anterior to anus ; length of basal rod about 4+—4 that of aedeagus, the rod extending anteriorly from base of aedeagus for about } of the distance to the apex of basal membranous area ; apex of sheath without membranous extension. The area from base of sheath to tip of aedeagus with 23-36 (average 29) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath. Aedeagus long (285-380, average 342 wu), penial sheath longer and basisternum shorter, ratios being I : 1:49-1-61 (average 1:55) and 1 : 0-43—0-54 (average 0-49) respectively.
Material examined : 10 specimens, collected by J. Rehdéek on Picea sp. on 12.iv.53 in PleSivec, Czechoslovakia ; remounted from Swann’s mountant.
P. piceae (Schr.) differs from P. insignicola (Craw) in having only 2 pairs of simple eyes. Moulton (1907) found 4 pairs in P. insignicola. P. piceae is also described as having 2 pairs of eyes by Jancke (1955).
RHODOCOCCUS Rhodococcus spiraeae (Borchsenius) (Text-figs. 8 and 9)
A short, robust species with comparatively long antennae and moderately long legs ; with numerous setae on the appendages, but few on the body itself. When mounted, total body length 1500-1600 (average 1557) uw; width at mesothorax 400-420 (average 411) wu. Wing expanse 2860-3000 (average 2940) wu.
Head subconical in dorsal view ; in lateral view obliquely dorsoventrally elongated, with the the anterodorsal bulge not pronounced ; length from apex to pronotal ridge 198-213 (average 206) uw, width across genae 194-232 (average 217) uw. Median crest broad posteriorly, weakly sclerotized and reticulated, with 3—6 (average 4-7) hair-like dorsal head setae. Midcranial ridge dorsally absent ; -ventrally narrow but well defined, reaching ocular sclerite posteriorly, surrounding area showing weak polygonal reticulation posteriorly. Genae large, sclerotized, not reticulated, without setae. Eyes: four pairs ; dorsal and ventral pairs large, subequal ; lateral pairs smaller, subequal ; corneae of dorsal eyes 19-23 (average 21) u in diameter and 4°4-5°6 (average 5-0) times as much apart ; those of the ventral eyes 17-23 (average 19) u in diameter and 1-5—2:2 (average 1-8) times as much apart. Ocellus small, situated laterally. Ocular sclerite well sclerotized and polygonally reticulated throughout ; dorsally widely separ- ated from median crest. Pveoculay ridge of variable length, ventral part usually extending about half-way from articular process to midcranial ridge. Postoculay ridge very weak dorsally, some- times missing posterior to ocellus ; well developed lateroventrally, but weak posteromedially ; below ocellus the ridge splits up, with the anterior branch partly surrounding ocellus. Jmnter- ocular ridge absent. Dorsal ocular setae absent ; ventral head setae : o-4 (average 2:1) hs., situated on or immediately anterior to margin of the ocular sclerite on each side of midcranial ridge. Preoral ridge present. Tendon-like apodeme long, with a broad base. Cranial apophysis broad, of medium length ; apex truncate with a central lobe, not reaching level of anterior margin of eyes. Mouth opening irregular. Anterior tentorial pits apparently present antero- lateral to mouth opening.
50 MORPHOLOGY AND TAXONOMY OF ADULT MALES
Antennae 1to-segmented, filiform ; 980-1017 (average 995) u long, i.e. longer than half body length (ratio I : 1:52-1:61, average 1-57), longer than posterior leg (ratio I : 1:14-1:21, average 1-17) and longer than penial sheath (ratio I : 2:44-2:63, average 2°53). Scape 38-51 (average 45) uw long and 46-49 (average 48) uw wide, with 3 h.s. Pedicel with distinct, polygonal dorsal reticulation ; 46-57 (average 54) w long and 42-48 (average 45) uw wide ; with 3-7 (average 5-3) f.s., 1-2 (average 1-4) h.s. and a sensillum placodeum. Segment III club-shaped, 2-4-3-7 times longer than wide (72-95, average 82 uw long and 23-34, average 29 u wide) ; with 2-4 (average 2-8) f.s. of medium length, 0-9-1-4 (average 1) times as long as width of segment ; with 2-4 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths of these segments (in uw) 148-179 (average 162), 152-175 (average 165), 122-171 (average 142), 114-137 (average 125), 80-87 (average 86) and 65-74 (average 68) respectively, all of about the same width, varying from 19 to 27 w; with 17-22 (average 20), 19-24 (average 22), 20-33 (average 24), 22-28 (average 25), 15-19 (average 17) and 13-17 (average 15) f.s. respectively, but no h.s. ; antennal bristles on segments VIII-IX not markedly different from f.s. Segment X : terminal part not constricted ; 61-72 (average 67) uw long and 17-23 (average 21) uw wide ; carrying 5-11 (average 8) f.s., 3 capitate subapical setae and 5 antennal bristles of which the 3 long ones are about half as long as the segment and the 2 shorter ones rather similar to the f.s. ; with 2 sensilla basonica ventrally, one near apex and the other more proximal.
Thorax 505-559 (average 520) yu long.
Prothovax. Pyronotal ridge strong, but medially interrupted by weak sclerotization. Lateral pronotal sclerites small, without setae. Medial pronotal setae absent. Post-tergites medium- sized, without striations and without setae. Pleural structures typical of the family. Sternum with strong transverse ridge, median ridge reduced to a basal stalk, and a triangular sclerite. Anteprosternal and prosternal setae absent.
= . PO Pree Naeger
DAS AA A NEA SS sss ‘ xe
nie,
0S C66 OE |
ee ___.60p
Fic. 8. Rhodococcus spivaeae (Borchs.), dorsal and ventral view.
OF THE FAMILY COCCIDAE 51
Mesothovax. Mesoprephragma with shallow emargination. Pyrescutum about twice as wide as long (average 156 and 82 u respectively) ; anterior margin slightly curved; laterally bounded by the prescutal ridges and posteriorly by the prescutal suture ; sometimes with weak polygonal reticulation. Scutum. Median membranous area subrectangular ; 49-65 (average 59) u long and 1:81-2:36 (average 2-01) times as wide (width 106-125, average 117 u) ; without setae. Scutellum large, 76-95 (average 84) u long and 125-144 (average 135) uw wide, the ratio being I : 1-5-1-7 (average 1-6); not tubular; without setae. Postnotum with anterior margin irregular and partly overlapped by the metathoracic fold ; postnotal apophysis and postalare well developed, the latter densely reticulated distally. A/esopostphragma with moderately deep emargination. Mesopleuron. Mesopleural ridge strong, but interrupted above coxal articulation ; pleural apophysis and pleural wing process well developed, the latter connected with episternum by a narrow basalare. Subalave small. Episternum not reticulated ; sub- episternal vidge well developed, but below membranous cleft indistinct and only marked by a band of dark sclerotization. Epimeron small. Lateropleurite partly bounded anteriorly by an extension from marginal ridge. Basisternum large, about 237 » wide and 160 y long, i.e. 2°4-3°4 (average 2-8) times longer than membranous area of scutum ; with strong median ridge and bounded by strong marginal and precoxal ridges. Furca well developed. Meso- thoracic spivacle with well developed peritreme ; postmesospivaculay setae absent. Tegula small, membranous bulge with a small weak sclerite posteriorly and 2~—5 (average 3:2) hs. Third axillary wing sclerite with a pronounced ventral projection at its base. Additional sclervite well defined. Antemetaspivacular setae absent.
Metathorax. Metanotum with thickening of posterior margin desclerotized medially ; suspensorial sclerites small, spot-like. Postnotum consisting of a transverse sclerite on each side. Metatergal setae absent. Pleural ridge well developed, though interrupted in the middle ; with small wing process. Episternum with anterior margin ridge-like in parts ; vestigial pre- coxal vidge present ; epimeron produced posteriorly. Metathovacic spivacle similar to meso- thoracic one. Dorsospivaculay setae absent. Postmetaspivaculay setae : occasionally one h:s. Metasternal plate weak and irregular. Anterior and posterior metasternal setae absent.
Wings hyaline ; long (1270-1350, average 1310 uw) and of medium width (480-540, average 512 uw), ratio width to length being 1 : 2:49-2:65 (average 2:57) ; alar lobe present ; alar setae : 0-3 (average 1-5) h.s. on each wing. Halteres well developed, 114—129 (average 121) u long and 27-42 (average 37) uw wide, each with 2 (rarely 3) apically hooked setae, which are about 69 yu long.
Legs moderately long and slender, with middle pair usually shortest and hind pair longest ; ratio length of hind leg to body length is 1 : 1-72—1-83 (average 1-78). Length of segments
(in yw) :
Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 65-76 99-110 205-228 308-315 76-95 27-30 798-834 (68) (103) (218) (312) (89) (29) (819)
Il 72-84 95-105 182—200 308-317 91-99 30-34 798-820 (81) (ror) (192) (311) (97) (31) (813)
Ill 76-87 99-110 186-209 312-352 103-106 30-34 834-882 (83) (104) (200) (338) (105) (32) (862)
Fic. 9. Rhodococcus spivaeae (Borchs.), lateral view.
52 MORPHOLOGY AND TAXONOMY OF ADULT MALES
F.s. slender and sometimes difficult to separate from h.s.
Coxae each with 4-9 f.s. and 10-19 h.s. ; fore coxa without coxal bristles ; apical seta about 2 as long as trochanter. Tvochanters 29-38 u wide ; with 6 oval sensilla ; with 5-10 f.s., and 8-10 (average 8-8), 10-14 (average 12) and 12-22 (average 16) h.s. on fore, middle and hind trochanter respectively, the h.s. including 2 minute setae near basal ridge, one small seta on outer margin and 2 long setae of which the longest (apical), on the fore trochanter, is 2-3—3-1 (average 2-9) times as long as width of trochanter. Femova of medium width (42-51) u, ratio width to length of hind femur being 1 : 3:8—4-4 (average 4-1) ; each with 6-14 f.s. and 11-25 h.s. Tibiae 23-40 w wide, ratio width to length of hind tibia being 1 : 11-4—14-2 (average 12°6) ; each with 45—68 setae of which 30-60 are h.s. and 8-18 f.s., the latter somewhat longer than width of tibia ; apical spur about the same size on all tibiae. Tavsi 21-27 uw wide, hind tarsus 4:3-4°7 (average 4:6) times longer than wide ; each with 4-8 f.s. and 15-26 his. ; tarsal digitules subequal, long, about 14 times as long as claw. Claws long, hind claw about 14 times as long as width of tarsus ; slightly curved, with small denticle near tip ; ungual digitules subequal, about as long as claw.
Abdomen 430-500 (average 474) u long and 300-350 (average 331) uw wide.
Segments I-VII : tergites on segments [I-III represented by a small sclerite on each side on the anterior margin, and on VI and VII by a weak plate ; steynites present on segments II, III and VII, represented by weak plates. Caudal extension of segment VII small, rounded, not sclerotized. Dorsal setae: f.s. absent ; h.s. absent on I and usually on II, but usually one on each side on segments III-VII. Pleural setae consisting of h.s. only, which include dorso- pleural setae : o-2 (average 1-0), 2-5 (average 3-7), 4-7 (average 6) and 4-8 (average 5:8) on segments III—VI respectively, and ventropleural setae : occasionally one on III and usually one on each of segments IV-VI. Segment VII with 6-15 (average 11) hs. Ventral setae: hss. only, occasionally one on II, usually 2 medially on III, and 4 on each of segments IV—VII.
Segment VIII with transverse tergite and sternite ; caudal extension forming a small, simple lobe ; glandular pouch with 2 long, pointed setae, whose protruding part is 13-2 times as long as section within pouch. A small IXth tergite sometimes perceptible. Avnte-anal setae : 1-2 (average 1-8) long h.s. Posterior margin with 2-3 (average 2-8) h.s. on each side.
Genital segment. Penal sheath long, about }+ total body length (ratio 1 : 3-8—4-1, average 3:9), 376-414 (average 399) uw long and 36-40 (average 38) uw wide ; lateral sclerotizations not joined anterior to anus ; length of basal vod about ? that of aedeagus, the rod extending anteriorly from base of aedeagus for about ? of the distance to apex of basal membranous area ; apex of sheath without membranous extension. The area from base of sheath to tip of aedeagus with 19-23 (average 21) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath. Aedeagus long (167-182, average 173 uw), penial sheath longer and basisternum shorter, the ratios being 1: 2:25—2-42 (average 2:30) and 1: 0-83—0-98 (average 0-92) respectively.
Material examined : 9g specimens, collected by G. Matesova on Spiraea hypersifolia L. in the ravine Talgar, Zailiiski Alatau, Kazakhstan, USSR, on 14.v1.57.
PALAEOLECANIUM Palaeolecanium bituberculatum (Targ.) (Text-figs. 10 and 11)
Living specimens coral-red in colour, with the sclerotized areas brown and the appendages light yellow, wings with a reddish tinge between anterior margin and first wing vein ; short, slender, with comparatively long antennae and short legs which carry many setae. When mounted, total body length 1380-1460 (average 1419) uw; width at mesothorax 290-320 (average 310) uw. Wing expanse 2470-2660 (average 2566) uw.
Head subconical in dorsal view ; in lateral view obliquely dorsoventrally elongated, with the anterodorsal bulge not pronounced ; length from apex to pronotal ridge 205—232 (average
a
OF THE FAMILY COCCIDAE 53
220) uw, width across genae 217-241 (average 233) uw. Median crest sclerotized and distinctly polygonally reticulated ; with 6-9 (average 7-4) hair-like dorsal head setae, arranged in a group of one or two near the anterior margin of the head and a group of 5~8 (average 6:2) more posteriorly. Midcranial ridge dorsally represented by a short, weak ridge near anterior margin; ventrally narrow, but well defined, reaching ocular sclerite posteriorly, surrounding area with weak, polygonal reticulation posteriorly. Genae large, sclerotized, not distinctly reticulated, without setae. Eyes : two pairs ; subequal ; corneae of dorsal eyes 21-25 (average 22) u in diameter and 3:4—4:2 (average 3-9) times as much apart ; those of the ventral eyes 22-27 (average 24) uw in diameter and 1-2~-1-8 (average 1-6) times as much apart. Ocellus small. Ocular sclerite well sclerotized and polygonally reticulated throughout. Pyreoculay ridge with ventral part almost reaching midcranial ridge. Postocular ridge weak and tapering dorsally, well developed lateroventrally, but weak posteromedially ; below ocellus the ridge splits up, with anterior branch partly surrounding ocellus. Interoculay vidge absent. Dorsal ocular setae absent ; ventral head setae : 4-7 (average 5-6) h.s., situated anterior to the ocular sclerite on each side of the midcranial ridge. Preoral ridge present. Tendon-like apodeme short. Cranial apophysis of medium length ; apex bifurcate with a central lobe, extending to around the level of the posterior margin of the ventral eyes. Mouth opening irregular. Anterior tentorial pits apparently absent.
Antennae to-segmented, filiform ; 705-883 (average 819) yu long, i.e. longer than half body length (ratio 1 : 1-63—-1-96, average 1-76), longer than posterior leg (ratio 1 : 1:15-1:22, average 1-19) and longer than penial sheath (ratio 1 : 2:83-3:26, average 3-09). Scape 46-57 (average 51) uw long and 40-46 (average 42) uw wide, with 3 h.s. Pedicel with distinct, polygonal, dorsal reticulation ; 42-51 (average 47) uw long and 38-42 (average 40) uw wide ; with 1-7 (average 4°3) f.s., 2-5 (average 3:5) f.s. and a sensillum placodeum. Segment III club-shaped, 2:7—3:6 (average 3-2) times longer than wide (76-87, average 81 uw long and 21-30, average 26 wu wide),
Fic. 10. Palaeolecanium bituberculatum (Targ.), dorsal and ventral view.
54 MORPHOLOGY AND TAXONOMY OF ADULT MALES
with 2-8 (average 4-8) f.s. of medium length, 1-3-1-7 (average 1-5) times as long as width of segment ; with 1-2 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths of these segments (in up.) 87-122 (average 109), 195-125 (average 118), 91-114 (average Io1), 80-110 (average 100), 65-91 (average 76) and 57—76 (average 68) respectively, width varying form 17 to 23 vp, with distal segments wider than proximal ones ; with 12-19 (average 15), 15-22 (average 18), 12-19 (average 15), 15-20 (average 16), 11-19 (average 14) and 11-14 (average 13) f.s. respec- tively, but no h.s. ; antennal bristles on segments VIII and IX not markedly different from f.s., sometimes somewhat thicker. Segment X : terminal part not constricted ; 53-78 (average 70) up. long and 19-23 (average 21) uw wide ; carrying 4-11 (average 8-6) f.s., 3 capitate subapical setae and 5 antennal bristles of which the 3 long ones are about half as long as the segment and the 2 shorter ones equal to or somewhat shorter than the f.s. ; with 2 sensilla basiconica ventrally, one near apex and the other more proximal.
Thorax 445-517 (average 497) u long.
Prothovax. Pyronotal ridge strong, but medially interrupted by weak sclerotization. Lateral pronotal sclerites medium-sized, without setae. Medial pronotal setae : 2 small h.s., situated close together on the median line. Post-teygites very small and weakly sclerotized, without striations and without setae. Pleural structuyes typical of family. Steynuwm with strong transverse ridge, median ridge reduced to a basal stalk or sometimes absent, and a triangular sclerite. Anteprosternal setae absent ; prostevnal setae : occasionally one h.s. present.
Mesothovax. Mesoprephragma with shallow emargination. Prescutum about twice as wide as long (average 145 and 74 yp respectively) ; anterior margin curved ; laterally bounded by the prescutal vidges and posteriorly by the prescutal suture ; weakly irregularly reticulated ; with a median ridge extending for some distance anteriorly from the prescutal suture. Scutum. Median membranous avea subrectangular ; 76-91 (average 82) uw long and 1-47—1-75 (average 1°57) times as wide (width 118-137, average 128) ; with 1-4 (average 2-7) h.s. Scutellum 57-68 (average 63) uw long and 118-133 (average 127) uw wide, ratio being I : 1:9-2-2 (average 2) ; not tubular ; occasionally with one or two h.s. Postnotum with anterior margin irregular and partly overlapped by the metathoracic fold ; postnotal apophysis and postalave well developed, the latter densely reticulated distally. Mesopostphvagma with deep emargination. Mesopleuron. Mesopleuval vidge strong, but interrupted above coxal articulation ; pleural apophysis and pleural wing process well developed, the latter connected with episternum by narrow basalave. Subalave small. Episternum not reticulated ; subepisternal ridge well developed, but below membranous cleft indistinct and only marked by a band of dark sclero- tization. Epimevon small. Latervopleurite anteriorly bounded by an extension from marginal ridge. Basisternum large, about 199 up wide and 148 yu long, i.e. 1:7-2:1 (average 1-8) times longer than membranous area of scutum ; with strong median ridge and bounded by strong marginal and precoxal vidges ; without setae. Furca well developed. Mesothorvacic spivacle with well developed peritreme ; postmesospivacular setae absent. Tegula small, membranous bulge with a small weak sclerite posteriorly and 4—7 (average 5) h.s. Third axillary wing sclerite with a pronounced ventral projection at its base. Additional sclervite well defined. Antemeta- spivaculay setae absent.
Fic. 11. Palaeolecaniwm bituberculatum (Targ.), lateral view.
OF THE FAMILY COCCIDAE 55
Metathorax. Metanotum with anterior margin desclerotized medially ; suspensorial sclerites small, spot-like. Postnotum consisting of a transverse sclerite on each side. Metatergal setae : one h.s. on each side. Pleural ridge well developed, though interrupted near the middle ; with a small wing process. Episternum with anterior margin not ridge-like ; vestigial precoval vyidge sometimes present ; epimeron produced posteriorly. Metathoracic spiracle similar to mesothoracic one. Dorsospivaculay setae absent. Postmetaspivaculay setae : occasionally one h.s. Metasternal plate weak and irregular. Anterior and posterior metasternal setae absent.
Wings hyaline ; long (1110-1190, average 1155 w) and of medium width (450-510, average 474 wv), ratio width to length being 1 : 2:33-2:58 (average 2-44) ; alar lobe present ; alar setae absent. Halteres well developed, 95-114 (average 103) p long and 20-30 (average 25) u wide, each with one apically hooked seta, which is about 50 yu long.
Legs short and slender, with the fore pair longest and the middle pair shortest ; ratio length of hind leg to body length is 1 : 1:94-2:06 (average 1-99). Length of segments (in yu) :
Leg Coxa —_ Trochanter Femur Tibia Tarsus Claw Total
I 57-61 76-84 171-186 255-285 89-99 25-27 695-735 (58) (80) (183) (274) (95) (26) (715)
Il 61-65 68-72 163-175 247-277 87-95 23-25 654-701 (63) (70) (169) (261) (90) (24) (678)
Ii 57-68 67-72 163-175 257-289 87-99 24-27 669-722 (65) (69) (170) (273) (94) (26) (697)
Coxae with 7-14 (average 10) f.s. on the fore and 14—19 on the middle and hind coxa, and each with 7-15 h.s. ; fore coxa with 1-2 (average 1-4) pointed coxal bristles, about as long as segment; apical seta about half as long as trochanter. Tyvochanters 24-29 u wide ; with 6 oval sensilla, 4-8 f.s. and 6-8 h.s., the latter including 2 minute setae near basal ridge, one small seta on outer margin and 2 long setae of which the longest (apical), on the fore trochanter, is 1-8—2-2 (average 2) times as long as width of trochanter. Femora of medium width (34-40 yp), ratio width to length of hind femur being 1 : 4-4-5-0 (4-7) ; each with 8-14 f.s. and 12-19 hs. Tuibiae 19-23 pw wide, ratio width to length of hind tibia being 1 : 12-3-14-6 (average 13-1) ; each with 35-51 setae of which 19-27 are h.s. and 15-30 f.s., latter somewhat longer than width of tibia ; apical spur about the same size on all tibiae. Tarsi 19-23 uw wide, hind tarsus 4:2—4-9 (average 4°4) times longer than wide ; each with 4-8 f.s. and 11-17 his. ; tarsal digitules subequal, longer than claw. Claws of medium length, a little longer than width of tarsus ; slightly curved, with small denticle near tip ; ungual digitules subequal, about as long as claw.
Abdomen 420-480 (average 461) uw long and 240-300 (average 263) uw wide.
Segments I-VIL : tergites on segments II-III represented by small sclerites on the anterior margin—one on each side and one medially on II, one on each side on III, and on segments VI-VII by weak transverse plates ; sternites present on all segments, represented by a weak transverse plate on the anterior and posterior segments and a small sclerite on each side on the intermediate segments. Caudal extension of segment VII small, rounded, not sclerotized. Dorsal setae : h.s. only, usually one on each side on all segments. Pleural setae consisting of h.s. only, which include dorsopleural setae : occasionally one on III and usually 2 on each of segments IV—VI, and ventyvopleural setae : occasionally one on each of segments IV—V and usually one on VI. Segment VII with 3-5 (average 3:8) h.s. Ventral setae: h.s. only, 2 medially on II, and 1-3 (average 2-4), 2-4 (average 3), 3-4 (average 3-5), 3-5 (average 3-9) and 3-4 (average 3-8) on segments III-VII respectively.
Segment VIII with transverse tergite and sternite ; caudal extension forming a small, simple lobe ; glandular pouch with 2 long, pointed setae, whose protruding part is 2-24 times as long as section within pouch. No IX tergite observed. Ante-anal setae: 2 long h.s. Posterior margin with 2-4 (average 2-6) h.s. on each side.
Genital segment. Penial sheath of medium length, about + total body length (ratio 1 : 5-09— 5°65, average 5:39), 247-277 (average 266) uw long and 46-61 (average 52) uw wide ; lateral sclerotizations not joined anterior to anus; length of basal vod 4-2 that of aedeagus, the rod
56 MORPHOLOGY AND TAXONOMY OF ADULT MALES
extending anteriorly from base of aedeagus to the apex of basal membranous area ; apex of sheath without membranous extension. The area from base of sheath to tip of aedeagus with 17-22 (average 19) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath. Aedeagus broad at its base and tapering towards tip ; long (129-148, average 141 uy), penial sheath longer and basisternum as long or longer, the ratios being I : 1-8—-2-0 (average I-g) and I : 0-99-1-14 (average 1-04) respectively.
Material examined : Io specimens, bred from material collected by myself on Crataegus sp. (hawthorn) on Putney Heath, London ; males emerged in the laboratory between the 19th and 22nd June, 1962. Four specimens, collected by J. Rehaéek on Crataegus oxyacantha L. in Novy Bydiov, Czechoslovakia on 18.vi.52 (remounted from Swann’s mountant) agreed well with the above description, but differed in the following respects : (i) 2 specimens were longer, measuring 1490 u each, (ii) the darker median sclerotization on the prescutum was less well developed and not ridge-like in appearance and (ili) the reticulation on the prescutum was more distinct.
PHYLLOSTROMA Phyllostroma myrtilli (Kaltenbach) (Text-figs. 12 and 13)
A small, slender species with comparatively long antennae and short legs ; with many setae on the appendages, but few on the body itself. When mounted, total body length 1460-1620 (average 1553) uw; width at mesothorax 370-410 (average 387) u. Wing expanse 2870-3010 (average 2940) uw.
Head subconical in dorsal view ; in lateral view obliquely dorsoventrally elongated, with anterodorsal bulge not pronounced ; length to pronotal ridge 209-247 (average 226) uw, width across genae 255-270 (average 262) u. Median crest sclerotized and distinctly polygonally reticulated ; with 6—10 (average 7-4) hair-like dorsal head setae, arranged in a group of 3-5 near anterior margin of head and a group of 2—5 more posteriorly. Midcvanial ridge dorsally absent; ventrally narrow but well defined, surrounding area weakly sclerotized and distinctly poly- gonally reticulated. Genae large, sclerotized, not reticulated, without setae. Eyes four pairs ; dorsal and ventral pairs large, subequal ; lateral pairs smaller, subequal ; corneae of dorsal eyes 23-27 (average 24) wu in diameter and 3-4-4°5 (average 4-1) times as much apart ; those of the ventral eyes 25-30 (average 28) pw in diameter and 1-3-1-7 (average 1-5) times as much apart. Ocellus small. Ocular sclerite well sclerotized and polygonally reticulated throughout. Preoculay ridge short, not extending far below articular process. Postocular ridge weak and tapering dorsally, but well developed lateroventrally and posteromedially ; below ocellus the ridge splits up, with anterior branch partly surrounding ocellus. Intevocular ridge absent. Dorsal ocular setae absent ; ventral head setae : 8-14 (average Io) hs., situated around posterior part of the midcranial ridge. Pyreoval vidge present. Tendon-like apodeme short. Cranial apophysis of medium length ; apex bifurcate with central lobe, not reaching level of anterior margin of ventral eyes. Mouth opening irregular. Anterior tentorial pits present, situated anterolateral to mouth opening.
Antennae to-segmented, filiform ; 910-1053 (average 986) u long, i.e. longer than half body length (ratio I : 1:43-1:65, average 1-57), longer than posterior leg (ratio I : 1:23-1-33, average 1-27) and longer than penial sheath (ratio I : 3:15-3:47, average 3:36). Scape long (length 65-68, average 66 uw) and 42-48 (average 45) uw wide, with 1-3 (average 2) h.s. Pedicel not reticulated, but with one or two wavy lines ; 61-68 (average 65) u long and 42-46 (average 44) u wide; with 5-10 (average 7:4) f.s., 3-5 (average 3-7) h.s., and a sensillum placodeum. Segment III club-shaped, 2:5—3:1 (average 2-8) times longer than wide (76-84, average 79 u long
OF THE FAMILY COCCIDAE 57
and 42-46, average 44 u wide), with 4-8 (average 6) f.s. of medium width, 1-2—1-5 (average 1-3) times as long as width of segment ; with 1-3 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths of these segments (in uw) 118-152 (average 138), 122-148 (average 137), 118-141 (average 131), 99-133 (average 111), 91-106 (average 97) and 80~99 (average 86) respectively, widths varying from 21-27 » with distal segments wider than proximal ones ; with 15-22 (average 18), 15-23 (average 20), 18-28 (average 23), 16-25 (average 20), 14-18 (average 16) and 15~—21 (average 18) f.s. respectively, and occasionally with one or two h.s. on segments IV-VI ; antennal bristles on segments VIII-IX not markedly different from f.s. Segment X : terminal part not constricted ; 67-86 (average 76) u long and 20-23 (average 22) uw wide ; carrying 5-8 (average 6-4) f.s., 3 capitate subapical setae, and 5 antennal bristles of which the 3 longest ones are about half as long as the segment and the 2 shorter ones rather similar to the f.s. ; with 2 sensilla basiconica ventrally, one near apex and the other more proximal.
Thorax 460-532 (average 503) pu long.
Prothovax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral pronotal sclerites small, without setae. Medial pronotal setae : usually 2 h.s. Post-tergites small and irregular, without striations and without setae. Pleural structures typical of family. Sternum with strong transverse and median ridges, well sclerotized triangular sclerite and rather distinct sternal apophyses. Anteprosternal setae absent ; prosternal setae : sometimes one h.s. on each side.
Mesothorvax. Mesoprephragma with shallow emergination. Prescutum about twice as wide as long (average 160 and 83 yp respectively) ; anterior margin strongly curved ; laterally bounded by the prescutal ridges and posteriorly by the prescutal suture ; slightly more heavily sclerotized medially but not reticulated. Scutum. Median membranous area subrectangular ;
PES SSP
4
Fic. 12. Phyllostroma myrtilli (Kalt.), dorsal and ventral view.
58 MORPHOLOGY AND TAXONOMY OF ADULT MALES
68-91 (average 78) u long and 1-79—2-17 (average 1-94) times as wide (width 137-163, average 150 uw) ; with 1-2 (average 1-7) h.s. Scutellum 57-68 (average 64) uv long and 160-171 (average 166) uw wide, the ratio being 1 : 1-8-2-2 (average 1-9) ; not tubular ; without setae. Post- notum with anterior margin irregular, not overlapped by the metathoracic fold ; postnotal apophysis and postalave well developed, the latter densely reticulated distally. Mesopost- phragma small, with moderately deep emargination. Mesopleuron. Mesopleural ridge strong, not interrupted above coxal articulation ; pleural apophysis and pleural wing process well developed, latter connected with episternum by a narrow basalare. Subalave small. Epi- steynum not reticulated ; subepisternal vidge well developed, but below membranous cleft in- distinct and only marked by a band of dark sclerotization. Epimeron small. Lateropleurite partly bounded anteriorly by an extension from marginal ridge. Basisteyvnum large, about 210 uw wide and 151 u long, i.e. 1-7—2-2 (average 1-9) times longer than membranous area of scutum ; with strong median ridge and bounded by strong marginal and precoval ridges ; without setae. Furca well developed. Mesothoracic spivacle with well developed peritreme ; postmesospiracular setae absent. Tegula small, membranous bulge with a small weak sclerite posteriorly and 0-4 (average 1-9) h.s. Third axillary wing sclerite with a small ventral projection at its base. Additional sclerite well defined. Antemetaspivacular setae absent.
Metathorax. Metanotum with anterior margin desclerotized medially ; suspensorial sclerites absent ; a small, additional sclerite sometimes present anterior to postnotum. Postnotum consisting of a transverse sclerite on each side. Metatergal setae : occasionally one h.s. on each side. Pleural ridge reduced, extending only for a short distance above coxal articulation. Vestigial precoxal ridge present. Episternum reduced to a small subtriangular plate ; epimeron produced posteriorly. Metathoracic spivacle similar to mesothoracic one. Dorsospivacular setae absent. Postmetaspivacular setae : 2-6 (average 4) f.s. and o-1 (average 0-7) h.s. Meta- steynum represented by a weak, transverse plate. Antevioy and posterioy metasternal setae absent.
Wings hyaline ; long (length 1260-1330, average 1295 wu.) and broad (width 570-600, average 585 uw), ratio width to length being I : 2:19-2:24 (average 2:22) ; alay lobe and alar setae absent. Halteres absent.
Legs short and moderately slender, with the fore pair usually longest and the middle pair shortest ; ratio length of hind leg to body length is 1 : 1-89—2-05 (average 1:99). Length of segments (in pu) :
Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 65-80 91-99 198-213 285-304 99-122 23-27 785-828 (74) (95) (207) (295) (107) (26) (804)
II 68-80 87-93 182-194 281-296 99-106 23-29 762-785 (75) (91) (189) (289) (102) (26) (771)
Ill 76-91 QI-—99 179-190 289-300 99-110 21-29 771-800 (83) (95) (184) (296) (106) (26) (788)
Coxae with 11-16 (average 14) f.s. on the fore and 16-22 on the middle and hind coxa, and each with 9-12 h.s. ; fore coxa without coxal bristles ; apical seta about 4 as long as trochanter. Trochanters 30-34 u wide ; with 6 oval sensilla, 6-14 f.s. and 5-8 h.s., the latter including 2
Fic. 13. Phyllostvoma myrtilli (Kalt.), lateral view.
OF THE FAMILY COCCIDAE 59
minute setae near basal ridge, one small seta on outer margin and a long apical seta which, on the fore trochanter, is 1-3-2-1 (average 1-8) times as long as width of trochanter. Femora of medium width (38-46 yw), ratio width to length of hind femur being 1 : 3-9-4-6 (average 4:2) ; each with 1o-r19 f.s., and with 15-24 (average 19) h.s. on the fore and 11-16 on the middle and hind femur. Tibiae 23-27 w wide, ratio width to length of hind tibia being 1 : 11-3~-12-7 (average 12) ; each with 36-56 setae, of which 17~32 are h.s. and 14-28 f.s., the latter somewhat longer than width of tibia ; apical spur about the same size on all tibiae. Tayrsi 22-27 uw wide, hind tarsus 4:0-4°8 (average 4-5) times longer than wide ; each with 2-7 f.s. and 1o-19 hss. ; tarsal digitules subequal, longer than claw. Claws of medium length, about as long as width of tarsus ; slightly curved, with very small denticle near tip ; ungual digitules subequal, about as long as claw.
Abdomen 490-580 (average 531) up long and 320-380 (average 346) uw wide.
Segments I-VII : tergites and sternites present on all segments ; tergites represented by a transverse sclerite on or near anterior margin (sometimes interrupted medially) ; sternites represented by a weak transverse plate in middle of each segment. Caudal extension of segment VII small, rounded, not sclerotized. Dorsal setae : h.s.only, usually one on each side on segments ITand V-VII. Pleural setae consisting of h.s. only, which include dorsopleural setae : usually 2 on each of segments III—VI, and ventropleural setae : occasionally one on each of III-V, and usually on segment VI. Segment VII with 1-6 (average 3) h.s. Ventral setae: hs. only, usually none on II, one on each side on III and IV, and 4 on each of segments V—VII.
Segment VIII with transverse tergite and sternite ; caudal extension forming a small, simple lobe ; glandular pouch with 2 long, pointed setae, whose protruding part is about twice as long as section within pouch. No IXth tergite observed. Ante-anal setae usually 2 (range 1-4) h.s. Posterior margin with 3—4 (average 3-5) h.s. on each side.
Genital segment. Penial sheath of medium length, about 4 total body length (ratio 1 : 4-97- 5°57, average 5:27), 285-304 (average 294) u long and 46—49 (average 48) uw wide ; lateral sclerotizations narrowly joined anterior to anus ; length of basal rod about } that of aedeagus, the rod extending anteriorly from base of aedeagus to the apex of the basal membranous area ; apex of sheath without membranous extension. The area from base of sheath to tip of aedeagus with 11-19 (average 15) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath. Aedeagus of medium length (118-137, average 127 wu), penial sheath and basisternum longer, the ratios being 1 : 2:2—2-6 (average 2:3) and 1 : 1-11—1-29 (average I-19) respectively.
Material examined : 8 specimens, collected by J. Rehdéek on Vaccinium myrtillus L. in Czechoslovakia on 6.vii.1953 (remounted from Swann’s mountant).
FILIPPIA Filippia viburni (Signoret) (Text-figs. 14 and 15)
A moderately long and robust species, with comparatively long antennae and moderately long legs ; with numerous setae on the appendages, but few on the body itself. When mounted, total body length 1940-2080 (average 2004) p ; width at mesothorax 440-490 (average 469) w. Wing expanse 3630-3940 (average 3798) uw.
Head subconical in dorsal view ; in lateral view obliquely dorsoventrally elongated, with anterodorsal bulge not pronounced ; length from apex to pronotal ridge 262-285 (average 277) u, width across genae 289-312 (average 297) u. Median crest sclerotized and striated posteriorly, but usually without polygonal reticulation ; with 7-12 (average 9-3) hair-like dorsal head setae. Midcranial ridge dorsally absent ; ventrally narrow but well defined, reaching postocular sclerite posteriorly, surrounding area with polygonal reticulation. Genae large, sclerotized ; polygonal reticulation enclosing weaker irregular reticulation ; without setae. Eyes: five pairs ; dorsal and ventral pairs large, subequal ; lateral eyes smaller, especially the middle one
60
MORPHOLOGY AND TAXONOMY OF ADULT MALES
6 2 ae / ase / 7] / = S
Ss \ “i | | . | \ Vv
\ \ 2332 | zs
oo
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Filippia viburmi (Sign.), dorsal and ventral view.
Fic. 14.
OF THE FAMILY COCCIDAE 61
which is about half as big as the other two ; corneae of dorsal eyes 27-30 (average 29) pu in diameter and 3-0-4-3 (average 3-6) times as much apart ; those of the ventral eyes 29-33 (average 30) » in diameter and 1-:2~-1-7 (average 1-4) times as much apart. Ocellus small. Ocular sclevite well sclerotized and polygonally reticulated throughout. Preocular ridge with ventral part almost reaching midcranial ridge. Postoculay ridge tapering dorsally, well developed lateroventrally, and narrow but well defined posteromedially ; below ocellus the ridge splits up, with anterior branch partly surrounding ocellus. Jnteroculay ridge absent. Dorsal ocular setae absent ; ventral head setae : 2-10 (average 6-3) h.s., situated medially just behind the preocular ridges. Preoval ridge present. Tendon-like apodeme long. Cranial apophysis of medium length ; apex bifurcate, not reaching the level of anterior margin of ventral eyes. Mouth opening irregular. Anterior tentorial pits absent.
Antennae to-segmented, filiform ; 1129-1379 (average 1284) u long, i.e. longer than half body length (ratio 1 : 1-47-1-75, average 1-58), longer than posterior leg (ratio 1 : 1-06—1-18, average 1-13) and longer than penial sheath (ratio 1 : 3:38-3:94, average 3-74). Scape 59-65 (average 62) uw long and 46-51 (average 48) uw wide, with 3-4 (average 3:3) h.s. Pedicel with distinct, polygonal, dorsal reticulation ; 57-65 (average 62) uw long and 49-59 (average 54) uw wide ; with 2-6 (average 3-6) f.s., 7-11 (average 8-4) h.s. and a sensillum placodeum. Segment III bulging in the middle, 1-9—2-6 (average 2-1) times longer than wide (76-87, average 80 wu long and 34-39, average 35 uw wide) ; with 4-9 (average 7-1) h.s. and 4~12 (average 7-9) f.s., the latter of medium length, o-9-1-2 (average 1-1) times as long as width of segment ; with 1-3 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths of these segments (in uw) 194-220 (average 205), 182-247 (average 223), 171-224 (average 204), 129-186 (average 165), 87-118 (average 108) and 72-99 (average 86) respectively, all of about the same width, varying from 21 to 29 uw; with 33-45 (average 38), 37-44 (average 41), 30-49 (average 41), 28-39 (average 33), 7-22 (average 16) and 10-19 (average 14) f.s., and 2-4 (average 3), 3-9 (average 5-1), 2-7 (average 4), 3-6 (average 4-2), 1-4 (average 2-9) and o-5 (average 2-3) h.s. respectively ; antennal bristles on segments VIII-IX distinctly larger than f.s. Segment X : terminal part not constricted ; 86-99 (average 92) up long and 23-26 (average 24) uw wide ; carrying 6-12 (average 9) f.s., o-3 (average 1-1) h.s., 3 capitate subapical setae and 5 antennal bristles of which the 3 long ones are up to half as long as the segment and the 2 shorter ones somewhat shorter but distinctly thicker than the f.s. ; with 2 sensilla basiconica ventrally, one near apex and the other more proximal.
Thorax 733-794 (aver age 766) p long.
Prothorax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral pronotal sclerites small, without setae. Medial pronotal setae : sometimes one or two (0-2, average 0-7) h.s. Post-tergites narrow, elongated, without striations and without setae. Pleural structures typical of the family. Sternwm with a strong transverse ridge, a long median ridge, which is usually weak and interrupted posteriorly, and a narrow triangular sclerite. Anteprosternal setae absent ; prosternal setae : 1-2 (average 1-7) h.s., usually one on each side.
Fic. 15. Filippia viburni (Sign.), lateral view.
62 MORPHOLOGY AND TAXONOMY OF ADULT MALES
Mesothovax. Mesoprephragma with shallow emargination. Pyrescutum more than twice as wide as long (average 237 and 106 u respectively) ; anterior margin strongly curved ; laterally bounded by the prescutal ridges and posteriorly by the prescutal suture ; slightly more heavily sclerotized medially and irregularly reticulated. Scutuwm. Median membranous area sub- rectangular ; 120-133 (average 128) uw long and 1:54-1:94 (average 1-68) times as wide (width 201-236, average 213 uw) ; with 11-19 (average 15) h.s. Scutellum 76-86 (average 82) uw long and 201-228 (210) u wide, the ratio being 1 : 2-4—2-8 (average 2-6) ; tubular, with large ventral foramen ; without setae. Postnotum with anterior margin usually regular and exposed ; postnotal apophysis and postalave well developed, the latter densely reticulated distally. Meso- postphragma with deep emargination. Mesopleuron. Mesopleural ridge strong, but interrupted above coxal articulation ; pleuval apophysis and pleural wing process well developed, the latter connected with episternum by a narrow basalare. Subalave small. Episteynum showing weak, irregular reticulation ; subepisternal ridge well developed, but below membranous cleft indistinct and only marked by a band of dark sclerotization. Epimevon small. Latervopleurite anteriorly bounded by an extension from marginal ridge. Basisternum large, about 293 u wide and 231 uw long, i.e. 1:66—-1-95 (average 1-81) times longer than membranous area of scutum ; with strong median ridge and bounded by strong marginal and precoxal ridges ; without setae. Furca well developed. Mesothovacic spivacle with well developed peritreme ; postmesospivacular setae absent. Tegula small, membranous bulge with a small weak sclerite posteriorly and 5-9 (average 7) h.s. Third axillary wing sclerite with a pronounced ventral projection at its base. Additional sclerite small, but well defined. Antemetaspivacular setae absent.
Metathovax. Metanotum with anterior margin usually weak medially ; suspensorial sclerites small, rod-like ; a small, additional sclerite present anterior to postnotum. Postnotum con- sisting of a transverse sclerite on each side. Metatergal setae : one h.s. on each side. Pleural vidge well developed, though interrupted in middle ; with small wing process. Episternum with anterior margin not ridge-like ; vestigial precoxal ridge present ; epimeyon produced posteriorly. Metathoracic spivacle similar to mesothoracic one. Dorsospivacular setae absent. Postmetaspivaculay setae: 4-9 (average 5:5) fs. and o-5 (average 2:5) hs. Metasternum represented by a weak, transverse plate. Antevioy and posterior metasternal setae : 0-3 (average 0-8) and o-1 (average 0-6) h.s. respectively.
Wings hyaline ; long (length 1650-1770, average 1713 w) and broad (width 750-860, average 818 uw), ratio width to length being 1 : 2:04-2:25 (average 2-09) ; alar lobe present ; alar setae absent. Halteves well developed, 106-133 (average 123) uw long and 23-33 (average 26) uw wide, each with one apically hooked seta which is about 68 y long.
Legs moderately long and slender, with middle pair shortest and hind pair longest ; ratio length of hind leg to body length is 1 : 1-76-1-85 (average 1:80). Length of segments (in p) :
Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 74-84 105-118 258-308 456-483 129-139 23-27 1062-1144 (78) (113) (284) (467) (133) (25) (rror)
II QI-103 105-118 250-277 418-448 129-137 23-26 1018-1100 (96) (113) (266) (439) (134) (25) (1072)
Ill 106-118 110-118 266-289 429-479 127-141 25-29 1068-1163 (113) (116) (276) (451) (136) (26) (1119)
Coxae each with 21-32 f.s. and 12-23 h.s. ; fore coxa with 1-3 (average 2) pointed coxal bristles which are about 4 as long as the segment ; apical seta about } as long as the trochanter. Tvochanters 34-38 » wide ; with 6 oval sensilla, 13-20 f.s. and 7-10 h.s., the latter including 2 minute setae near basal ridge, one small seta on outer margin and a long apical seta which, on the fore trochanter, is 2-6—-3-0 (average 2-7) times as long as width of the trochanter. Femova of medium width (46-49 yw), ratio width to length of hind femur being I : 5:5-6:3 (average 5:8); each with 34-45 fs. and 18-34 hs. Tuibiae 27-30 uw wide, ratio width to length of hind tibia being 1 : 14-1-17-3 (average 15-4) ; each with 94-130 setae, of which 36-50 are h.s. and 56-80 f.s., the latter about as long as width of
OF THE FAMILY COCCIDAE 63
tibia ; apical spur about the same size on all tibiae. Tayrsi 23-27 uw wide, hind tarsus 5-1-5-3 (average 5-2) times longer than wide ; each with 13-19 f.s. and 15-22 h.s. ; tarsal digitules subequal, longer than claw. Claws of medium length, about as long as width of tarsus ; slightly curved, with small denticle near tip ; wngual digitules subequal, about as long as claw.
Abdomen 620-710 (average 669) u long and 400-500 (average 446) uw wide.
Segments I-VI1 : tergites on segments II-III represented by 3 small sclerites on anterior margin, one medially and one on each side, and on VII by a weak transverse plate in middle of segment ; sternites present on all segments, represented by a weak transverse plate on anterior and posterior segments and a small sclerite on each side on intermediate segments. Caudal extension of segment VII small, rounded, not sclerotized. Dorsal setae : h.s. only, usually one on each side on each of segments I and V-VII and sometimes one or two on IV. Pleural setae consisting of h.s. only which include dorsopleural setae : up to 3 on II, 2—3 on III and usually 4 (range 3-5) on each of segments IV—VI, and ventropleural setae : usually one on segment VI. Segment VII with 6-10 (average 7:3) h.s. Ventral setae : h.s. only, occasionally one on each side on II, usually one on each side on III and IV, and 4 on each side of segments V—VII.
Segment VIII with transverse tergite and sternite ; caudal extension forming a small, simple lobe ; glandular pouch with 2 long, pointed setae, whose protruding part is about twice as long as section within pouch. No IXth tergite observed. Ante-anal setae ; 2 long h.s. Posterior margin with 3 h.s. on each side.
Genital segment. Penial sheath short, about } total body length (ratio 1 : 5-49-6-00, average 5°86), 327-357 (average 342) uw long and 56-63 (average 60) » wide ; lateral sclerotizations not joined anterior to anus ; length of basal vod about half that of aedeagus, the rod extending anteriorly from base of aedeagus to the apex of the basal membranous area ; apex of sheath without membranous extension. The area from base of sheath to tip of aedeagus with 28-42 (average 33) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath. Aedeagus long (160-186, average 174 uw), penial sheath and basisternum longer, the ratios being I : 1:8-2-1 (average 2) and 1 : 1:23-1-39 (average 1-34) respectively.
Material examined : 10 specimens, collected by J. M. Cherret on Hedera helix L. (ivy) in Bangor, Wales during May, 1962. Two specimens collected by N. S. Borchsenius in the Crimea, USSR on 24.v.54 agreed well with the above description, but were somewhat smaller, measuring 1680 and 1720 uw. The same is true for 4 specimens collected by J. Rehdéek on Hedera helix L. during May, 1953 in Czecho- slovakia ; they measured 1660, 1680, 1760 and 1830 u.
CTENOCHITON Ctenochiton sp. (Text-figs. 16 and 17)
A moderately small and slender species, with comparatively long antennae and moderately long legs ; with numerous setae on the appendages, but few on the body itself. When mounted, total body length 1601-1929 (average 1754) u ; width at mesothorax 240-280 (average 258) u. Wing expanse 3052-3265 (average 3178) u.
Head subconical in dorsal view ; in lateral view obliquely dorsoventrally elongated, with anterodorsal bulge not pronounced ; length from apex to pronotal ridge 220-258 (average 237) u, width across genae 240-280 (average 256) u. Median crest sclerotized and striated posteriorly, but not polygonally reticulated ; with 9-15 (average 12) hair-like dorsal head setae. Muid- cranial ridge dorsally absent ; ventrally narrow but well defined, reaching postocular ridge posteriorly, surrounding area not reticulated. Genae large, sclerotized ; polygonal reticulation enclosing weaker, irregular reticulation ; without setae. Eyes: four pairs; dorsal and
ventral pairs large, subequal ; lateral pairs smaller, subequal ; corneae of dorsal eyes 19-27
64 MORPHOLOGY AND TAXONOMY OF ADULT MALES
(average 23) uw in diameter and 3-1—5-8 (average 4-6) times as much apart ; those of the ventral eyes 20-27 (average 23) u in diameter and 1-2-1-6 (average 1-5) times as much apart. Ocellus small. Ocular sclerite well sclerotized and polygonally reticulated throughout. Pvreoculay ridge with ventral part reaching or almost reaching midcranial ridge. Postoculay ridge thin and tapering dorsally, well developed lateroventrally, and narrow but well defined posteromedially ; below ocellus the ridge splits up, with anterior branch partly surrounding ocellus. Intevoculay vidge absent. Dorsal oculay setae : 1-4 (average 2:2) h.s. on each side ; ventral head setae : 3-5 (average 4°5) h.s., 2 of which are usually situated anterior to preocular ridge and 2-3 posterior to it. Pvreoral ridge present. Tendon-like apodeme long. Cranial apophysis short ; apex bifurcate with a central lobe, not reaching level of posterior margin of ventral eyes. Mouth opening irregular. Anterior tentorial pits absent.
Antennae to-segmented, filiform ; 1024-1144 (average 1070) yu long, i.e. longer than half body length (ratio I : 1:51-1:69, average 1-63), longer than posterior leg (ratio I : 1-01I—1-05, average 1-04) and longer than penial sheath (ratio 1 : 3:30-3:69, average 3:48). Scape 52-60 (average 58) uw long and 40-48 (average 46) uw wide, with 3 h.s. Pedicel with weak polygonal, dorsal reticulation ; 44-52 (average 46) u long and 40-48 (average 45) u wide ; with 8-14 (average 10) f.s., 4-8 (average 6-3) h.s. and a sensillum placodeum. Segment III bulging in the middle, 2:0-2°4 (average 2-2) times longer than wide (52-68, average 61 u long and 26-32, average 29 uw wide) ; with 2-4 (average 3-1) h.s. and 6-10 (average 8-3) f.s., the latter of medium length, I-I-I-4 (average 1-3) times as long as width of segment ; with 2-4 usual sensilla basiconica. Segments IV-I1X cylindrical ; lengths of these segments (in w) 152-180 (average 166), 148-164 (average 155), 148-180 (average 166), 124-164 (average 147), Ioo-124 (average 111) and 72-94
Tic. 16. Ctenochiton sp., dorsal and ventral view.
OF THE FAMILY COCCIDAE 65
(average 80) respectively, all of about the same width, varying from 19 to 25 pu ; with 27-38 (average 33), 26-32 (average 28), 26-36 (average 30), 21-33 (average 27), 18-26 (average 22) and 11-21 (average 17) f.s., and 1-4 (average 2-3), I-4 (average 2-2), I-3 (average 2-3), I-3 (average 2-1), 0-2 (average 0-6) and o~2 (average 1) h.s. respectively ; antennal bristles on segments VIII-IX distinctly larger than fleshy setae. Segment X : terminal part not con- stricted ; 76-84 (average 80) u long and 22-26 (average 24) uw wide ; carrying 6-12 (average Io) f.s., 3 capitate subapical setae and 5 antennal bristles of which the 3 long ones are about half as long as the segment and the 2 shorter ones somewhat shorter than most fleshy setae, but distinctly thicker ; with 2 sensilla basiconica ventrally, one near apex and the other more proximal.
Thorax 584-664 (average 619) p long.
Prothorax. Pyronotal ridge strong, but medially interrupted by weak sclerotization. Lateral pronotal sclerites small, without seate. Medial pronotal setae absent ; 3-7 (average 6-1) circular pores present on each side posterior to pronotal sclerite. Post-tergites small, without striations, and without setae. Pleural structures typical of family. Steynum with a strong transverse ridge, a narrow but uninterrupted median ridge and a triangular sclerite. Anteprosternal setae absent ; prosternal setae : occasionally one h.s.
Mesothovax. Mesoprephragma with shallow emargination. Pyrescutum about twice as wide as long (average 201 and 98 p respectively) ; anterior margin strongly curved ; laterally bounded by the prescutal ridges and posteriorly by the prescutal suture ; weakly, irregularly reticulated ; slightly more heavily sclerotized medially. Scutwm. Median membranous area subrectangular; 108-128 (average 116) u long and 1-28—1-78 (average 1-49) times as wide (width 152-192 average 173 w) ; with 7-13 (average 10) h.s. Scutellum 61-87 (average 72) u long and 160—192 (average 175) w wide, the ratio being 1 : 2:2—2-8 (average 2-4) ; tubular, with ventral foramen of medium size ; without setae. Postnotum with anterior margin usually regular and exposed ; postnotal apophysis and postalare well developed, the latter densely reticulated distally. Mesopost- phragma with shallow emargination. Mesopleuron. Mesopleural ridge strong, but interrupted above coxal articulation ; pleural apophysis and pleural wing process well developed, the latter connected with episternum by a narrow basalare. Subalare small. Episternum showing weak, irregular reticulation ; subepisternal ridge well developed, but below membranous cleft indistinct and only marked by a band of dark sclerotization. Epimeron small. Lateropleurite bounded anteriorly by an extension from marginal ridge. Basisternuwm large, about 253 w wide and 207 u long, i.e. 1-60-1-89 (average 1-74) times longer than membranous area of scutum ; with strong mediun ridge and bounded by strong marginal and precoxal ridges ; without setae. Furca well developed. Mesothoracic spiracle with well developed peritreme ; postmesospivaculay setae absent. Tegula small, membranous bulge with a small weak sclerite posteriorly and 4-7 (average 5) h.s. Third avillary wing sclerite with a pronounced ventral projection at its base. Additional sclerite small, but well defined. Antemetaspivacular setae absent.
Metathovax. Metanotum with anterior margin usually strong medially ; suspensorial sclevites small, spot-like ; a small, additional sclerite usually present anterior to postnotum. Postnotum consisting of a transverse sclerite on each side. Metatergal setae : one h.s. on each side, 3-11 (average 6-3) circular pores occurring near each seta. Pleural ridge well developed,
Fic. 17. Ctenochiton sp., lateral view.
66 MORPHOLOGY AND TAXONOMY OF ADULT MALES
though interrupted in middle ; with small wing process. Episternum with anterior margin net ridge-like ; vestigial precoxal vidge present ; epimervon posteriorly directed. Metathoracic spivacle similar to mesothoracic one. Dorsospivacular setae absent. Postmetaspivacular setae ? 2-6 (average 3:5) f.s. and 1-7 (average 4) h.s. Metasteynum represented by a weak, transverse plate. Anterior metasternal setae : 1-3 (average 1-9) medially situated h.s. ; posterior meta- steynal setae : occasionally one h.s.
Wings hyaline ; long (length 1357-1463, average 1427 uw) and of medium width (530-615, average 578 u), ratio width to length being 1 : 2:36-2:56 (average 2-46) ; alar lobe present ; alay setae : one or occasionally twoh.s. oneach wing. Halteves well developed, 104-120 (average 111) w long and 20-28 (average 24) w wide, each with one apically hooked seta which is about 64 uw long.
Legs moderately long, and slender, with middle or fore pairs shortest and hind pair longest ; ratio length of hind leg to body length is 1 : 1:68-1:73 (average 1-71). Length of segments (in w) :
Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 68-76 92-106 198-222 359-422 110-128 23-26 903-1013 (71) (100) (211) (391) (119) (24) (964)
II 72-84 92-106 178-207 331-414 115-133 24-27 859-IOIO (81) (99) (193) (377) (124) (25) (947)
III 80-91 95-116 190-218 361-426 112-135 24-27 906-1053 (85) (104) (204) (393) (125) (25) (985)
Coxae each with 13-25 f.s. and 13-19 h.s. ; fore coxa with 2-5 (average 3-8) pointed coxal bristles, the latter about $4 as long as the segment ; apical seta about } as long as trochanter. Trochanters 26-30 w wide ; with 6 oval sensilla, 10-16 f.s. and 7-10 h.s., the latter including 2 minute setae near basal ridge, one small seta on outer margin and a long apical seta which, on the fore trochanter, is I-4—-1-7 (average 1-5) times as long as width of trochanter. Femora of medium width (38-42 uw), the ratio width to length of hind femur being I : 4-7—-5-4 (average 5:1) ; each with 22-34 f.s. and 13-26 h.s. Tuibiae 19-25 uw wide, the ratio width to length of hind tibia being 1 : 16-7—20-0 (average 18-4) ; each with 79-102 setae, of which 30-48 are h:s. and 44-60 f.s., the latter about as long as width of tibia ; apical spur about the same size on all tibiae. Tarsi 18-21 (average 19) w wide, hind tarsus 5-7~7:0 (average 6-3) times longer than wide ; with 9-15 f.s. and 19-32 h.s. ; tarsal digitules subequal, about as long as claw. Claws of medium length, somewhat longer than width of tarsus ; slightly curved, with a small denticle near tip ; wngual digitules subequal, about as long as claw.
Abdomen 551-721 (average 613) uw long and 371-477 (average 411) u wide.
Segments I-VII : tergites represented by a small sclerite on each side on anterior margin of segments II-III and a weak transverse plate on VII; steynites present on all segments, represented by a weak transverse plate on anterior and posterior segments and a sclerite on each side on intermediate segments. Caudal extension of segment VII small, somewhat pointed, not sclerotized. Dorsal setae: h.s. only, usually one on each side on segments I, and III-VII ; 1-7 (average 4-1) circular pores occur near each seta on segment I. Pleural setae consisting of h.s. only ; dorsopleural setae : 2-5 (average 3-3), 3-5 (average 4-1), 3-6 (average 4:6) and 3-5 (average 4:3) on segments III—VI respectively ; ventropleural setae absent or incorporated into the dorsopleural group. Segment VII with 5—8 (average 6) h.s., of which the posterior ones are usually longer than the rest. Ventral setae : h.s. only, usually one on each side on III-IV, and 4 on each of segments V—VII.
Segment VIII with transverse tergite and sternite ; caudal extension forming a small, simple lobe ; glandular pouch with 2 long setae, each with a small apical knob, the protruding part of these setae 3-4 times as long as section within pouch. No IXth tergite observed. Ante-anal setae : 2 long and occasionally one small h.s. ; 9-17 (average 14) circular pores present anterior to ante-anal setae. Posterior margin with 3-4 (average 3-3) h.s. on each side.
Genital segment. Penial sheath short, about } total body length (ratio 1 : 5:20-6:12, average 5°75), 285-323 (average 304) p long and 42-49 (average 46) uw wide ; lateral sclerotizations
OF THE FAMILY COCCIDAE 67
narrowly joined anterior to anus ; length of basal rod %-¢ as long as aedeagus and extending anteriorly from the base of aedeagus for about % of the length to the apex of the basal mem- branous area ; apex of sheath without membranous extension. The area from base of sheath to tip of aedeagus with 17-26 (average 20) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath. Aedeagus of medium length (101-120, average 114 yu), penial sheath and basisternum longer, the ratios being 1 : 2:6—3-0 (average 2-7) and I : 1-74-1-92 (average 1-85) respectively.
Material examined : 10 specimens, collected by myself on Antizoma capensis (Thunb.) in Stellenbosch, South Africa during August, 1961 ; identified by G. De Lotto.
ERICERUS Ericerus pela (Chavannes) (Text-figs. 18 and 19)
A very large and moderately robust species, with comparatively long antennae and legs ; with numerous long setae on the appendages, but few on the body itself. When mounted, total body length 2500-3100 (average 2864) » ; width at mesothorax 570-740 (average 680) u. Wing expanse 5330-5700 (average 5563) wu.
Fic. 18. Evicerus pela (Chav.), dorsal and ventral view.
68 MORPHOLOGY AND TAXONOMY OF ADULT MALES
Head subconical in dorsal view ; in lateral view dorsoventrally elongated ; long and narrow, length from apex to pronotal ridge 357-456 (average 420) w and width across genae 285-342 (average 315) uw. Median crest sclerotized and densely polygonally reticulated ; with 5-7 (average 5:7) hair-like dorsal head setae anteriorly. Muidcyranial ridge dorsally absent ; ventrally narrow but well defined, reaching postocular sclerite posteriorly, surrounding area polygonally reticulated. Genae large, sclerotized ; polygonal reticulation enclosing weaker, irregular reticulation ; without setae. Eyes: five pairs ; dorsal and ventral pairs large, subequal ; lateral pairs smaller, subequal ; corneae of dorsal eyes 40-47 (average 43) u in diameter and I-4-2:I (average 1-7) times as much apart, those of the ventral eyes 42-49 (average 46) u in diameter and o-9-1-1 (average 1) times as much apart. Ocellus small. Ocular sclerite well sclerotized and densely polygonally reticulated throughout. Preoculay ridge with ventral part reaching or almost reaching midcranial ridge. Postoculay ridge weak dorsally, well developed lateroventrally, and narrow but well defined posteromedially ; below ocellus the ridge splits up, with anterior branch partly surrounding ocellus. Jnteroculay vidge absent. Dorsal ocular setae absent ; ventval head setae : 3—5 (average 4-4) h.s., situated anterior to ocular sclerite on or near midcranial ridge. Pyveoval ridge present. Tendon-like apodeme long. Cranial apophysis short ; apex truncate, not reaching level of anterior margin of ventral eyes. Mouth opening irregular. Anterior tentorial pits situated anterolateral to mouth opening.
Antennae 10-segmented, filiform ; 1691-2120 (average 1922) u long, i.e. longer than half body length (ratio 1 : 1-46-1-55, average 1-50), longer than posterior leg (ratio I : 1-I0-1-17, average 1-13) and longer than penial sheath (ratio I : 3:50-4:04, average 3°89). Scape 80-99 (average 93) p long and 76-95 (average 86) u wide, with 3 h.s. and 2-4 (average 2-6) f.s. Pedicel with distinct, polygonal, dorsal reticulation ; small, 61-84 (average 73) uw long and 57-68 (average 64) » wide ; with 1-3 (average 2:1) f.s., 2-4 (average 2-6) h.s. and a sensillum placo- deum. Segment III long, cylindrical, 4-6-6-3 (average 5-3) times longer than wide (209-258, average 228 uw long and 38-48, average 43 uw wide) ; with 6-11 (average 8-3) h.s. and 15-26 (average 21) f.s., the latter, as on the other antennal segments, very long, 4-5 (average 4°5) times as long as width of segment III ; with 1-3 usual sensilla basiconica. Segments 1V-IX cylindrical ; lengths of these segments (in uw) 213-274 (average 248), 235-281 (average 264), 232-312 (average 277), 194-274 (average 235), 156-205 (average 186) and 118-156 (average 145) respectively, all of about the same width, varying from 27 to 38 u ; with 15-28 (average 21), 17-26 (average 23), 18-31 (average 24), 18-26 (average 22), 12-18 (average 15) and 9-14 (average 12) f.s. respectively, but no h.s. ; antennal bristles on segments VIII—-IX distinctly thicker than f.s. ; one or two f.s. near apex of each segment short, shorter than width of segment. Segment X : terminal part not constricted ; 125-213 (average 172) uw long, and 29-34 (average 31) w wide ; carrying 6-11 (average 8-2) f.s., 3 capitate subapical setae and 5 antennal bristles of which the 3 long ones are about ? as long as the segment and the 2 short ones thin and shorter than the f.s. (one shorter than the other, about as long as width of segment) ; with 2 sensilla basiconica ventrally, one near apex and the other more proximal.
Thovax 832-1113 (average 1015) u long.
Fic. 19. Evicerus pela (Chav.), lateral view.
OF THE FAMILY COGCIDAE 69
Prothovax. Pronotal ridge strong, but narrowly constricted medially. Lateral pronotal sclerites small, without setae. Medial pronotal setae absent. Post-tergites elongated, with wavy striations and without setae. Pleural structures typical of the family. Steynwm with trans- verse ridge strong, median ridge reduced to a short basal stalk and triangular sclerite well sclerotized. Anteprosternal setae absent ; prosternal setae : o-4 (average 1-8) hs.
Mesothorax. Mesoprephragma with shallow emargination. Pyrescutuwm less than twice as wide as long (average 287 and 185 u respectively) ; anterior margin curved ; laterally bounded by the prescutal ridges and posteriorly by the prescutal suture ; with weak, irregular reticula- tion ; slightly more heavily sclerotized medially. Scutuwm. Median membranous area trapezoidal ; 108-186 (average 156) uw long and 1-57—2-40 (average 1-95) times as wide (width 247-334, average 298 uw) ; without setae. Scwutellum large, 76-114 (average 96) p» long and 209-293 (average 250) w wide, ratio being 1 : 1-4-1°8 (average 1-6) ; not tubular ; without setae. Postnotum with anterior margin irregular and partly overlapped by metathoracic fold ; postnotal apophysis and postalare well developed, the latter densely reticulated distally. _Meso- postphragma with deep emargination. Mesopleuron. Mesopleural ridge strong, but interrupted above coxal articulation ; plewral apophysis and pleural wing process well developed, the latter connected with episternum by a narrow basalare. Subalare small. Episternum not reticulated ; subepisternal ridge well developed, but below membranous cleft indistinct and only marked by a band of dark sclerotization. Epimeron small. Lateropleurite partly bounded anteriorly by an extension from marginal ridge. Basisternum large, about 430 u wide and 356 yp long, i.e. 2-o8—2-81 (average 2-31) times longer than membranous area of scutum; with strong median ridge and bounded by strong marginal and precoxal ridges ; without setae. Furca well developed. Mesothoracic spiracle with well developed peritreme; postmesospivacular setae absent. Tegula small, membranous bulge with a weak sclerite posteriorly and 2-7 (average 4:7) h.s. Third axillary wing sclerite with a pronounced ventral projection at its base. Additional sclerite elongated, well defined. Antemetaspiraculay setae absent.
Metathovax. Metanotum with anterior margin usually strong medially ; suspensorial sclerites small, spot-like ; a small, additional sclerite present anterior to postnotum. Post- notum consisting of a narrow, transverse sclerite on each side. Metatergal setae absent. Pleural ridge well developed, though interrupted in middle ; with small wing process. Epi- sternum with anterior margin not ridge-like ; pyrecoxal ridge absent ; epimeron produced posteriorly. Metathoracic spivacle similar to mesothoracic one. Dorsospivaculay setae absent. Postmetaspivaculay setae : o-3 (average 1) f.s. and 4-8 (average 5-6) h.s. ; of the latter 3~7 are situated dorsally and one or two more ventrally. Metasternwm represented by a weak trans- verse plate. Anterior and posterior metasternal setae absent.
Wings hyaline ; long (length 2350-2530, average 2490 uw) and of medium width (890-950, average 9II uw), ratio width to length being 1 : 2:64-2:77 (average 2-68) ; alar lobe present ; alay setae absent. Halteres well developed, 163-228 (average 202) yp long and 42-61 (average 52) w wide, each with 1-4 (average 2-3) apically hooked setae which are about 97 yu long.
Legs long and slender, with middle pair shortest and hind pair longest ; ratio length of hind leg to body is 1 : 1-54-1-73 (average 1:65). Length of segments (in y) :
Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 106-137 114-186 334-426 562-760 194-266 38-48 1379-1804 (119) (t71) (389) (678) (242) (45) (1643)
Hin 106-141 114-186 308-388 543-749 179-251 40-49 1321-1763 (124) (164) (353) (669) (227) (46) (1584)
UU 114-156 160-198 334-419 608-798 188-304 40-49 1444-1908 (136) (176) (384) (734) (249) (47) (1726)
Cowxae with 14-22 f.s. on fore and middle coxa and 17—32 (average 25) on the hind coxa, and with 12-23 (average 18) h.s. on fore coxa and 8-15 on the middle and hind coxa ; fore coxa without coxal bristles ; apical setae about half as long as the trochanter. Tvochanters 42-68 u wide, with 6-8 oval sensilla, 7-12 f.s. and 7-9 h.s., the latter including 2 minute setae near basal
70 MORPHOLOGY AND TAXONOMY OF ADULT MALES
ridge, one small seta on the outer margin and a long apical seta which, on the fore trochanter, is I-5—I-9 (average 1-7) times as long as width of trochanter. Femorva of medium width (60-87 y), ratio width to length of hind femur being 1 : 4-7-5:5 (average 5-1) ; each with 25-43 f.s., with 16-21 (average 18) h.s. on the fore and 10-16 on the middle and hind legs. Tibiae 34-49 u wide, the ratio width to length of hind tibia being I : 16-2-18-2 (average 17-1) ; each with 72-99 setae of which 17—29 are h.s. and 53-72 f.s. ; the latter, as on the other segments, very long, about 4—5 times as long as width of tibia ; with a number of spurs near apex, apical spur about the same size on all tibiae. Tavsi 29-38 uw wide, hind tarsus 6-2—8-9 (average 7-3) times longer than wide ; each with 12-19 f.s. and 8-15 hs. ; tarsal digitules subequal, longer than claw. Claws of medium length, somewhat longer than width of tarsus ; slightly curved, with small denticle near tip ; ungual digitules subequal, about as long as claw.
Abdomen 800-1060 (average 982) uw long and 600-780 (average 681) wu wide.
Segments I-VII : tergites on segments II-III represented by a small sclerite on each side, and on VII by a weak transverse plate ; stevmites on segments V—VII represented by a weak transverse plate. Caudal extension of segment VII small, somewhat pointed, not sclerotized. Dorsal setae : h.s. only, sometimes one on each side on segment IV and usually so on V—VII. Pleural setae consisting of h.s. only, which include dorsopleural setae : 1, 1-2 (average 1-2), I-3 (average 1-7) and 3 on segments III—VI respectively, and ventvopleural setae: usually one on each of segments IV-VI. Segment VII with 4 hs. Ventral setae : h.s. only, usually one on each side on each of segments II-IV, and 3-5 on V—VII.
Segment VIII with transverse tergite and sternite ; caudal extension forming a small, simple lobe ; glandular pouch with 2 long setae, each with a small apical knob, the protruding part of these setae 1-1} times as long as section within pouch. NoIXthtergite observed. Ante-anal setae : 2 long h.s. Posterior margin with 3 h.s. on each side.
Genital segment. Penial sheath short, about } total body length (ratio 1 : 5:26-6:20, average 5:81), 441-536 (average 492) »w long and 61-68 (average 65) uw wide ; lateral sclerotizations not joined anterior to anus ; length of basal vod about equal that of aedeagus, extending anteriorly from base of aedeagus to apex of the basal membranous area ; apex of sheath without mem- branous extension. The area from base of sheath to tip of aedeagus with 26-41 (average 34) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath. Aedeagus short (length 137-171, average 162 yu), penial sheath and basisternum longer, ratios being I : 2:8-3-2 (average 3) and 1 : 1-81—2-44 (average 2:17) respectively.
Material examined : Io specimens, collected by N. S. Borchsenius on Ligustrum sp. in China on 14.1x.54.
Genus A sp. (Text-figs. 20 and 21)
A medium-sized, robust species with comparatively long antennae and moderately long legs ; with numerous short, thick setae on the appendages but with only a few h.s. on the body itself. When mounted, total body length 1630-1830 (average 1757) uw ; width at mesothorax 455-500 (average 477) uw. Wing expanse 3140-3220 (average 3172) u.
Head subconical in dorsal view ; rounded in lateral view, with anterodorsal bulge not pro- nounced ; length from apex to pronotal ridge 213-236 (average 218) uw, width across genae 277-304 (average 286) uw. Median crest well sclerotized, especially anteriorly and around the posterior margin ; central part polygonally reticulated ; with 7-10 (average 8-5) hair-like dorsal head setae. Midcvanial ridge dorsally absent ; ventrally narrow but well defined, reaching postocular sclerite posteriorly, surrounding area sometimes weakly sclerotized near ridge, but not reticulated. Genae large, sclerotized, not reticulated, without setae. Eyes : four pairs ; dorsal and ventral pairs large, subequal ; lateral pairs smaller, subequal ; corneae of dorsal eyes 21-23 (average 22) wu in diameter and 6:8-7-8 (average 7-2) times as much apart ;
OF THE FAMILY COCCIDAE 71
those of the ventral eyes 19-23 (average 21) uw in diameter and 2-8—3-3 (average 3-1) times as much apart. Ocellus small. Ocular sclerite well sclerotized and polygonally reticulated throughout. Pyreoculay ridge with ventral part reaching midcranial ridge. Postoculay ridge weak dorsally, well developed lateroventrally and narrow but well defined posteromedially ; below ocellus the ridge splits up, with the posterior part weak or lost entirely and the anterior part strong, extending anterior to ocellus. Jntevoculay ridge usually present, narrow, connecting pre- and postocular ridges between lateral eyes (ridge sometimes absent on one side and occasionally on both sides). Dorsal ocular setae absent ; ventral head setae : 3-5 (average 4) h.s., situated on or just anterior to preocular ridge, a pair of these being considerably longer than the others. Preoral ridge absent. Tendon-like apodeme long, arising from the anterior part of an elongated ventral sclerite. Cranial apophysis very short, apex truncate with a large, irregular central lobe. Mouth opening irregular. Anterior tentorial pits situated anterolateral to mouth opening.
Antennae 1o-segmented, filiform ; 1003-1144 (average 1101) yu long, i.e. longer than half body length (ratio 1 : 1-62—1-74, average 1-68), longer than posterior leg (ratio I : 1-16—1-19, average 1-17) and longer than penial sheath (1 : 3-06—3-40, average 3:18). Scape 61-72 (average 66) uw long and 57-63 (average 60) w wide, with 3-6 (average 4-1) h.s. Pedicel with distinct, poly- gonal, dorsal reticulation ; 57-68 (average 64) uw long and 53-63 (average 59) uw wide ; with 7-14 (average 10) f.s., 8-15 (average 11) h.s. and a sensillum placodeum. Segment III bulging in middle, 1-7—2-1 (average 1-9) times longer than wide (76-95, average 84 u long and 38-46, average 43 uw wide) ; with 6-8 (average 7) h.s. and 21-31 (average 24) f.s., the latter, as on the other antennal segments, very short, about 0-4—0-6 (average 0-5) times as long as width of segment III ; with 1-3 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths of
Fic. 20. Genus A, dorsal and ventral view.
72. MORPHOLOGY AND TAXONOMY OF ADULT MALES
these segments (in w) 137-164 (average 149), I41I-175 (average 155), 156-175 (average 165), 118-148 (average 135), 95-122 (average 113) and 76-91 (average 85) respectively, all of about the same width, varying from 29 to 38 uw ; with 35-48 (average 42), 40-51 (average 47), 46-53 (average 48), 36-48 (average 42), 30-41 (average 33) and 20-28 (average 25) f.s., and 5-10 (average 8), 5-8 (average 6-7) 5-12 (average 7-7), 4-6 (average 4-6), 3-5 (average 4) and o-4 (average 2:3) h.s. respectively ; antennal bristles on segments VIII-IX not distinctly different from the f.s. Segment X : terminal part not constricted ; 84-91 (average 86) w long and 57-68 (average 63) w wide ; carrying 5—10 (average 7-3) f.s., o—-2 (average 1-5) h.s., 4-6 (average 5) capitate subapical setae and 5 antennal bristles of which the 3 long ones are less than half as long as the segment and the 2 short ones about as long as the f.s. though thinner ; with 2 sensilla basiconica ventrally, one near apex and the other more proximal.
Thorax 593-665 (average 637) u long.
Pyrothovax. Pyonotal vidges strong, but medially interrupted by weak sclerotization. Lateral pronotal sclevites small, without setae. Medial pronotal setae absent. Post-tergites small, without striations and without setae. Pleural styuctuves typical of family. Steynum with strong transverse ridge, a median ridge which is occasionally interrupted in the middle, and a well sclerotized triangular sclerite. Anteprosternal setae absent ; pyrosternal setae : occasionally one h.s. on each side.
Mesothovax. Mesoprephragma with shallow emargination. Prescutum more than twice as wide as long (average 228 and 90 p respectively) ; anterior margin curved ; laterally bounded by the prescutal ridges and posteriorly by the prescutal suture ; with polygonal reticulation ; slightly more heavily sclerotized medially. Scutum. Median membranous area subrectangular; 96-114 (average 100) u long and 2:00-2:82 (average 2-38) times as wide (width 205-236, average 221 uw) ; with 9-13 (average 12) h.s., usually with one on each side situated posterolaterally immediately outside the membranous area. Scutellwm large, 110-125 (average 118) uw long and 201-235 (average 220) pu wide, the ratio being 1 : 1-8—2-1 (average 1:9) ; not tubular; with o-2 (average 1-6) h.s. Postnotum with anterior margin irregular and partly overlapped by metathoracic fold ; postnotal apophysis and postalave well developed, the latter densely reticulated distally. Mesopostphragma with deep emargination. Mesopleuron. Mesopleural vidge strong, but interrupted above coxal articulation ; pleural apophysis and pleural wing process well developed, the latter connected with episternum by a narrow basalave. Subalare small. Episteynum not reticulated ; subepisternal ridge well developed but below membranous cleft indistinct and only marked by a band of dark sclerotization. Epimevon small. Latero- pleurite partly bounded anteriorly by an extension from marginal ridge. Basisteynum large, about 262 » wide and 183 yu long, i.e. 1:57-2:49 (average 1:92) times longer than membranous area of scutum ; with strong median ridge and bounded by strong marginal and precoxal ridges ; without setae. Furca well developed. Mesothovacic spivacle with well developed peritreme ; postmesospivaculay setae absent. Tegula small, membranous bulge with a weak sclerite posteriorly and 7-11 (average 8-8) h.s. Third axillary wing sclerite with a large ventral projection at its base. Additional sclerite elongated, well defined. Antemetaspivacular setae absent.
Fic. 21. Genus 4A, lateral view.
OF THE FAMILY COCCIDAE 73
Metathorax. Metanotum with anterior margin usually strong medially ; suspensorial sclerites small, spot-like ; a small, additional sclerite occasionally present anterior to post- notum. Postnotum consisting of a transverse sclerite, which is narrow medially but usually not interrupted. Metatergal setae : one or two h.s. on each side. Pleural ridge well developed, but usually interrupted in the middle ; with a small wing process. Precoxal vidge absent. Epi- sternum ventrally interrupted so that a small, separate sclerite is formed ; anterior margin not ridge-like. Epimervon produced posteriorly. Metathoracic spivacle similar to mesothoracic one. Dorsospiracular setae absent. Postmetaspivacular setae : occasionally one or two h.s. Meta- sternum represented by a transverse plate. Anterior and posterior metasternal setae : usually one median h.s. each.
Wings hyaline ; of medium length (1370-1420, average 1394 wu) and broad (610-655, average 630 yw), the ratio width to length being 1 : 2-17—2-26 (average 2-21) ; alar lobe present ; alar setae absent. Halteres well developed, 125-144, (average 136) u long and 30-36 (average 34) u wide, each with one apically hooked seta which is about 70 pu long.
Legs moderately long and slender, with fore pair shortest and the middle or hind pair longest ; ratio length of hind leg to body length is 1 : 1-82—2-o1 (average 1-89). Length of segments (in y) :
Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 76-87 87-103 209-224 338-384 QI-103 25-27 842-920 (82) (99) (217) (364) (100) (26) (888)
Il 76-91 93-106 191-231 376-440 103-110 25-29 865-954 (87) (102) (209) (395) (108) (27) (928)
Ill 76-91 QI-I14 192-227 357-418 108-112 25-27 859-969 (87) (106) (215) (390) (110) (26) (927)
Coxae with 10-13 (average 11), 11-19 (average 15) and 14-25 (average 18) f.s. on the fore, middle and hind coxa respectively, and with 11-21 h.s. each ; fore coxa with 2—4 (average 3) coxal bristles ; apical seta about half as long as trochanter. Tyvochantervs 30-40 uw wide ; with 6 oval sensilla, 5—10 f.s. and 6-11 h.s., the latter including 2 minute setae near basal ridge, one small seta on outer margin and a long apical seta which, on the fore trochanter, is 1-2—1°5 (average 1-4) times as long as width of trochanter. Femora very broad (46-65 yp), ratio width to length of hind femur being 1 : 3:2—3-8 (average 3-5) ; each with 32-54 f.s. and 15-31 hs. Tibiae 30-34 w wide, ratio width to length of hind tibia being 1 : 10-4-12-2 (average 11-5) Setae : 83-135, of which 35~—59 are h.s. and 48-81 f.s., the latter, as on the other segments, very short and thick, about % as long as width of tibia ; apical spur about the same size on all tibiae. Tarsi 25-30 w wide, hind tarsus 3:6—4-1 (average 3-7) times longer than wide ; each with 21-40 h.s., with o—5 (average 1-8) f.s. on the fore and none on the middle and hind tarsi ; tarsal digitules subequal, longer than claw. Claws of medium length, about as long as width of tarsus ; slightly curved, with small denticle near tip ; ungual digitules subequal, about as long as claw. Abdomen 540-620 (average 577) » long and 430-510 (average 459) u wide.
Segments I-VII : tergites represented by a transverse sclerite on anterior margin of segments II, III and sometimes IV ; sternites represented by a weak transverse plate on segments II, III and V-VII. Caudal extension of segment VII small, somewhat pointed, not sclerotized. Dorsal setae : h.s. only, usually one on each segment. Pleural setae : consisting of h.s. only, which include dorsoplewral setae : o-2 (average 1:6), 1-4 (average 2-1), 2-4 (average 3-3) and o-4 (average 2-9) on segments III—VI respectively, and ventropleural setae : sometimes one on each of segments IV-VI. Segment VII with 3-9 (average 5-4) h.s. Ventral setae : h.s. only, usually one on each side on each of segments III-V, 4 on VI and 4-6 (average (5-6) on VIL ; two of the median setae on segment VII considerably longer than rest.
Segment VIII with small tergite and transverse sternite ; glandular pouch with 2 long setae, each with a small apical knob, the protruding part of these setae 2-2} times as long as section within pouch. NoIXthtergite observed. Ante-anal setaeabsent. Posterior margin with 3 h:s. on each side.
74 MORPHOLOGY AND TAXONOMY OF ADULT MALES
Genital segment. Penal sheath of medium length, about + total body length (ratio I : 4-80— 5°32, average 5:04), 315-369 (average 349) uw long and 53-61 (average 57) uw wide ; lateral sclerotizations narrowly joined anterior to anus ; length of basal vod about 32 to equal that of aedeagus and extending anteriorly from base of the latter to apex of the basal membranous area ; apex of sheath without membranous extension. Area from base of sheath to tip of aedeagus with 36-46 (average 40) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath. Aedeagus long (125-148, average 140 uw) and broad, penial sheath and basisternum longer, ratios being I : 2-4—2-6 (average 2:5) and I : 1:21-1-42 (average 1-31) respectively.
Material examined : 7 specimens, collected by C. J. Joubert on “ katdoring ”’ (Asparagus sp.) during August, 1956 in Stanford, South Africa; received from J. G. Theron.
SPHAEROLECANIUM Sphaerolecanium prunastri (Fonscolombe)
(Text-figs. 22 and 23)
A medium-sized, robust species with comparatively short antennae and legs ; with many setae on the appendages, but few on the body itself. When mounted, total body length 1640-1810 (average 1739) uw ; width at mesothorax 360-430 (average 408) up. Wing expanse 2750-2940 (average 2816) u.
Fic. 22. Sphaevolecanium prunastri (Fonsc.), dorsal and ventral view.
OF THE FAMILY COCCIDAE 75
Head subconical in dorsal view ; in lateral view obliquely elongated dorsoventrally, with, anterodorsal bulge somewhat pronounced ; length from apex to pronotal ridge 205-247 (average 222) p, width across genae 243-266 (average 253) u. Median crest sclerotized and distinctly polygonally reticulated ; with 9-12 (average ro) hair-like dorsal head setae, arranged in a group of 3-5 near anterior margin of head and a group of 5-7 very short setae more posteriorly. Midcranial ridge dorsally absent ; ventrally narrow but well defined, reaching ocular sclerite posteriorly, surrounding area with polygonal reticulation. Genae large, sclerotized, polygonally reticulated, without setae. Eyes: two pairs; subequal; corneae of dorsal eyes 19-27 (average 25) up in diameter and 3-4~—4-2 (average 3-9) times as much apart ; those of the ventral eyes 19-27 (average 26) » in diameter and 1-2—1-6 (average 1-3) times as much apart. Ocellus small, bulging. Ocular sclerite well sclerotized and polygonally reticulated throughout. Pre- ocular ridge short, not extending far below articular process. Postoculay ridge well developed dorsally and lateroventrally, but weak posteromedially ; below ocellus the ridge splits up, with anterior branch partly surrounding ocellus. Jnteroculary vidge absent, but sometimes indicated by a line of dark sclerotization. Dorsal ocular setae absent; ventral head setae : 4-6 (average 5) h.s., situated on the anterior margin or just anterior to the ocular sclerite