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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

OF

“MOIA [PIJUDA pure [esIop

‘(¥

1)

avyy wnimnoayny

“OI

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
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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 ;
with one pair of setae distinctly longer than others. Preoval ridge present. Tendon-like
apodeme long. Cranial apophysis of medium length ; apex bifurcate with a central lobe, not
reaching level of anterior margin of ventral eyes. Mouth opening irregular. Anterior tentorial
pits apparently present anterolateral to mouth opening.

Antennae 1o-segmented, filiform ; 671-809 (average 770) w long, i.e. shorter than half body
length (ratio 1 : 2-13—2°45, average 2-28), longer than posterior leg (ratio 1 : 0-95—0-96) and
longer than penial sheath (ratio 1 : 1°55-1-73, average 1:62). Scape 45-53 (average 47) u long
and 46-49 (average 47) w wide, with 3-4 (average 3-4) hs. Pedicel with weak, polygonal,
dorsal reticulation ; 55-68 (average 63) w long and 32-42 (average 38) pw wide, with o-3
(average 1-2) f.s., 4-9 (average 6-8) h.s. and a sensillum placodeum. Segment III club-shaped,
2'7-3'7 (average 3-1) times longer than wide (68-86, average 78 w long and 23-29, average
26 uw wide) ; with 4-7 (average 6) f.s. of medium length, 1-1-1-3 (average 1-2) times as long as
width of segment ; with 1-3 usual sensilla basiconica. Segments [V-IX cylindrical ; lengths of
these segments (in uw) 106-133 (average 121), 76-133 (average 95), 72-91 (average 84), 70-95
(average 83), 61-76 (average 67) and 53-72 (average 66) respectively, widths varying from
21 to 27 uw, with distal segments wider than proximal ones ; with 18-27 (average 21), 13-24
(average 16), 13-23 (average 17), 12-18 (average 16), 12-19 (average 15) and 10-17 (average 15)
f.s. respectively, but no h.s. ; antennal bristles on segments VIII-IX not distinctly different
from f.s., sometimes slightly thicker. Segment X : terminal part not constricted ; 57-72
(average 65) u long and 21-27 (average 23) uw wide ; carrying 9-15 (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 not distinctly different from the f.s. ; with 2 sensilla basiconica
ventrally, one near apex and the other more proximal.

Thorax 513-597 (average 566) u long.

Prothorax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites medium-sized, without setae. Medial pronotal setae: usually 2 h.s. Post-

oo de, Hegel tee
i jane —400p

Fic. 23. Sphaerolecanium prunastri (Fonsc.), lateral view.

70 MORPHOLOGY AND TAXONOMY OF ADULT MALES

tergites very small and weakly sclerotized, without striations and without setae. Pleural
stvuctures typical of family. Steynwm with transverse ridge strong, median ridge absent (occa-
sionally a small basal stalk may be present) and triangular sclerite with lines arching latero-
anteriorly. Anteprosternal setae absent ; prosternal setae: 7-16 (average 11) f.s. and o-2
(average 0-7) h.s., scattered over the sternal area but not occurring anterior to the spiracles.

Mesothorvax. Mesoprephragma with shallow emargination. Pyrescutwm more than twice as
wide as long (average 181 and 82 uw respectively) ; anterior margin curved ; laterally bounded
by the prescutal ridges and posteriorly by the prescutal suture ; slightly more heavily sclerotized
medially but not reticulated. Scutwm. Median membranous avea subrectangular ; 72-91
(average 83) wu long and 1-92—2-48 (average 2-11) times as wide (width 152-220, average 175 yu);
with 2-4 (average 2°8) h.s. Scutellum 65-80 (average 74) up long and 141-182 (average 163) pv
wide, ratio being 1 : 2:0-2-3 (average 2-2) ; tubular, with a large ventral foramen ; occasion-
ally 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. Mesopostphragma with moderately deep emargination. Mesopleuron.
Mesopleural ridge strong, not 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 ; subepisternal ridge well developed, but below
membranous cleft indistinct and only marked by a band of dark sclerotization. Epimeron
small. Latevopleurite not bounded by an extension from marginal ridge. Basisternum large,
about 236 uw wide and 170 yu long, i.e. 1-8—2-2 (average 2-1) 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 ;
postmesospiraculay setae absent. Tegula small, membranous bulge with a small weak sclerite
posteriorly and 4-10 (average 7:3) h.s. Third avillavy wing sclerite with a pronounced ventral
projection at its base. Additional sclerite well defined. Antemetaspivacular setae absent.

Metathovax. Metanotum with anterior margin desclerotized medially ; suspensorial sclerites
absent. Postnotum consisting of a transverse sclerite on each side. Metatergal setae : usually
one h.s. on each side. Pleural ridge considerably reduced, extending only a short distance
above coxal articulation. Vestigial precoval ridge present. Episteynum reduced to a small
subtriangular plate ; epimevon produced posteriorly. Metathoracic spivacle similar to meso-
thoracic one. Dorsospivaculay setae absent. Postmetaspivacular setae : 0-15 (average 7:5) f.s.
and 1-3 (average 1:2) h.s. Metasternuwm represented by two small, irregular sclerites. Anterior
metasternal setae : 11-18 (average 14) f.s. and occasionally one h.s. ; posterior metasternal
setae : 0-3 (average 1-6) f.s.

Wings hyaline ; short (length 1200-1280, average 1231 w) and of medium width (500-530,
average 516 w), the ratio width to length being 1 : 2:34-2:42 (average 2-38) ; alar lobe and
alav setae absent. Halteves absent.

Legs short and moderately slender, with fore pair usually shortest and hind pair longest ;
ratio length of hind leg to body length is 1 : 2:02-2:34 (average 2-17). Length of segments

(in p) :

Leg Coxa Trochanter Femur Tibia Tarsus Claw Total

I 59-72 76-95 175-209 239-293 QI-110 26-28 667-803
(67) (87) (197) (280) (103) (27) (759)

Il 62-76 84-103 160-190 251-304 106-118 27-30 690-813
(70) (95) (179) (281) (113) (28) (766)

Ill 65-82 84-106 171-201 255-317 IOI-125 27-32 701-851
(76) (100) (189) (295) (116) (30) (805)

Cowae 3-6 (average 4-6) f.s. on the fore and 10-20 on the middle and hind coxa, and 10-20
h.s. on each ; fore coxa with 3-6 (average 4) pointed coxal bristles (about as long as segment) ;
apical seta about 3 as long as trochanter. Tvochanters 30-34 u wide ; with 6 oval sensilla,
6-10 f.s. and 5-9 h.s., the latter including 2 minute setae near basal ridge, one small seta on

OF THE FAMILY COCCIDAE 77

outer margin and a long apical seta which, on the fore trochanter, is 2-1-2-9 (average 2-4) times
as long as width of trochanter. Jemova very wide (46-57 uw), ratio width to length of hind
femur being 1 : 3-3-3°8 (average 3:5) ; each with 8-16 f.s. and 17-28 hs. Tibiae 23-30 u
wide, ratio width to length of hind tibia being 1 : 10-3—11-5 (average 10-9) ; each with 47-75
setae of which 31-49 are h.s. and 10-26 f.s., the latter about as long as width of tibia ; apical
spur about the same size on all tibiae. Tarsi 21-25 w wide, hind tarsus 4-8-5-5 (average 5-2)
times longer than wide ; each with 1-6 f.s. and 15-27 h.s. ; tarsal digitules subequal, longer
than claw. Claws of medium length, longer than width of tarsus ; slightly curved, with small
denticle near tip ; wngual digitules subequal, about as long as claw.

Abdomen 470-530 (average 498) uw long and 320-420 (average 375) u wide.

Segments I-VII : tergites represented by a small sclerite on each side on anterior margin of
segments II-III and by a weak plate on VI-VII ; sternites represented by a small sclerite on
each side on segments II-III and a transverse plate on VI. Caudal extension of segment VII
small, pointed, not sclerotized. Dorsal setae : h.s. only, usually one on each side on I and each
of segments V—VII. Pleural setae on segments III—VI consisting of o-1 (average 0-1), 0-1
(average 0-1), o—2 (average 0-5) and o-2 (average 0-9) f.s., and 0-2 (average 0-7), 0-3 (average
1-8), 1-3 (average 2-1) and 1~—4 (average 1-9) hair-like dorsopleural setae respectively ; ventro-
pleural setae absent or incorporated into the dorsopleural group. Segment VII with o-2
(average 1-1) f.s. and 3~—5 (average 3-4) h.s. Ventral setae : 1-5 (average 2-3) f.s. on segment II,
occasionally one on III, and usually one h.s. on each side on II-IV and 4 on each of segments
V-VII.

Segment VIII with transverse tergite and sternite, and with a large, cylindrical, caudal exten-
sion ; glandular pouch with 2 long, pointed setae, whose protruding part is 24-3 times 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 long, about ? total body length (ratio 1 : 3-42—3-85, average
3°68), 433-517 (average 473) u long and 46-51 (average 48) uw wide ; lateral sclerotizations
narrowly joined anterior to anus ; length of basal rod 1}-2 times that of aedeagus and extend-
ing 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 27-39
(average 32) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath.
Aedeagus of medium length (114-152, average 140) uw, penial sheath and basisternum longer, the
ratios being 1 : 3:1-3°8 (average 3-4) and I : 1-15-1-32 (average 1-23) respectively.

Material examined : 10 specimens, collected by B. Ogaza on Prunus spinosa L.
in Cracow, Poland during March, 1962 ; received from Z. Kawecki. A number of
specimens received from N. S. Borchsenius (collected by A. Kiritchenko in the
Odessa district, USSR in 1928) agreed well with the above description, but the
medial pronotal setae were not observed in any of the 10 specimens examined.
These specimens were generally in a poor condition (they were kept dry on cotton
wool), with many setae broken off.

THe ERIOPELTIS Group
ERIOPELTIS
Eriopeltis sp.

(Text-figs. 24 and 25)

Living specimens light reddish brown ; short, slender with comparatively long antennae and
short legs ; with many setae on the body and appendages. When mounted, total body length
1510-1590 (average 1550) uw ; width at mesothorax 260-290 (average 281) u. Wing expanse
2730-2930 (average 2839) u.

78 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Head subconical in dorsal view ; in lateral view flat, not obliquely elongated dorsoventrally,
anterodorsal bulge not pronounced ; length from apex to pronotal ridge 186-205 (average 199)
u, width across genae 182-198 (average 190) uw. Median crest sclerotized, distinctly polygonally
reticulated, with 7-11 (average 9) hair-like dorsal head setae. Midcvanial ridge dorsally absent ;
ventrally reduced to a shorter or longer median bar with lateral arms, surrounding area sclero-
tized and showing distinct polygonal reticulation. Genae large, weakly sclerotized, with distinct
polygonal reticulation ; without setae. Eyes : two pairs, subequal ; corneae of dorsal eyes
19-25 (average 22) uw in diameter and 2-:7-3-4 (average 3) times as much apart ; those of the
ventral eyes 19-25 (average 22) p in diameter and 1-2-1-8 (average 1-4) times as much apart.
Ocellus small, situated on a distinct protuberance which overhangs the postocular ridge. Ocular
sclerite well sclerotized, distinctly polygonally reticulated throughout. Pveocular ridge long, the
ridges of each side ventrally meeting or almost meeting each other medially. Postocular ridge
well developed throughout. Jnteroculay vidge broadly joining pre- and postocular ridges below
ocellus. Dorsal ocular setae absent. Ventral head setae : 9-16 (average 13) f.s. and 5-9 (average
6-9) h.s., arranged in a broad band on anterior part of ocular sclerite, occasionally with some
(o-3, average 0-7) f.s. and some (0-1, average 0-5) h.s. occurring in the area anterior to this
sclerite, but with none between the eyes. Pyveoval ridge absent. Tendon-like apodeme variable,
occasionally fully developed, but usually reduced or absent. Cvanial apophysis of medium
length ; apex trifurcate, extending to around the margin of the posterior level of the ventral
eyes. Mouth opening irregular. Anterior tentorial pits present, situated anterolateral to mouth
opening.

Antennae to-segmented, filiform ; 1030-1284 (average 1125) u long, i.e. longer than half
body length (ratio I : 1:23—1-51, average 1-38), longer than posterior leg (ratio 1 : I-42—1°52,

Fic. 24. Eviopeltis sp., dorsal and ventral view.

OF THE FAMILY COCCIDAE 79

average 1°46) and longer than penial sheath (ratio 1 : 3-04-3-86, average 3-39). Scape 44-53
(average 48) u long and 45-50 (average 46) wu wide, with 3 h.s. Pedicel with distinct, poly-
gonal, dorsal reticulation ; 49-61 (average 55) wu long and 34-42 (average 37) uw wide ; with
3-6 (average 4:2) f.s., 1-5 (average 2-9) h.s. anda sensillum placodeum. Segment III somewhat
club-shaped with a rugose surface ; 4:0-4-9 (average 4:6) times longer than wide (99-125,
average 114 uw long and 23-27, average 25 uw wide) ; with 5-11 (average 7:7) f.s. of medium
length, 1-3-1-7 (average 1-5) times longer than width of segment ; with one or two usual
sensilla basiconica. Segments IV-IX cylindrical, rugose ; lengths of these segments (in wp)
141-198 (average 162), 152-205 (average 171), 141-194 (average 159), 114-171 (average 135),
95-118 (average 106) and 84-95 (average 90) respectively, all of about the same width, varying
from 19 to 23 uw; with 14-23 (average 20), 20-30 (average 24), 19-28 (average 23), 14-23
(average 20), 14—20 (average 18) and 13~19 (average 16) f.s. respectively, but no h.s. ; antennal
bristles on segments VIII-IX not distinctly different from the f.s. Segment X : terminal part
not constricted ; 80-95 (average 86) p long and 19-21 (average 19) w wide ; carrying 10-14
(average 11°5) f.s., 3 capitate subapical setae and 5 antennal bristles of which the three long ones
are nearly half as long as the segment and the 2 short ones less than half as long as the f.s. ;
with 2 sensilla basiconica ventrally, one near apex and the other more proximal.

Thorax 448-486 (average 469) pv long.

Prothovax. Pyronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, with an occasional fleshy lateral pronotal seta occurring immediately
behind them. Medial pronotal setae and pores absent. Post-tergites very small, if present ; derm
irregularly striated, without setae. Pleural structures typical of the family. Sternum with
strong transverse ridge, usually narrowed medially ; median ridge only represented on anterior
part of well sclerotized triangular sclerite. Amnteprosternal setae absent ; prosternal setae :
1-7 (average 4:2) f.s., occurring medially around anterior part of the sternum.

Mesothorvax. Mesoprephragma with shallow emargination. Pyrescutuwm about twice as wide
as long (average 128 and 68 u respectively) ; anterior margin 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 weakly polygonally reticu-
lated ; subrectangular, 65-76 (average 73) u long and 1:47-1:79 (average 1-62) times as wide
(width 106-129, average 117 up) ; with 4-10 (average 7) h.s. Anterior arms of scutum irregularly
reticulated. Scutellum 38-46 (average 41) p long and 114-137 (average 127) uw wide, ratio
being 1 : 2:8—3-5 (average 3-1) ; tubular, with a small ventral foramen ; without setae. Post-
notum with anterior margin irregular, weakly sclerotized and not overlapped by metathoracic
fold ; postnotal apophysis and postalave well developed, the latter densely reticulated distally.
Mesopostphragma short, with shallow emargination. Mesopleuron. Mesopleural ridge strong,
but becoming weaker immediately above coxal articulation ; pleural apophysis and pleural
wing process well developed, the latter connected with episternum by a narrow basalare.
Subalare small. Episternum without polygonal reticulation ; subepisternal ridge becoming
broader ventrally, but below membranous cleft indistinct and only marked by a band of dark
sclerotization. Epimeron very small. Latevopleurite partly bounded anteriorly by an exten-

400p

Fic. 25. Eriopeltis sp., lateral view: also referable for lateral view of E. ?festucae (Fonsc.).

80 MORPHOLOGY AND TAXONOMY OF ADULT MALES

sion from marginal ridge. Basisteynum large, about 176 » wide and 130 yu long, i.e. 1:68-1:96
(average 1-79) times longer than membranous area of scutum ; median ridge reduced to a few
vestiges of which the combined length is less than half the length of the basisternum ; marginal
and precoxal ridges well developed ; without setae. Furca well developed. Mesothoracic
spivacle with well developed peritreme ; postmesospivaculay setae absent. Tegula small,
membranous bulge with a small weak sclerite posteriorly and with 1-5 (average 2-8) h.s. Third
axillary wing sclerite with small, rounded ventral projection at its base. Additional sclerite
well sclerotized. Antemetaspivacular setae absent.

Metathovax. Metanotum with posterior margin thickened throughout ; suspensorial sclerites
absent. Postnotum consisting of a large, subtriangular sclerite on each side. Metatergal setae :
usually one h.s. on each side. Pleural vidge considerably reduced, extending only a short
distance above coxal articulation. Episteynum reduced to a small subtriangular plate, ep7-
meyvon produced posteriorly. Metathoracic spiracle similar to mesothoracic one. Dorsospivacular
setae absent. Postmetaspivacular setae: 5-13 (average g-1) f.s. and o-2 (average o-4) hss.
Metasternal plate transverse, irregular ; weak irregular sclerotization present in the area
anterior to the sternum. Anterior metasternal setae : 14-23 (average 18) f.s. and o-3 (average
0:6) h.s. ; posterior metasternal setae : 8-15 (average 11) fs.

Wings hyaline ; long (1255-1340, average 1303 ») and narrow (width 395-430, average 413 wy),
ratio width to length being 1 : 3:02-3:29 (average 3:16) ; alar lobe and alar setae absent.
Halteres absent.

Legs short and moderately slender, with fore legs usually longest and middle pair shortest ;
ratio length of hind leg to body length is 1 : 1:96-2:09 (average 2:04). Length of segments

(in pw) :

Leg Coxa Trochanter Femur Tibia Tarsus Claw Total

I 59-68 76-84 186-198 272-319 110-122 30-34 736-806
(61) (79) (191) (292) (116) (33) (772)

II 57-07 72-80 175-198 270-315 103-114 28-31 711-789
(60) (75) (183) (291) (109) (29) (748)

Ill 61-68 72-84 177-217 277-323 99-114 30-34 722-809
(66) (76) (183) (296) (106) (32) (758)

Coxae with 6-9 (average 7:6), 10-13 (average 12) and 13-18 (average 16) f.s. on the fore,
middle and hind coxa respectively, and each with 5-12 h.s. ; fore coxa without coxal bristles ;
apical seta long, about as long as trochanter. Tvochanters 25-30 w wide ; with 6 oval sensilla,
5-10 f.s. and 7-10 h.s., the latter including 2 minute setae near basal ridge, one small seta on
outer margin and a very long apical seta which, on the fore trochanter, is 3-0—3-6 (average 3-3)
times as long as the width of the trochanter. Femora of medium width (34—42 yu), ratio width to
length of hind femur being I : 4-4—4:7 (average 4-6) ; with 14-22 (average 18) f.s. and 17-20
(average 19) h.s. on the fore, 10-18 (average 14) f.s. and 13-15 (average 14) h.s. on the middle,
and 11-15 (average 12) f.s. and 11-17 (average 15) h.s. on the hind femur. Tzibiae 19-23 wu
wide, ratio width to length of hind tibia being 1 : 12:2-14:2 (average 13:2) ; each with 42-58
setae, of which 22-34 are h.s. and 17-33 f.s., the latter about 14 times longer than width of tibia;
apical spur about the same size on all tibiae. Tarsus 19-23 u wide, hind tarsus 4-5—5-5 (average
4:9) times longer than wide ; each with 2-8 f.s. and 13-25 h.s. ; tarsal digitules subequal,
somewhat longer than claw. Claws long, 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 520-600 (average 572) w long and 240-300 (average 270) w wide.

Segments I-VIIL : a small tergite occasionally present on each side on anterior margin of
segment II ; sternites consisting of transverse plates on all segments, these usually less well
sclerotized medially. Caudal extension of segment VII small, rounded, weakly sclerotized
lateroventrally. Dorsal setae : o-2 (average 0-4), 0-3 (average 1-1), 3-10 (average 6-3), 3-10
(average 5-9), 2-5 (average 3:2) and o-4 (average 2-5) f.s. on segments II—-VII respectively, and
usually with one h.s. on each side on segments I and IV-VII. Pleural setae absent on segments

OF THE FAMILY COCCIDAE 81

I-III and on IV—-VI represented by h.s. only, which include the dorsopleural setae : usually
one on IV and 2 on segments V—VI, and ventropleural setae : occasionally one on V and usually
one on segment VI. Segment VII with o—6 (average 2-8) f.s. and 5—7 (average 5-8) h.s., some
of the posterior h.s. somewhat longer than rest. Ventral setae : segments II—-VII with 10-18
(average 15), 7-16 (average 12), 7-13 (average 10), 7-14 (average 11), 7-12 (average 9) and
8-14 (average 11) f.s. respectively ; h.s. : none on II, usually one on each side of III-IV, and
usually 4 (range 3—5) on each of segments V—VII.

Segment VIII with irregular tergite and transverse sternite, the latter carrying o—2 (average
1-3) f.s. ; caudal extension forming a small, simple lobe ; glandular pouch with 2 long, pointed
setae, whose protruding part is 4-6 times as long as section within pouch. Small I Xth tergite
present. Ante-anal setae: 4-9 (average 5°8) f.s. and an occasional h.s. Posterior margin
with 3-4 (average 3-1) h.s. on each side.

Genital segment. Penial sheath of medium length, about j—-} total body length (ratio
I : 4°5-4°9, average 4:7), 314-342 (average 332) u long and 29—36 (average 32) uw wide ; lateral
sclerotizations narrowly joined anterior to anus ; basal rod about 1} times as long as aedeagus,
extending anteriorly from base of aedeagus to between $ and ? of the length 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 27-37 (average 32) small setae ; a cluster of
small sensilla occurring ventrally near apex of sheath. Aedeagus short (68-80, average 75 wu
long), penial sheath and basisternum longer, the ratios being 1 : 4-1-4-9 (average 4-5) and
I : 1°57-1°94 (average 1-74) respectively.

Material examined : 10 specimens, bred in the laboratory from material collected
by myself on leaves of Agrostis tenuis Sibth. at the Imperial College Field Station,
Silwood Park, Sunninghill, Berks. ; males emerged during September, 1962 ; the
species was found to be bivoltine in the field.

Eriopeltis ?festucae (Fonscolombe)
(Text-figs. 26 and 25)

Living specimens light reddish brown ; short, slender, with comparatively long antennae and
short legs ; with many setae on body and appendages. When mounted, total body length
1360-1570 (average 1486) pu ; width at mesothorax 280-310 (average 293) uy. Wing expanse
2730-2920 (average 2820) u.

Head subconical in dorsal view ; in lateral view flat, not obliquely elongated dorsoventrally,
anterodorsal bulge not pronounced ; length from apex to pronotal ridge 175-201 (average 190)
u, width across genae 182-201 (average 195) u. Median crest sclerotized, distinctly polygonally
reticulated, with 6—9 (average 7-8) hair-like dorsal head setae. Midcranial ridge dorsally absent;
ventrally reduced to a shorter or longer median bar without lateral arms, surrounding area
sclerotized and showing distinct polygonal reticulation. Genae large, weakly sclerotized, with
distinct polygonal reticulation ; without setae. Eyes: two pairs, subequal ; corneae of
dorsal eyes 17-23 (average 20) u in diameter and 3-3—4:2 (average 3-7) times as much apart ;
those of the ventral eyes 19-23 (average 21) uw in diameter and 1-5-1-9 (average 1-7) times as
much apart. Ocellus small, situated on a distinct protuberance which overhangs the postocular
ridge. Ocular sclerite well sclerotized, distinctly polygonally reticulated throughout. Pyre-
ocular ridge long, the ridges of each side ventrally meeting or almost meeting each other medially.
Postocular ridge well developed and distinct throughout. Intevocular ridge broadly joining pre-
and postocular ridges below ocellus. Dorsal ocular setae absent. Ventral head setae: 12-25
(average 18) f.s. and 7-17 (average 12) h.s., arranged in a broad band on anterior part of the
ocular sclerite, usually also with some (o—2, average 1) f.s. and some (0-3, average 1) h.s. occurring
in the area anterior to this sclerite, but with none between the eyes. Preoval ridge absent.

82 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Tendon-like apodeme absent or very short. Cvanial apophysis of medium length ; apex trun-
cate or rounded, not reaching level of anterior margin of ventraleyes. Mouth opening irregular.
Anterior tentorial pits present, situated anterolateral to mouth opening.

Antennae 1o-segmented, filiform ; 996-1060 (average 1018) yu long, i.e. longer than half body
length (ratio I : 1-37-1-52, average 1-45), longer than posterior leg (ratio I : 1:38-1:45, average
1-41) and longer than penial sheath (ratio I : 3:31-3:49, average 3-41). Scape 42-49 (average 47)
uw long and 44-48 (average 46) pp wide, with 3 h.s. Pedicel with distinct, polygonal, dorsal
reticulation ; 49-69 (average 57) u long and 38—42 (average 39) uw wide ; with 2-5 (average 3-4) f.s.,
3-7 (average 4-9) h.s. and a sensillum placodeum. Segment III somewhat club-shaped, witha
rugose surface ; 4:3—6-0 (average 5) times longer than wide (99-118, average 113 p long and
19-25, average 22 uw wide) ; with 3-9 (average 5-4) f.s. of medium length, 1-2-1-7 (average
1-4) times longer than width of segment ; with 1-3 usual sensilla basiconica. Segments 1V-IX
cylindrical, rugose ; lengths of these segments (in uw) 137-156 (average 148), 133-156 (average
145), 133-163 (average 146), 103-118 (average 111), 84-99 (average 90) and 72-87 (average 82)
respectively, all of about the same width, varying from 19 to 23 up and with 13-21 (average 18),
19-23 (average 21), 20-32 (average 26), 18-27 (average 22), 13-25 (average 18) and 11-17
(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 ; 72-89 (average 80) yu long and
19-22 (average 20) uw wide ; carrying 6-12 (average 9-1) 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 about half as long as the f.s. ; with 2 sensilla basiconica ventrally, one near apex and the
other more proximal.

Thorax 448-502 (average 473) pu long.

Fic. 26: Eviopeltis ?festucae (Fonsc.), dorsal and ventral view.

OF THE FAMILY COCCIDAE 83

Prothorax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, with an occasional f.s. occurring on or immediately behind them.
Medial pronotal setae rarely present, pores absent. Posttergites small, without wavy striations
and without setae. Pleural structures typical of the family. Steynuwm with strong transverse
ridge, usually somewhat narrowed medially ; median ridge sometimes complete but usually
interrupted ; triangular sclerite well sclerotized. Anteprosternal setae absent ; prosternal setae
consisting of o—2 (average o-7) f.s. occurring medially around anterior part of sternum.

Mesothovax. Mesoprephragma without emargination. Prescutum about 14 times as wide as
long (average 125 and 81 yp respectively) ; anterior margin curved ; laterally bounded by the
prescutal ridges and posteriorly by the prescutal suture ; slightly more heavily sclerotized
medially but not reticulated. Scutwm. Median membranous area without reticulation ;
subrectangular, 61-72 (average 64) uw long and 1-47-2-00 (average 1-71) times as wide (width
99-122, average 109 uw) ; with 2-10 (average 6) h.s. Anterior arms of scutum not reticulated.
Scutellum 42-49 (average 45) uw long and 114-129 (average 122) uw wide, the ratio being
I : 2-5-3°0 (average 2-7) ; tubular, with ventral foramen of medium size ; setae absent.
Postnotum with the anterior margin irregular, weakly sclerotized and not overlapped by meta-
thoracic fold ; postnotal apophysis and postalare well developed, the latter densely reticulated
distally. Mesopostphragma with shallow emargination. Mesopleuron. Mesopleural ridge
strong, but becoming weaker immediately above coxal articulation ; pleuval apophysis and
pleural wing process well developed, the latter connected with episternum by a narrow basalare.
Subalare small. Episternum without polygonal reticulation ; subepisternal ridge becoming
broader ventrally, but below the membranous cleft indistinct and only marked by a band of
dark sclerotization. Epimeron very small. Lateropleurite partly bounded anteriorly by an
extension from marginal ridge. Basisternum large, about 171 w wide and 128 yw long, i.e.
1°74-2°19 (average 2-01) times longer than membranous area of scutum ; median ridge some-
times complete but usually interrupted, the combined length of the vestiges more than half
the length of the basisternum ; marginal and precoxal ridges well developed ; without setae.
Furca well developed. Mesothoracic spiracle with well developed peritreme ; postmesospivacular
setae absent. Tegula small, its membranous bulge with a small weak sclerite posteriorly and
with 2-6 (average 3:6) h.s. Third axillary wing sclerite with small, rounded ventral projection
at its base. Additional sclerite well sclerotized. Antemetaspivaculay setae absent.

Metathorax. Metanotum with posterior margin thick and well developed throughout ;
suspensorial sclerites absent ; a small sclerite sometimes present on each side anterior to post-
notum. Postnotum consisting of a large, subtriangular sclerite on each side. Metatergal setae :
one h.s. oneach side. Plewral ridge considerably reduced, extending only a short distance above
coxal articulation. Episternum reduced to a small subtriangular plate ; epimeron produced
posteriorly. Metathoracic spiracle similar to mesothoracic one. Dorsospivaculay setae absent.
Postmetaspivacular setae : 8-11 (average 9:2) f.s. and o-1 (average 0-3) h.s. Metasternal plate
transverse, irregular ; no sclerotization present in the area anterior to sternum. Anterior and
posterior metasternal setae : 8-17 (average 13) and 7—15 (average 10) f.s. respectively.

Wings hyaline; long (1240-1320, average 1280 uw) and narrow (width 400-460, average 422 uy),
the ratio being 1 : 2:87-3:15 (average 3-04) ; alay lobe and alar setae absent. Halteres absent.

Legs short and moderately slender, with fore legs the longest and either middle or hind legs
shortest ; ratio length of hind leg to body length is 1 : 1-98—2-12 (average 2-05). Length of
segments (in y) :

Leg Coxa ‘Trochanter Femur Tibia Tarsus Claw Total

I 53-68 76-84 186-194 258-293 I10-125 29-34 726-775
(62) (80) (190) (277) (114) (32) (754)

Il 61-68 72-80 171-190 251-274 106-122 27-32 695-751
(62) (76) (180) (262) (111) (29) (720)

III 61-68 72-78 167-186 251-274 99-108 30-32 686-731

(64) (75) (177) (265) (104) (31) (714)

84 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Coxae each with 7-15 f.s. and 9-15 h.s. ; fore coxa without coxal bristles ; apical seta long,
about as long as trochanter. Tvochanters 26-30 u wide ; with 6 oval sensilla, 5-12 f.s. and
7-11 h.s., the latter including 2 minute setae near basal ridge, one small seta on the outer margin
and one long apical seta which, on the fore trochanter, is 3-I-5-o0 (average 4-1) times as long as
width of trochanter. Femova of medium width (34-42 yu), ratio width to length of hind femur
being 1 : 4-0-4-9 (average 4-6), each with 6-14 f.s. and 14-32 h.s. Tuibiae 21-25 w wide, ratio
width to length of hind tibia being 1 : 10-3-13-1 (average 11-6) ; each with 48-62 setae, of
which 30-52 are h.s. and 9-18 f.s., the latter about 14 times as long as width of tibia ; apical
spur about the same size on all tibiae. Tarvsi 19-23 wu wide, hind tarsus 4-3—4:8 (average 4-6)
times longer than wide ; each with o-3 f.s. and 21-30 h.s. ; tarsal digitules subequal, slightly
longer than claw. Claws long, about 1} times as long as width of tarsus ; slightly curved, with
small denticle near tip ; ungual digitules subequal, about as long as claw.

Abdomen 470-600 (average 563) u. long and 250-300 (average 272) uw wide.

Segments I-VI1 : tergites absent ; steynites consisting of transverse plates on all segments,
these usually less well sclerotized medially. Caudal extension of segment VII small, rounded
and weakly sclerotized lateroventrally. Dorsal setae : o-3 (average 0-8), 0-6 (average 2-5),
7-16 (average 9-7), 3-8 (average 4:9), 2-6 (average 3-4) and 2-6 (average 3-6) f.s. on segments
II-VII respectively, and usually with one h.s. on each side on segments I and IV—-VII. Pleural
setae absent on segments I-III, and on IV—VI represented by h.s. only, which include dorso-
pleural setae : sometimes one on IV and usually 2 on segments V—VI, and ventropleural setae :
usually one on each of segments V and VI. Segment VII with 1-8 (average 3-6) f.s. and 3-6
(average 4-8) h.s., some of the posterior h.s. somewhat longer than rest. Ventral setae :
segments II to VII with 8—22 (average 14), 6-21 (average 13), 8-18 (average 11), 6-15 (average
12), 9-14 (average 12) and 8-15 (average 13) f.s. repectively ; h.s. ; none on II, sometimes one
on each side on III and IV, and usually 4 (4-5) on each of segments V—VII.

Segment VIII with irregular tergite and transverse sternite, the latter carrying 0-3 (average
1-4) f.s. ; caudal extension forming a small, simple lobe ; glandular pouch with 2 long, pointed
setae, whose protruding part is 4-6 times as long as section within pouch. Small IXth tergite
present. Ante-anal setae: 1-4 (average 2-7) f.s. and occasionally one short or long h:s.
Posterior margin with 3 h.s. on each side.

Gemital segment. Penial sheath of medium length, about + total body length (ratio 1 : 4-53—
5:16 (average 4-95), 289-308 (average 300) u long and 30-36 (average 34) uw wide, lateral sclero-
tizations narrowly joined anterior to anus ; basal vod 14-2 times as long as aedeagus, extending
anteriorly from the base of the latter for $—? of the distance to the apex of the basal membranous
area ; apex of sheath without membranous extension. The area from base of the sheath to
tip of aedeagus with 25-36 (average 30) small setae ; a cluster of small sensilla occurring
ventrally near apex of sheath. Aedeagus short (61-69, average 65u long), penial sheath and
basisternum longer, ratios being I : 4:3-5-1 (average 4-7) and I : 1-78—2-19 (average 1-99)
respectively.

Material examined : 10 specimens, bred in the laboratory from material collected
by myself on leaves of Festuca rubra L. at the Imperial College Field Station, Silwood
Park, Sunninghill, Berks. ; males emerged during September, 1963 ; the species
was found to be bivoltine in the field.

This species can readily be separated from Eviopeltis sp. by the truncate or rounded
apex of the cranial apophysis, the absence of the lateral arms of the midcranial ridge
and the better developed median ridge of the mesosternum. In addition the post-
tergital region, and the anterior arms and membranous area of the mesoscutum are
not reticulated.

OF THE FAMILY COCCIDAE 85

LUZULASPIS
Luzulaspis luzulae (Dufour)
(Text-figs. 27 and 28)

Living specimens light reddish brown ; short, slender, with comparatively long antennae and
short legs ; with many setae on the body and appendages. When mounted, total body length
1020-1290 (average 1141) u ; width at mesothorax 240-270 (average 255) u. Wing expanse
2090-2350 (average 2213) wu.

Head subconical in dorsal view ; in lateral view flat, not obliquely elongated dorsoventrally,
anterodorsal bulge not pronounced ; length from apex to pronotal ridge 148-179 (average 164)
u, width across genae 167-182 (average 175) u. Median crest sclerotized, distinctly polygonally
reticulated, with 7-10 (average 8-7) hair-like dorsal head setae. Midcranial ridge dorsally
absent ; ventrally reduced to a shorter or longer median bar with lateral arms, surrounding
area not sclerotized, but showing distinct polygonal reticulation. Genae large, weakly sclero-
tized, with distinct polygonal reticulation ; without setae. Eyes : two pairs, small, subequal;
corneae of dorsal eyes 14-17 (average 15) in diameter and 4-0-4-5 (average 4-2) times as much
apart ; those of the ventral eyes 15-20 (average 16) uw in diameter and 2-2—2-5 (average 2-3)
times as much apart. Ocellus small. Ocular sclerite well sclerotized, distinctly polygonally
reticulated throughout. Preoculay ridge long, the ridges of each side ventrally meeting or
almost meeting each other medially. Postocular ridge well developed, but tapering dorsally.
Interocular ridge broadly joining pre- and postocular ridges below ocellus. Dorsal ocular setae
absent. Ventral head setae : 11-30 (average 25) f.s. and 6-16 (average 10) h.s., arranged in a
broad band on anterior part of ocular sclerite, frequently with some (o—6, average 2-6) f.s.
occurring in the area anterior to the sclerite and occasionally with a seta just behind anterior
level of eyes ; a pair of median h.s. in the middle of the band distinctly longer than the rest.

Fic. 27. Luzulaspis luzulae (Dufour), dorsal and ventral view.

86 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Preoval vidge absent. Tendon-like apodeme short. Cvanial apophysis long ; apex truncate,
occasionally bifurcate, extending to around the level of the anterior margin of the ventral eyes.
Mouth opening irregular. Anterior tentorial pits present, situated anterolateral to mouth
opening.

Antennae 10-segmented, filiform ; 729-901 (average 822) u long, i.e. longer than half body
length (ratio I : 1:28-1-56, average 1-39), longer than posterior leg (ratio I : I1-36—-1-40, average
1-38) and longer than penial sheath (ratio I : 4:36-6:02, average 4:87). Scape 36-46 (average
41) p long and 36-40 (average 38) uw wide, with 3 h.s. and occasionally one f.s. Pedicel with
weak, polygonal, dorsal reticulation ; 42-49 (average 45) u long and 31-34 (average 33) uw wide;
with 5-10 f.s., 1-5 (average 3-6) h.s., and a sensillum placodeum. Segment III somewhat club-
shaped, 3:0-3-7 (average 3-3) times longer than wide (68-84, average 75 uv long and 23 yp wide);
with o—2 (average 0-7) h.s. and 1-6 (average 3-3) f.s., the latter of medium length, 1-2-1-6
(average 1-4) times longer than width of segment ; with 1 or 2 usual sensilla basiconica.
Segments IV-IX cylindrical ; lengths of these segments (in uw) 114-158 (average 131), 105-137
(average 122), 95-137 (average 118), 82-106 (average 97), 67-82 (average 74) and 57-72
(average 63) respectively, widths varying from 15 to 23 uw, with segment IX somewhat wider than
proximal ones ; with 11-24 (average 18), 13-25 (average 20), 20-30 (average 23), 18-28
(average 22), 13-19 (average 16) and 12-20 (average 16) f.s. respectively, but no h.s. ; antennal
bristles on segments VIII and IX not always distinctly different from f.s. Segment X : terminal
part not constricted ; 46-61 (average 54) uw long and 19-23 (average 20) uw wide ; carrying
4-14 (average 8) 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 shorter than the f.s. ; with 2 sensilla
basiconica ventrally, one near apex and the other more proximal.

Thorax 357-424 (average 387) wu long.

Prothovax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, without setae. Medial pronotal setae and pores absent. Post-tergites
small, weakly sclerotized, with irregular wavy strictions, without setae. Pleural structures
typical of family. Steynuwm with transverse ridge narrowed medially, median ridge represented
by a short basal stalk, and triangular sclerite weakly sclerotized, with lines arching latero-
anteriorly from basal stalk ; a small apophysis sometimes present anterior to this sclerite.
Anteprosternal setae frequently absent, but sometimes up to 2 present on each side ; prosternal
setae : 0-25 (average 14), but usually 12-21 f.s. and o-1 (average 0-5) h.s., none of these occurring
anterior to spiracles.

Mesothovax. Mesoprephvagma short, with shallow emargination. Pvrescutum about twice as
wide as long (average 116 and 60 u respectively) ; anterior margin curved ; laterally bounded
by strong prescutal vidges and posteriorly by the prescutal suture ; slightly more heavily sclero-
tized medially ; polygonally reticulated. Scutum. Median membranous area subrectangular ;
49-72 (average 60) uw long and 1-42~1-92 (average 1-70) times as wide (width 97-114, average
102 uw); with 3-8 (average 4:7) h.s. Anterior arms of scutum irregularly reticulated.
Scutellum 29-34 (average 32) u long and 95-114 (average 102) uw wide, ratio being I : 3:0-3°8
(average 3-4) ; tubular, with small ventral foramen ; usually without setae. Postnotum with

Fic. 28. Luzulaspis luzulae (Dufour), lateral view.

OF THE FAMILY COCCIDAE 87

anterior margin irregular, weakly sclerotized, and partly overlapped by metathoracic fold ;
postnotal apophysis and postalare well developed, the latter densely reticulated distally. Meso-
postphragma short, with shallow emargination. Mesopleuron. Mesopleural ridge strong, not
interrupted above coxal articulation ; pleural apophysis and pleural wing process well developed,
the latter connected with episternum by a narrow basalare. Subalave small. Episternum
without polygonal reticulation ; subepisternal ridge broadening ventrally, but below mem-
branous cleft indistinct and only marked by a band of dark sclerotization. Epimeron very
small. Lateropleurite partly bounded anteriorly by an extension from marginal ridge. Basit-
sternum large, about 163 uw wide and gt yp long, i.e. 1:26—1-72 (average 1-52) 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 ; postmesospiracular setae absent. Tegula small, membranous bulge with
a small weak sclerite posteriorly and with 1-5 (average 2-8) h.s. Third axillary wing sclerite
with a small, rounded ventral projection at its base. Additional sclerite weak, but distinct.
Antemetaspivacular setae absent.

Metathovax. Metanotum with posterior margin thickened throughout ; suspensorial sclerite
absent. Postnotum large, transverse, weak or interrupted medially. Metatergal setae : one
h.s. on each side. Pleural ridge considerably reduced, extending only a short distance above
coxal articulation. Episternum reduced to a small subtriangular plate ; epimeron produced
posteriorly. Metathoracic spiracle similar to mesothoracic one. Dorsospivaculay setae absent.
Postmetaspivaculay setae 6-14 (average 10) f.s. and o~—2 (average 0-7) h.s. Metasternal plate
weak, but more heavily sclerotized around anterior margin. Anterior metasternal setae :
16-23 (average 19) f.s. and no his. ; posterior metasternal setae : 7-19 (average 13) f.s.

Wings hyaline ; long (g10-1050, average 969 w) and narrow (295-360, average 336 u wide),
ratio width to length being 1 : 2:80-3:01 (average 2-90) ; alay lobe and alay setae absent.
Halteres absent.

Legs short and moderately slender, all 3 pairs subequal in size, fore legs being only slightly
shorter ; ratio length of hind leg to body length is 1 : 1-89-2-o1 (average 1-98). Length of
segments (in y) :

Leg Coxa ‘Trochanter Femur Tibia Tarsus Claw Total

I 43-57 61-70 125-152 186-217 76-91 26-28 518-599
(49) (65) (140) (203) (82) (27) (566)

Il 49-60 62-72 129-148 186-219 84-89 26-28 540-616
(54) (69) (139) (206) (86) (27) (578)

III 40-57 61-68 125-152 190-224 80-87 25-27 536-613
(52) (65) (140) (209) (84) (26) (577)

Coxae each with 10-17 f.s. and 10-18 h.s. ; fore coxa without coxal bristles ; apical seta long,
about as long as trochanter. Tyvochanters 21-24 » wide ; with 6 oval sensilla, 7-12 (average 9)
f.s. and 7-9 (average 7:6) 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 1-7—2:2 (average
1-9) times as long as width of trochanter. Femora of medium width (30~—32) yu, ratio width to
length of hind femur being 1 : 4-1—-5:0 (average 4-6) ; each with 8-16 f.s. and 9-17h.s. Tuibiae
17-23 w wide, ratio width to length of hind tibia being 1 : 9:1-11-6 (average 10-5) ; each with
20-39 h.s. and 8-22 f.s., the latter about 14 times longer than width of tibia ; apical spur about
the same size on all tibiae. Tavs: 19-21 u wide, hind tarsus 4-0—4-3 (average 4-2) times longer
than wide ; each with 2-6 f.s. and 17-23 hs. ; tarsal digitules subequal, slightly longer than
claw. Claws distinctly longer than width of tarsus, slightly curved, with small denticle near
tip ; ungual digitules subequal, about as long as claw.

Abdomen 350-505 (average 430) u long and 200—260 (average 227) uw wide.

Segments I-VII : tergites represented by a weak sclerite on or near anterior margin of each
segment ; steynites consisting of a weak transverse sclerite on segments II-III, a small sclerite
on each side of V and VI, and a transverse plate on segment VII. Caudal extension of segment

88 MORPHOLOGY AND TAXONOMY OF ADULT MALES

VII small, rounded and weakly sclerotized latero-ventrally. Dorsal setae : f.s. occasionally
present on segments I (0-2, average 0-6) and V—VII (up to 3) ; usually one h.s. on I and each of
segments IV—VII. Pleural setae : absent onsegments I, II and usually III, on segments IV—VI
consisting of dorsopleural setae : usually 2 h.s. on each of segments IV—VI, and ventvopleural setae :
usually one h.s. and occasionally one f.s. on segments V-VI. Segment VII with 4-11 (average
8) f.s. and 3-5 (average 4) h.s., some of the posterior h.s. somewhat longer than rest. Ventral
setae : 3-14 (average 9), 3-14 (average 9), 0-9 (average 3-7), 0-7 (average 2-6), o—-6 (average 3-3)
and 1-9 (average 4-9) f.s. respectively ; h.s. : none on II, usually one on each side on III and
IV, and usually 4 on each of segments V—VII.

Segment VIII with weak tergite and transverse sternite ; caudal extension forming a small,
simple lobe ; glandulay pouch absent, replaced by a shallow depression and one long pointed
seta. No IXth tergite observed. Ante-anal setae absent, but 2-6 (average 3-9) small, circular
pores present. Posterior margin with 2-4 (average 3) h.s. on each side, one of these situated
more laterally on caudal extension.

Genital segment. Penial sheath short, about 4 to 4 total body length (ratio 1 : 6-1-7:3,
average 6-7), 152-186 (average 173) w long and 30-38 (average 34) uw wide ; lateral sclerotiza-
tions narrowly joined anterior to anus ; basal vod about as long as aedeagus, extending anter-
iorly from base of aedeagus to between 4 and # of the distance to apex of the basal membranous
area ; apex of sheath without membranous extension. The area from base of sheath to tip of
aedeagus with 18-26 (average 22) small setae ; a cluster of small sensilla occurring ventrally
near apex of sheath. Aedeagus short (40-53, average 46 uw long), penial sheath and basisternum
longer, the ratios being I : 3:2-4-3 (average 3:8) and I : 1-57—2-29 (average 1-97) respectively.

Material examined : 10 specimens, bred in the laboratory from material collected
by myself on Luzula campestris (L.) at the Imperial College Field Station, Silwood
Park, Sunninghill, Berks. ; males emerged during May, 1962 ; the species was
found to be bivoltine in the field.

THE ITNGLISIA Group

INGLISIA
Inglisia theobromae Newstead
(Text-figs. 29 and 30)

A medium-sized and moderately robust species, with comparatively long antennae and
moderately long legs ; with numerous setae on the body and appendages. When mounted,
total body length 1610-1790 (average 1710) uw ; width at mesothorax 370-420 (average 399) wu.
Wing expanse 2870-3270 (average 3032) uw.

Head subconical in dorsal view ; in lateral view obliquely elongated, with the anterodorsal
bulge not pronounced ; length from apex to pronotal ridge 209-262 (average 245) yw, width
across genae 251-277 (average 266) u. Median crest well sclerotized, especially around the
posterior margin, which is obtuse ; weakly polygonally reticulated ; with 4—11 (average 7-2)
fleshy and 10-14 (average 13) hair-like dorsal head setae. Midcranial ridge dorsally absent ;
ventral part with median bar reduced posteriorly, the lateral arms widely diverging, and the
surrounding area weakly sclerotized and reticulated. Genae large, sclerotized, not reticulated ;
each with 5-11 (average 7) fleshy and o-4 (average 1-2) hair-like genal setae. Eyes: four
pairs ; dorsal and ventral pairs large, subequal ; lateral pairs large, only slightly smaller than
dorsal and ventral eyes, subequal ; corneae of dorsal eyes 32-37 (average 35) » in diameter
and 1-6—-2:1 (average 1-8) times as much apart ; those of the ventral eyes 30-36 (average 34) u
in diameter and 0-6-0-8 (average 0-7) times as much apart. Ocellus small. Ocular sclerite

OF THE FAMILY COCCIDAE 89

weakly sclerotized ; reticulated only dorsally and between and anterior to the ventral eyes.
Preoculay ridge with ventral part not extending far below articular process. Postocular ridge
well developed throughout ; below ocellus the ridge splits up, completely enclosing the ocellus.
Interoculay ridge absent, apparently represented by a small posteriorly directed process below
the articular process of the preocular ridge. Dorsal ocular setae : on each side 0-3 (average
1-7) f.s. and occasionally one h.s. ; ventral head setae : 18-27 (average 23) f.s. and 7-11 (average
9) h.s., of which 1-4 (average 2) f.s. and o—3 (average 1-1) h.s. occur anteriorly, around the
midcranial ridge, and 1-4 (average 3) f.s. and o—2 (average 1) h.s. between and behind the
ventral eyes. Pyreoval ridge present. Tendon-like apodeme long. Cranial apophysis of medium
length ; apex bifurcate, not reaching level of anterior margin of ventral eyes. Mouth opening
irregular. Anterior tentorial pits absent.

Antennae 1o-segmented, filiform ; 903-1047 (average 986) u long, i.e. longer than half body
length (ratio 1 : 1-67—1-82, average 1-73), longer than posterior leg (ratio I : 1-07—-1-13, average
1-10) and longer than penial sheath (ratio 1 : 2:53-2:78, average 2-66). Scape 46-53 (average
47) uw long and 49-53 (average 51) w wide, with 3 h.s. Pedicel not reticulated ; 38-48 (average
43) uw long and 34-38 u wide ; with 1-4 (average 2) f.s., 4-9 (average 6-3) h.s. and a sensillum
placodeum. Segment III1 somewhat club-shaped, 3-6—4°5 (average 4) times longer than wide
(103-110, average 107 uw long and 25~—29, average 27 uw wide) ; with 1-4 (average 2-2) h.s. and
6-12 (average Io) f.s., the latter of medium width, o-8-1-1 (average 0-9) times as long as width
of segment ; with 1-3 usual sensilla basiconica. Segments [V-IX cylindrical ; lengths of these
segments (in uw) 114-141 (average 127), 114-137 (average 127), 125-152 (average 138), 116-129
(average 120), 89-122 (average 105) and 76-95 (average 82) respectively, all of about the same

Nays }

\
Pas

Wy

Fic. 29. Inglisia theobromae (Newst.), dorsal and ventral view.

go MORPHOLOGY AND TAXONOMY OF ADULT MALES

width, varying from 17 to 25 uw; with 22-25 (average 24), 22-30 (average 25), 22-34 (average
28), 20-28 (average 24), 17-27 (average 21) and 15-21 (average 17) f.s., respectively, and
occasionally each with one or two h.s. ; antennal bristles on segments VIII-IX distinctly
thicker than f.s. Segment X : terminal part not constricted ; 76-97 (average go) uw long and
42-46 (average 43) uw wide ; carrying 9-16 (average 13) f.s., 2 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
shorter ones not distinctly different from the f.s. ; with 2 sensilla basiconica ventrally, one near
apex and the other more proximal.

Thorax 562-635 (average 600) u long.

Pyothovax. Pyronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, without setae. Medial pronotal setae : occasionally one or two h.s.
Post-tergites very small, without striations and without setae. Pleural structures typical of the
family. Steynum with transverse ridge strong, median ridge reduced to a short basal stalk and
triangular sclerite bearing numerous small projections. Anteprosternal setae absent ; prosternal
setae : 5-11 (average 7-8) f.s. and o-2 (average 0-9) h.s., the setae not occurring in front of
spiracles.

Mesothovax. Mesoprephragma with shallow emargination. Prescutuwm about twice as wide
as long (average 171 and 90 uw respectively) ; anterior margin almost straight ; 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 ; 61-80
(average 70) u long and 2-10—2-88 (average 2-51) times as wide (width 156-179, average I7I yu);
with 3-15 (average 6-6) f.s. and 7-15 (average 12) h.s. Scutellum 53-80 (average 70) uw long
and 144-175 (average 163) uw wide, the ratio being I : 2-1-2-9 (average 2-4) ; not tubular.
Postnotum with anterior margin irregular and usually partly overlapped by metathoracic fold ;
postnotal apophysis and postalave well developed, the latter densely reticulated distally. Meso-
postphragma with deep emargination. _Mesopleuron. Mesopleural ridge strong, not interrupted
above coxal articulation ; pleural apophysis and pleural wing process well developed, the latter
connected with episternum by a narrow basalave. Subalavesmall. Episteynum not reticulated;
subepisternal ridge well developed, but below membranous cleft indistinct and only marked by
a band of dark sclerotization. Epimeyvon small. Lateropleurite partly bound anteriorly by
an extension from marginal ridge. Basisternum large, about 253 w wide and 162 uy long, i.e.
2:00-2:63 (average 2-33) times longer than membranous area of scutum ; median ridge almost
completely absent, represented by somewhat darker sclerotization medially which sometimes
appears ridge-like over short distances ; bounded by strong marginal and precoxal ridges ;
without setae. Furca well developed. Mesothoracic spivacle with well developed peritreme ;
postmesospivacular setae : 20-28 (average 24) f.s. and o-3 (average 1) h.s., arranged in a band
behind the spiracles and prosternum. Tegula small, membranous bulge with a weak sclerite
posteriorly, with 2-6 (average 4-2) h.s. and occasionally one f.s. Third axillary wing sclerite
with a small ventral projection at its base. Additional sclerite welldefined. Antemetaspivaculay
setae : 1-6 (average 3:8) f.s.

Fic. 30. Inglisia theobromae (Newst.), lateral view.

OF THE FAMILY COCCIDAE gl

Metathorax. Metanotum with anterior margin strong medially ; suspensorial sclerites
absent ; a small, additional sclerite occasionally present anterior to postnotum. Postnotum
consisting of two narrow, transverse sclerites in a tandem alignment on each side. Metatergal
setae : 0-2 (average 0-7) f.s. and o—2 (average 0-8) h.s. on each side. Pleural ridge considerably
reduced, extending only a short distance above coxal articulation. Vestigial precoxal ridge
present. Episternum reduced to a subtriangular plate ; epimeron produced posteriorly.
Metathoracic spivacle similar to mesothoracic one. Dorsospivacular setae : 1-8 (average 3-6)
{.s. and o-2 (average 0-3) h.s. Postmetaspivacular setae: 6-11 (average 8-2) f.s. and 0-5
(average 1-7) h.s. Metasternum represented by a transverse plate. Anterior metasternal setae :
13-24 (average 19) f.s. and 0-3 h.s. Posterior metasternal setae : 5-16 (average 8-4) f.s.

Wings hyaline; of medium length (1270-1370, average 1333 uw) and width (540-580, average
561 uw), ratio width to length being 1 : 2-33—2-45 (average 2-38) ; alay lobe and alar setae absent.
Halteres absent.

Legs moderately long and slender, with middle pair shortest and fore or hind pair longest ;
ratio length of hind leg to body length is 1 : 1-87-1-98 (average 1-91). Length of segments
(in p) :

Leg Coxa _Trochanter Femur Tibia Tarsus Claw Total

I 57-61 68-84 224-255 308-369 133-164 20-25 821-952
(60) (80) (239) (339) (148) (22) (888)

Il 57-97 65-76 201-232 285-338 150-175 23-27 787-901
(64) (71) (217) (315) (169) (24) (860)

TEE 61-72 76-84 201-232 304-357 160-179 25-29 827-937
(67) (79) (219) (333) (173) (26) (896)

Coxae : inner surface of front coxa with small projections ; with 3-7 (average 5) f.s. on the
fore and 14-21 on the middle and hind coxa, and each with 6-12 h.s. ; fore coxa without
coxal bristles ; apical seta about } as long as trochanter. Tyrochanters 23-29 u wide ; with 6
oval sensilla, 7-15 f.s. and 6-9 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 2-0—2-4 (average
2-3) times as long as width of trochanter. Femora of medium width (36—44 yu), the ratio width
to length of hind femur being 1 : 4-8-5-4 (average 5-2) ; each with 18-33 f.s. and 13-25 hss.
Tibiae 22-26 u wide, ratio width to length of hind tibia being 1 : 12-9-15-7 (average 14) ; each
with 64-99 setae of which 20-35 are h.s. and 39-68 f.s., the latter about as long as width of
tibia ; apical spur on fore tibia short, about half as long as on the other tibiae. Tarsi 19-27 u
wide, hind tarsus 6-4—7°8 (average 7-2) times longer than wide ; each with 12—28 f.s. and as many
h.s. ; tarsal digitules subequal, about as long as 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 490-610 (average 558) uw long and 320~420 (average 380) p wide.

Segments I-VII : tergites on segments II and III represented by 3 small sclerites on anterior
margin—one medially and one on each side, and on segment IV by a small median sclerite.
Pleurally a continuous band of sclerotization extends from segments IV—VII, with a separate
small sclerite situated ventral to this band on each of segments IV-VII. Sternites on segments
II-VII represented by a transverse plate, which is weak or interrupted medially. Caudal
extension of segment VII small, rounded. Dorsal setae : sometimes one or two f.s. on segment
I; usually one h.s. on each side on I and each of segments V—VII, occasionally also a single
h.s. on segments II-IV. Pleural setae consisting of dorsopleural setae : o-3 (average 0:5),
0-2 (average 0-6) and o~2 (average 0-8) f.s. on segments III and VI and o-1 (average 0-7) 1, I,
I-2 (average 1-2), o-2 (average 1-3) and 1-3 (average 1-8) h.s. on segments I-VI respectively,
and of ventropleurval setae : o-1 (average 0-4), 0-3 (average 1-5) and o-5 (average 2:1) f.s. on
segments IV—VI and o-1 (average 0-4), o-1 (average 0-5), I-2 (average 1-2), 0-1 (average 0-9)
and 1 h.s. on segments II-VI respectively. Segment VII with 3-9 (average 5-4) f.s. and 1-6
(average 3-7) h.s. Ventral setae : 2-15 f.s. on each of segments II-VII ; usually 4 h.s. on each
of II-V, and 2 median ones on each of segments VI-VII.

92 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Segment VIII with small tergite and transverse sternite, the latter carrying 7-15 (average 11)
f.s. ; glandular pouch with 2 very long, pointed setae, whose protruding part is 4-6 times as
long as section within pouch. NoIXth tergite observed. Ante-anal setae : occasionally one or
two f.s. and 0-4 (average 1:5) small h.s. present. Posterior margin with 2-3 (average 2-6) f.s.
on each side.

Genital segment. Penial sheath of medium length, about + total body length (ratio 1 : 4-30—
4°86, average 4:60), 353-380 (average 371) w long and 34-40 (average 37) uw wide ; lateral
sclerotizations not joined anterior to anus ; length of basal vod about 2 that of aedeagus and
extending anteriorly from base of the aedeagus for 3—? of the distance to the apex of the basal
membranous area ; apex of sheath without membranous extension. Area from base of sheath
to tip of aedeagus with 28-44 (average 35) small setae ; a cluster of small sensilla occurring
ventrally near apex of sheath. Aedeagus long (350-380, average 371 w), penial sheath longer
and basisternum shorter, ratios being I : 1-9-2:1 (average 2) and I : 0:72-0:93 (average 0-85)
respectively.

Material examined : 10 specimens, collected by G. De Lotto on Pelargonium sp.
at Limuru, Kenya on 16.1.63.

A number of the peculiarities of this species are shared by Ceroplastodes chiton
Green, of which 3 imperfect specimens were available. The most striking of these
are: the presence of 4 large simple eyes, the numerous projections on the pro-
sternum, the absence of the median ridge of the mesobasisternum, the presence of
a pleural band of sclerotization on the abdomen and the presence of numerous f.s.
(about 18 in C. chiton) on the 8th abdominal sternite. In addition, C. chiton also
has f.s. on all the regions of the head and prothorax, on the membranous area of the
mesoscutum and all the regions of the metathorax and abdomen ; the setae of the
glandular pouch are also very long, the protruding part being about 4 times as long
as the section within the pouch.

C. chiton differs from I. theobromae in the following respects : the midcranial
ridge is reduced to a Y-shaped sclerotized area ventrally, and sometimes represented
dorsally by a short median ridge on the posterior part of the median crest ; the
postocular ridge is missing behind the ocellus ; a number of f.s. are situated beyond
the posterolateral corners of the median membranous area ; the anterior margin
of the mesopostnotum is exposed ; the tegular bulge carries a large number (more
than 5) f.s. ; the postnotum is represented by a single transverse sclerite ; the
presence of numerous (about 10) fleshy ante-anal setae ; the apical spur of the
tibia is about the same size on all 3 tibiae.

The antennae of C. chiton were not available.

The specimens of C. chiton were collected by G. Charles on Carica papaya L. in
Port Moresby, New Guinea on 23.vii.59, and received from D. J. Williams.

THE COCCUS Group
cOoccUS
Coccus hesperidum Linnaeus
(Text-figs. 31 and 32)

A short, slender species with comparatively short antennae and long legs ; with numerous
setae covering the body and appendages. When mounted, total body length 1380-1520

OF THE FAMILY COCCIDAE 93

(average 1450) uw ; width at mesothorax 280-330 (average 309) u. Wing expanse 2200-2650
(average 2446) uw.

Head subconical in dorsal view ; in lateral view obliquely elongated, with anterodorsal bulge
pronounced ; length from apex to pronotal ridge 198-239 (average 216) uw, width across genae
186-205 (average 196) uw. Median crest sclerotized and distinctly polygonally reticulated ;
with numerous fleshy dorsal head setae and 5-9 (average 6-8) hair-like ones. Midcranial ridge
dorsally absent ; ventrally narrow but well defined, reaching ocular sclerite posteriorly,
surrounding area showing distinct polygonal reticulation. Genae large, sclerotized, with poly-
gonal reticulation ; each with 19-29 (average 23) fleshy and 3-7 (average 4-9) hair-like genal
setae. Eyes : two pairs, subequal ; corneae of dorsal eyes 21-32 (average 27) u in diameter
and 1-0-2-1 (average 1-3) times as much apart ; those of the ventral eyes 22-32 (average 29) u
in diameter and 0-6-1-6 (average o-g) times as much apart. Ocellus small. Ocular sclerite
well sclerotized dorsally and laterally, but weak ventrally except for an area around the eyes ;
polygonally reticulated throughout. Preocular ridge extending only a short distance below
articular process. Postocular ridge well developed throughout ; dorsal and ventral to ocellus
the ridge usually splits up, with the anterior branch partly surrounding ocellus. Jnterocular
vidge absent, although the area between ocellus and preocular ridge is strongly sclerotized.
Dorsal ocular setae : 2-8 (average 4:9) f.s. and o—4 (average 1-6) h.s. on each side. Ventral head
setae : 75-98 (average 83) f.s. and 3-4 (average 3-8) h.s., scattered over the ocular sclerite,
always with some (9-14, average 12) f.s. occurring in front of this sclerite and with 8-16 (average

amt So

C H
J K ue
ADEH
BCFCIOR
JKL

Vp

Fic. 31. Coccus hesperidum L., dorsal and ventral view.

94 MORPHOLOGY AND TAXONOMY OF ADULT MALES

12) f.s. between and behind the eyes. Pyveoval ridge present. Tendon-like apodeme long,
Cranial apophysis long ; apex deeply bifurcate, extending to around anterior level of ventral
eyes. Mouth opening irregular. Anterior tentorial pits absent.

Antennae to-segmented, filiform ; 576-732 (average 663) u long, i.e. shorter than half body
length (ratio 1 : 2:01-2:63, average 2-26), shorter than posterior leg (ratio I : 0:58—0-74, average
0-69) and longer than penial sheath (ratio I : 2:10-2:43, average 2:27). Scape 38-46 (average
43) uw long and 27-31 (average 30) uw wide, with 3-4 (average 3:1) h.s. Pedicel with wavy stria-
tion dorsally ; 46-53 (average 50) uw long and 27-33 (average 31) uw wide ; with 6-13 (average
9:7) f.s., 2-7 (average 4-2) h.s. anda sensillum placodeum. Segment III somewhat club-shaped,
3:0-3°7 (average 3-3) times longer than wide (57-72, average 60 uw long and 19-21, average 20 u
wide) ; with 10-13 (average 11) f.s. of medium length, 1-3—1-8 (average 1-5) times longer than
width of segment ; with 1-3 usual sensilla basiconica. Segments I1V—IX cylindrical ; lengths
of these segments (in ») 84-103 (average 95), 68-91 (average 81), 49-76 (average 67), 68-84
(average 75), 53-76 (average 62) and 55-68 (average 60) respectively, widths varying from 15
to 21 uw, with distal segments slightly wider than proximal ones ; with 20-25 (average 22),
18-25 (average 22), 12-23 (average 18), 19-23 (average 20), 15-21 (average 18) and 13-20
(average 17) f.s. respectively, but no h.s. ; antennal bristles on segments VIII-IX distinctly
larger than f.s. Segment X : terminal $—} constricted ; 57-78 (average 69) u long and 49-55
(average 52) wu wide (near base) ; carrying 6-9 (average 7:5) 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
about half as long as the f.s. ; with 2 sensilla basiconica ventrally, one near apex and the other
more proximal.

Thorax 448-540 (average 516) u long.

Prothovax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, without setae. Medial pronotal setae : 2 h.s., widely separated, with
o-I (average o-4) circular pores near each. Post-tergites relatively large, with irregular wavy
striations and with 4-13 (average 8-3) fleshy post-tergital setae occurring onor be hind the sclerite
on each side. Pleural structures typical of the family. Steynum with strong transverse and
median ridges and a small oval sclerite ; the area on each side of the median ridge showing
distinct polygonal reticulation. Anéeprosternal setae : 1-4 (average 3:2) f.s. on each side ;
prosternal setae : 29-37 (average 33) f.s., scattered over the sternal area and spreading into the
area anterior to the spiracles, and usually with one h.s. (0-2, average 0-9) on each side of median
ridge.

Mesothovax. Mesoprephragma with deep emargination. Pyvescutum about twice as wide as
long (average 159 and 83 py respectively) ; anterior margin strongly curved ; laterally bounded
by the prescutal yvidges and posteriorly by the prescutal suture ; slightly more heavily sclerotized
medially ; with dense irregular reticulation. Scutum. Median membranous area subrectan-
gular, 99-125 (average 116) uv long and 1-23-1-37 (average 1-30) times as wide (width 135-160,
average 146 uw) ; with 10-32 (average 24) f.s. and 11-15 (average 13) h.s. Scutellum 34-42
(average 38) w long and 141-160 (average 154) uw wide, ratio being I : 3-8-4-2 (average 4) ;
tubular, with a small ventral foramen ; setae absent. Postnotwm with anterior margin weakly

oll
U/) 4 \
ae a)

Sooo

Fic. 32. Coccus hesperidum L., lateral view.

OF THE FAMILY COCCIDAE 95

sclerotized, irregular and exposed ; postnotal apophysis and postalare well developed, the latter
densely reticulated distally and without setae. Mesopostphragma with deep emargination.
Mesopleuron. Mesopleural ridge strong, not interrupted above coxal articulation ; pleural
apophysis and pleural wing process well developed. Basalare vestigial, incorporated into pleural
wing process, not joining the latter with episternum. Swubalare small. Episternum showing
irregular reticulation dorsally and polygonal reticulation ventrally ; subepisternal ridge well
developed, but below membranous cleft indistinct and only marked by a band of darker sclero-
tization. Epimeron small. Lateropleurite not bounded anteriorly by an extension from
marginal ridge. Basisternum large, about 182 uw wide and 147 yu long, i.e. 1-20-1-35 (average
1:27) 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
spivacle with well developed peritreme ; postmesospiraculay setae numerous, consisting of
52-75 (average 61) f.s., arranged in a broad band behind the spiracles and prosternum, and with
a few on the episternae. Tegula small, membranous bulge with a small weak sclerite posteriorly
and with 2-6 (average 4:2) h.s. Third axillary wing sclerite with a small, pointed, ventral
projection at its base. Additional sclerite small, well sclerotized. Antemetaspivacular setae :
3-4 (average 3-6) f.s.

Metathorax. Metanotum with posterior margin strong medially ; suspensorial sclerites
absent. Postnotum consisting of a small sclerite on each side. Metatergal setae : 1-5 (average
3‘1) f.s., and usually one (o—2, average 1) h.s. on each side. Pleural ridge considerably reduced,
extending only a short distance above coxal articulation. Vestigial precoval ridge present.
Episteyrnum reduced to a small subtriangular plate ; epimeron produced posteriorly. Meta-
thoracic spiracle similar to mesothoracic one. Dorsospiraculay setae : 10-18 (average 15) f.s.
and occasionally one h.s. Postmetaspiracular setae : 8-14 (average 12) f.s. and usually one
h.s. ; some f.s. occurring anterior to rest, close behind spiracles. Metasternal plate weak and
irregular, but more heavily sclerotized anteriorly. Anterior and posterior metasternal setae :
39-54 (average 45) and 12-26 (average 19) f.s. respectively.

Wings hyaline; of medium length (980-1200, average 1097 w), but comparatively broad
(width 430-520, average 481 yu), ratio width to length being 1 : 2:19-2:31 (average 2-27) ;
alay lobe and alar setae absent. Halterves absent.

Legs long and slender, with fore pair shortest and hind pair longest ; ratio length of hind leg
to body length is 1 : 1-58-1:65 (average 1-62). Length of segments (in p) :

Leg Coxa ‘Trochanter Femur Tibia Tarsus Claw Total

I 38-49 57-72 171-200 291-369 117-152 17-18 692-850
(46) (66) (190) (341) (136) (17°5) (795)

Il 69-80 76-84 163-190 293-350 127-148 17-19 749-864
(75) (81) (180) (322) (140) (18) (814)

Ill 76-84 76-87 178-203 342-395 141-160 19-20 834-943
(81) (84) (191) (369) (152) (19-5) (895)

Coxae with 10-15 (average 12) f.s. on the fore and 16-29 on the middle and hind coxa, and
each with 7-10 (average 7°8) h.s. ; fore coxa with 2-3 (average 2-3) capitate coxal bristles ;
apical seta about % as long as trochanter. Tyvochanters 19-23 w wide ; with 6 oval sensilla,
9-15 f.s. and 5-7 h.s., the latter including 2 minute setae near basal ridge, one small seta on
outer margin and one long apical seta which, on the fore trochanter, is 1-2-1-4 (average 1-3)
times as long as width of trochanter. Femora very slender, 27—34 uw wide, ratio width to length
of hind femur being 1 : 6-1-6-7 (average 6-4) ; with 27-46 f.s. and 8-14 hs. Tuibiae 17-21 u
wide, ratio width to length of hind tibia being 1 : 18-o—20-8 (average 19-1) ; each with 89-114
setae of which 17-28 are h.s. and 89-114 f.s., the latter about twice as long as width of tibia ;
apical spur about the same size on all tibiae. Tarsi 17-21 uw wide, hind tarsus 7:8-8-4 (average
8-2) times longer than wide ; with 24-42 f.s. and 13-23 h:s. ; tarsal digitules subequal, somewhat
longer than claw. Claws of medium length, a little longer than width of tarsus ; slightly curved,
with very small denticle near tip ; ungual digitules subequal, about as long as claw.

Abdomen 400-490 (average 448) u long and 230-290 (average 263) u wide.

96 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Segments I-VII : tergites present on segment II only, represented by 3 small sclerites on
anterior margin, one medially and one on each side ; sternites represented by transverse plates
on segments II, III and V-VII. Caudal extension of segment VII large, reaching or extending
beyond the level of posterior margin of segment VIII, tapering, weakly sclerotized. Dorsal
setae: up to 8, 7, 9, 7, © and 3 f.s. on segments I-VI respectively, but none on VII ; usually
one h.s. on each side on segment I and each of segments IV—VII. Pleural setae consisting of dorso-
pleural setae : 4-12 (average 8-1), 2-7 (average 4-4), 6-9 (average 7-5), 5-9 (average 6-4), 3-8
(average 4:8) and 1-6 (average 3:9) f.s. on I-VI, and o-1 (average 0-5), 0-3 (average 1-6), I-3
(average 2-4), 2-4 (average 2-4) and o—3 (average 1-6) h.s. on segments II-VI respectively, and
of ventropleural setae (sometimes difficult to separate from dorsopleural group) : usually one
f.s. on II-III and up to 5 on segments IV—VI, and usually one h.s. on segments IV-VI. Seg-
ment VII with 10-17 (average 14) f.s. and 4-6 (average 5-3) h.s. ; some of the posterior h.s.
usually longer than rest. Ventral setae: 16-21 (average 17), 14-26 (average 20), 11-25
(average 16), 12-23 (average 16), 8-17 (average 11) and 5-12 (average 9) f.s. on segments
II—VII respectively ; usually one h.s. on each side on segments III-IV and 4 on each of seg-
ments V—VII.

Segment VIII with a weak tergite and transverse sternite ; caudal extension forming a
sclerotized cylindrical lobe with a large, weakly reticulated, membranous cicatrix posteriorly
on the lobe ; glandular pouch with 2 long, pointed setae, whose protruding part is 43-6 times as
long as section within pouch. Small IXth tergite present. Ante-anal setae : usually 2 strong
h.s. Posterior margin with 2-3 (average 2-9) h.s. on each side.

Gemital segment. Penial sheath of medium length, about + total body length (ratio 1 : 4-8—
5-1, average 4:9), 274-308 (average 296) uv long and 30-34 (average 32) w wide ; lateral sclero-
tizations narrowly joined anterior to anus ; length of basal vod about 1} times that of aedeagus
and extending anteriorly from base of aedeagus for about 4—} of the distance to the apex of the
basal membranous area ; apex of sheath with a small but distinct membranous extension.
Area from base of sheath to tip of aedeagus with 15-23 (average 20) small setae ; a cluster of
small sensilla occurring ventrally near apex of sheath. Aedeagus short (57-67, average 62 u
long), penial sheath and basisternum longer, ratios being I : 4:5—5-3 (average 4:8) and 1 : 2-12—
2-80 (average 2-39) respectively.

Material examined : g specimens, 4 from Laurus nobilis L., collected inLeningrad,
USSR, on 11.vi.60 and 5 from “ citrus ’’, collected in Leningrad on 3. x1.61 ; both
samples received from N. S. Borchsenius.

Genus B. sp. (near PULVINARIA)
(Text-figs. 33 and 34)

A moderately long, slender species with comparatively short antennae and long legs ; with
numerous setae covering the body and appendages. When mounted, total body length 1960—
2100 (average 2044) uw; width at mesothorax 3440-3560 (average 3515) uw. Wing expanse
3440-3560 (average 3515) wu.

Head subconical in dorsal view ; in lateral view dorsoventrally elongated, with anterodorsal
bulge pronounced ; length from apex to pronotal ridge 289-308 (average 301) uw, width across
genae 353-385 (average 366) u. Median crest sclerotized and distinctly polygonally reticulated ;
with numerous (10-22, average 16) fleshy dorsal head setae and 6-9 (average 7-1) hair-like ones.
Midcranial ridge dorsally absent ; ventrally narrow but well defined, reaching ocular sclerite
posteriorly, surrounding area showing distinct polygonal reticulation. Genae large, sclerotized,
with polygonal reticulation ; each with 12-24 (average 17) fleshy and 3-8 (average 4-6) hair-
like genal setae. Eyes : two pairs, subequal ; corneae of dorsal eyes 27-30 (average 28) yu in
diameter and 1-5—2:3 (average 1-9) times as much apart ; those of the ventral eyes 27-30
(average 29) uw in diameter and 1-o-1-8 (average 1-4) times as much apart. Ocellus small.

OF THE FAMILY COCCIDAE 97

Ocular sclerite well sclerotized dorsally and laterally, but weak ventrally ; polygonally reticu-
lated throughout. Preocular ridge extending only a very short distance below articular process.
Postoculay ridge well developed throughout ; below ocellus the ridge splits up with the anterior
branch partly surrounding ocellus. IJntervoculay ridge absent. Dorsal ocular setae: 1-4
(average 2-3) f.s. and 1-4 (average 2-3) h.s. on each side. Ventral head setae : 45-78 (average
62) f.s. and 3-8 (average 5-4) h.s., scattered over the ocular sclerite, always with some (3-8,
average 5-4) f.s. occurring in front of this sclerite and with 1-8 (average 4-8) between and behind
the eyes (when present in small numbers situated well behind the level of the anterior margin of
the eyes). Preoral ridge present. Tendon-like apodeme long. Cranial apophysis long ; apex
deeply bifurcate, extending to around the level of the anterior margin of the ventral eyes.
Mouth opening irregular. Anterior tentorial pits absent.

Antennae to-segmented, filiform ; 979-1094 (average 1028) pw long, i.e. about half body
length (ratio 1 : 1-86—2-13, average 1-99), shorter than posterior leg (ratio 1 : 0-77—0-82, average
0-79) and longer than penial sheath (ratio 1 : 2-67—3-13, average 2:94). Scape 57-61 (average
60) wu long and 40-45 (average 42) p wide, with 3 h.s. Pedicel with distinct, polygonal, dorsal
reticulation ; 65-72 (average 70) p. long and 40—45 (average 52) p wide ; with 9-13 (average 12)
f.s. 3-6 (average 4-7) h.s. and a sensillum placodeum. Segment III somewhat club-shaped, 2-6—
3'I (average 2-8) times longer than wide (68-84, average 78 uw long and 27-30, average 28 u

: J
Vs NAN }
“Ty AW |

Ye ar yh

Fic. 33. Genus B, dorsal and ventral view.

98 MORPHOLOGY AND TAXONOMY OF ADULT MALES

wide) ; with 11-17 (average 13) f.s. of medium length, 1-6—2:3 (average 1-9) times longer than
width of segment ; with 1-3 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths
of these segments (in ) 205-228 (average 218), 137-171 (average 153), 98-129 (average 113),
95-118 (average 104), 72-91 (average 82) and 65-80 (average 73) respectively, widths varying
from 21 to 27 u, with distal segments slightly wider than proximal ones ; with 38—49 (average
42), 28-35 (average 32), 24-31 (average 27), 22-33 (average 26), 17-25 (average 21) and 15-19
(average 17) f.s. respectively, but no h.s. ; antennal bristles on segments VIII-IX distinctly
larger than f.s. Segment X : terminal $ constricted ; 72-95 (average 81) u long and 24-27
(average 25) wu wide (near base) ; carrying 4-8 (average 6-1) f.s., 3 capitate subapical setae and
5 antennal bristles of which the 3 long ones are about as long as segment and the 2 shorter ones
about as long as f.s. ; with 2 sensilla basiconica ventrally, one near apex and the other more
proximal.

Thovax 730-779 (average 755) u long.

Prothovax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclevites medium-sized, without setae. Medial pronotal setae occasionally present,
consisting of o—2 (average 0-6) f.s. and o—2 (average 0-6) h.s. ; 2-5 (average 3-6) circular pores
present on each side posterior to pronotal sclerite. Post-tergites relatively large, with irregular
wavy striation and usually with some (0-3, average 1-4) fleshy post-tergital setae on each side,
occurring on or behind the sclerite. Pleural styvuctures typical of family. Steynum with strong
transverse ridge, a strong but interrupted median ridge, and an oval-shaped sclerite. <Ante-
prosternal setae : 1-3 (average 2-4) f.s. on each side ; prosternal setae : 15-28 (average 21)
f.s., scattered over the sternal area and spreading into the area anterior to the spiracles, and
usually one (0-2, average 1) h.s. on each side of median ridge.

Mesothovax. Mesoprephvagma with deep emargination. Pvescutum about 1? times as wide
as long (average 219 and 121 yp respectively) ; anterior margin strongly curved ; laterally
bounded by the prescutal vidges and posteriorly by the prescutal suture ; slightly more heavily
sclerotized medially ; with dense irregular reticulation. Scutum. Median membranous area
subrectangular ; 152-169 (average 157) pw long and 1-40-1-53 (average 1-45) times as wide
(width 213-243, average 227 uw) ; with 7-18 (average 11) f.s. and 14-22 (average 18) hs.
Scutellum 65-72 v. long and 220-247 (average 229) u wide, the ratio being 1 : 3-1-3-8 (average
3:4) ; tubular, with small ventral foramen ; setae absent. Postnotwm with anterior margin
weakly sclerotized, and usually regular and exposed ; postnotal apophysis and postalare well
developed, the latter densely reticulated distally and without setae. Mesopostphrvagma with
deep emargination. Mesopleuron. Mesopleural ridge strong, not interrupted above coxal
articulation ; pleural apophysis and pleural wing process well developed. Basalare vestigial,
incorporated into pleural wing process, not joining the latter with episternum. Swubalave small.
Episteynum showing irregular reticulation dorsally and polygonal reticulation ventrally ;
subepisternal vidge well developed, but below membranous cleft indistinct and only marked by a
band of dark sclerotization. Epimevon small. Lateropleurite not bounded anteriorly by an
extension from marginal ridge. Basisternum large, about 268 » wide and 228 yw long, Le.
1°39-1°53 (average 1-45) times longer than membranous area of scutum ; with strong median
ridge and bounded by strong marginal and precoxal ridges ; with 2-9 (average 5-4) f.s. posteriorly

1 OV a aaah 9 pay yy Oe
a Os sf BD ee

eer | = Zoo"
= se <<) Sy =
‘ ree a ee 0

Fic. 34. Genus B, lateral view.

OF THE FAMILY COCCIDAE 99

on or near median ridge. Jurca well developed. Mesothoracic spiracle with well developed
peritreme ; postmesospiracular setae numerous, consisting of 50-87 (average 66) f.s., arranged in
a broad band behind the spiracles and prosternum, often with a few on the episternae. Tegula
small, membranous bulge with a small weak sclerite posteriorly and with 2—6 (average 3-1) h.s.
Third axillary wing sclerite with a small, pointed, ventral projection at its base. Additional
sclerite small, well sclerotized. Antemetaspivacular setae : 2-4 (average 3:1) f.s.

Metathovax. Metanotum with posterior margin usually strong medially ; suspensorial
sclerites absent ; occasionally a small, additional sclerite present anterior to postnotum. Post-
notum consisting of 2 small sclerites, one on each side. Metatergal setae : usually 2 h.s. on each
side, one laterally and one more medially, and usually with one (o—3) f.s. near the lateral h.s.
Pleural ridge considerably reduced, extending only a short distance above coxal articulation.
Vestigial precoxal ridge present. Episternum reduced to a small subtriangular plate ; epimeron
produced posteriorly. Metathoracic spiracle similar to mesothoracic one. Dorsospiraculay
setae : 9-19 (average 13) f.s. and in one case 2 h.s. Postmetaspiracular setae : 14-23 (average
18) f.s. and usually one (o—2) h.s., one or more f.s. occurring anterior to rest, close behind spiracle.
Metasternal plate weak and irregular, but more heavily sclerotized anteriorly. Anterior meta-
sternal setae : 47-78 (average 63) f.s., occasionally one h.s. ; posterior metasternal setae : 18-41
(average 27) f.s.

Wings hyaline ; of medium length (1540-1600, average 1571 uw), but comparatively broad
(720-780, average 763 uw wide), the ratio width to length being 1 : 2-03—2-14 (average 2-06) ;
alay lobe and alar setae absent. Halteres absent.

Legs long and slender, with fore pair shortest and hind pair longest ; ratio length of hind
leg to body length is 1 : 1-51~-1-63 (average 1-56). Length of segments (in py) :

Leg Coxa Trochanter Femur Tibia Tarsus Claw Total

I 65-80 76-89 304-334 498-536 194-217 25-30 1180-1286
(74) (84) (317) (516) (203) (28) (1221)

II 99-122 106-118 277-300 490-543 186-209 28-30 1203-1301
(111) (111) (289) (517) (197) (29) (1253)

Ill IOI-I14 114-125 277-304 532-597 200-213 27-31 1277-1382
(111) (117) (290) (560) (206) (30) (1314)

Coxae : 16-24 (average 19) f.s. on the fore and 26—34 on the middle and hind coxa, and each
with 6-12 h.s. ; fore coxa with 2-4 (average 2-6) capitate coxal bristles ; apical seta about }
as long as trochanter. Tvochanters 29-34 » wide ; with 6 oval sensilla ; with 8—1o (average 9)
f.s. on the fore and 14—20 on the middle and hind trochanter ; each with 6-8 h.s. which include
2 minute setae near basal ridge, one small seta on outer margin and a long apical seta which, on
the fore trochanter, is 1-7—1-:9 (average 1-8) times as long as width of trochanter. Femora
slender, 42-49 uw wide, the ratio width to length of hind femur being 1 : 5-98-6-52 (average
6:24) ; with 35-49 f.s., and with 17~—28 (average 22) h.s. on the fore and ro—18 on the middle and
hind femur. Tibiae 23-27 u wide ; ratio width to length of hind tibia being 1 : 20-6—22-4
(average 21-2) ; each with 105-141 setae of which 19-37 are h.s. and 80-107 f.s., the latter
about twice as long as width of tibia ; apical spur about the same size on all tibiae. Tarsi
20-25 uw wide, hind tarsus 8-1-9-3 (average 9) times longer than wide ; each with 30—46 f.s.
and 13-25 h.s. ; tarsal digitules subequal, slightly longer than claw. Claws of medium length,
somewhat longer than width of tarsus ; slightly curved, with small denticle near tip ; wngual
digitules subequal, about as long as claw.

Abdomen 620-700 (average 676) u long and 370-500 (average 405) uw wide.

Segments I-VII : tergites present on anterior margin of segments II-III, represented by 3
small sclerites—one medially and one on each side ; sterynites represented by a transverse plate
on segments IT, III, V-VII. Caudal extension of segment VII large, reaching level of posterior
margin of segment VIII, tapering, weakly sclerotized. Dorsal setae: usually a few (0-5,
average 2-2) f.s. on the rst and occasionally one or two on each of segments II—V ; usually one
h.s. on each side on allsegments. Pleural setae : consisting of dorsopleural setae : 5-16 (average

100 MORPHOLOGY AND TAXONOMY OF ADULT MALES

10), 4-9 (average 5-9), 4-11 (average 7-2), 4-11 (average 6-8) 2-9 (average 5) and I—5 (average
2-6) f.s. on II-VI, and o-1 (average 0-3), 1-3 (average 1-9), 2-4 (average 3:I), 2-4 (average 3-1)
and 2-5 (average 3:2) h.s. on segments III—VI respectively, and of ventropleural setae : occa-
sionally one or two f.s. on IV, 0-3 (average 1-5) on V and o-5 (average 2-4) on the VI, with
occasionally one h.s. on III and usually one on each of segments [LV-VI. Segment VII with
10-16 (average 14) f.s. and 3—6 (average 4-7) h.s., some of the posterior h.s. usually longer than
rest. Ventral setae : 17-26 (average 21), 18-28 (average 24), 10-21 (average 16), 9-15 (average
12), 4-10 (average 6-7) and 3-10 (average 6-2) f.s. on segments II—VII respectively ; usually
one h.s. on each side on segments II-IV and 4 on each of V—VII.

Segment VIII with weak tergite and transverse sternite ; caudal extension forming a genicu-
late lobe, with a large, weakly reticulated, membranous cicatrix posteriorly ; glandular pouch
with 2 long, pointed setae, whose protruding part is 2-24 times as long as section within pouch.
Small I Xth tergite present. Ante-anal setae : usually 2 strong h.s., with occasionally one or two
additional small h.s. and with 1-5 (average 2-5) small, circular pores. Posterior margin with
2-3 (average 2-8) h.s. on each side.

Genital segment. Penial sheath short, about } total body length (ratio I : 5-6-6-1, average
5°9), 334-367 (average 352) u long and 34-42 (average 38) uw wide ; lateral sclerotizations
narrowly joined anterior to anus ; length of basal vod 13-2 times that of aedeagus, extending
anteriorly from the base of the aedeagus for about ? of the distance 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 23-34 (average 28) small setae ; a cluster of small sensilla occur-
ring ventrally near apex of sheath. Aedeagus short (57-72, average 67 uw long), penial sheath
and basisternum longer, the ratios being 1 : 4:8-6-1 (average 5-3) and I : 3:11-3:93 (average
3°43) respectively.

Material examined : 10 specimens ; from leaves of unknown plant ; collected
in Goba, Ethiopia on 6.viii.62 by members of Imperial College Expedition ; identifi-
cation by K. Boratynski.

PULVINARIA
Pulvinaria ?betulae (Linnaeus)
(Text-figs. 35 and 36)

A moderately long and robust species, with comparatively short antennae and long legs ;
with numerous setae covering the body and appendages ; wings with a faint purplish tinge
between anterior margin and ist wing vein. When mounted, total body length 1800-2010
(average 1885) uw; width at mesothorax 415-460 (average 434) u. Wing expanse 3070-3320
(average 3170) wu.

Head subconical in dorsal view ; in lateral view dorsoventrally elongated, with anterodorsal
bulge pronounced ; length from apex to pronotal ridge 247-288 (average 269) uw, width across
genae 247-266 (average 257) u. Median crest sclerotized and distinctly polygonally reticulated ;
with numerous (20-34, average 28) fleshy dorsal head setae and 5-8 (average 6-7) hair-like ones.
Midcranial ridge dorsally weak and irregular ; ventrally narrow, but well defined, reaching
ocular sclerite posteriorly, area surrounding ventral part showing weak polygonal reticulation.
Genae large, weakly sclerotized and without polygonal reticulation ; each with 11-28 (average
20) fleshy and 1-5 (average 3) hair-like genal setae. Eyes : two pairs, subequal ; corneae of
dorsal eyes 30-34 (average 32) p in diameter and 1-1-1-8 (average 1-5) times as much apart ;
those of the ventral eyes 30-34 (average 32) uw in diameter and 0-6—I-o (average 0-7) times as
much apart. Ocellus small. Ocular sclevite well sclerotized dorsally, but weak ventrally ;
polygonally reticulated throughout. Pyveoculay vidge extending only a very short distance
below articular process. Postoculay ridge well developed throughout ; below ocellus the ridge
splits up, with the anterior branch partly surrounding the ocellus. Interoculay ridge absent.

OF THE FAMILY COCCIDAE IOI

Dorsal ocular setae : o—-5 (average 2:5) f.s. and o-2 (average 0:6) h.s. oneach side. Ventral head
setae : 61-82 (average 71) f.s. and 4-6 (average 4-7) h.s., scattered over the ocular sclerite,
always with some (2-6, average 4) f.s. occurring in front of this sclerite and with 13-18 (average
14) f.s. between and behind the eyes. Preoval ridge present. Tendon-like apodeme long
Cranial apophysis long ; apex deeply bifurcate, extending to around the level of the anterior
margin of the ventral eyes. Mouth opening irregular. Anterior tentorial pits absent.

Antennae 1o-segmented, filiform ; 840-901 (average 871) uv long, i.e. shorter than half body
length (ratio 1 : 2:10-2:23, aversge 2-18), shorter than posterior leg (ratio 1 : 0-87—0-92, average
o-go) and longer than penial sheath (ratio 1 : 2-30—-2-35, average 2°33). Scape 46-53 (average
47) w long and 38-46 (average 43) » wide, with 3 h.s. Pedicel : dorsal reticulation very weak
or absent ; 51-61 (average 56) uw long and 42~-49 (average 45) uw wide ; with 7-11 (average 8-6)
f.s., 4-6 (average 5:1) h.s. and a sensillum placodeum. Segment III somewhat barrel-shaped,
1-9-2-4 (average 2-1) times longer than wide (57-65, average 60 uw long and 27-30, average 29 u
wide), with 5-15 (average 9-3) f.s. of medium length, 1-3-1°5 (average 1-4) times longer than
width of segment ; with 2 or 3 usual sensilla basiconica. Segments [V-IX cylindrical ; lengths
of these segments (in w) 106-182 (average 144), 110-131 (average 119), 106-129 (average 118),
87-103 (average 95), 76-91 (average 83) and 68-84 (average 76) respectively, widths varying
from 19 to 27 u, with distal segments slightly wider than proximal ones ; with 23-44 (average
31), 25-30 (average 28), 22-32 (average 27), 20-30 (average 24), 15-20 (average 19) and 16-18
(average 18) f.s. respectively, but no h.s. ; antennal bristles on segments VIII-IX distinctly
larger than f.s. Segment X : terminal part not constricted ; 72-76 (average 74) uw long and
25-29 (average 26) uw wide ; caryring 6-11 (average 8:3) f.s., 3 capitate subapical setae and 5
antennal bristles of which the 3 long ones are almost as long as the segment and the 2 shorter

4 b —

e;

4y ' “Ah
i ee Ht
4 Aye iT }
A yy) % a
| r 3 fay ) yy) A
¥ : AWN a AS
S i { } MW

a 4 I } A
cb: \ } ae AN) Kx Nt
SM j < OWT SF Say
i ee Pee Ss) a

=A t Pe. i \\y ) y if
Q : 4¢ - = j
\

< ; Ihey Wty A .

; AW) y) 4 ;

} rue 4) N

eu 4, easly AN
Ve ‘, ap 4

vy \ Dr i
ADEHM. : 400 | ;

his > ee
—— cnc e

Pulvinaria ?betulae (L.), dorsal and ventral view.

102 MORPHOLOGY AND TAXONOMY OF ADULT MALES

ones distinctly larger than the f.s. ; with 2 sensilla basiconica ventrally, one near apex and the
other more proximal.

Thorax 650-749 (average 703) u long.

Prothovax. Pyronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, without setae. Medial pronotal setae and pores absent. Post-tergites
medium-sized, not striated, with 4-13 (average7°8) fleshy post-tergital setae occurring on and behind
the sclerite on each side. Pleural stvuctures typical of family. Steynum with strong transverse
ridge, irregular median ridge and a small triangular sclerite. Amnteprosternal setae: o-6
(average 2-9) f.s. on each side ; prosternal setae : 10-54 (average 37) f.s. and o—2 (average 1)
h.s., scattered over the sternal area and spreading into the area anterior to the spiracles.

Mesothovax. Mesoprephvagma with shallow emargination. Pyvescutum about 1} times as
wide as long (average 208 and 119 yu respectively) ; anterior margin strongly curved ; laterally
bounded by the prescutal vidges and posteriorly by the prescutal suture ; slightly more
heavily sclerotized medially, reticulation weak or absent. Scutuwm. Median membranous
avea subrectangular ; 118-125 (average 123) uw long and 1:52-1:67 (average 1°59) times as
wide (width 179-209, average 196 uw) ; with 15-28 (average 20) h.s. but no fs. Scutellum
65-68 (average 67) u long and 179-209 (average 196) u wide, the ratio being I : 3:0-3-4 (average
3:2) ; tubular, with small ventral foramen ; without setae. Postnotum with anterior margin
irregular, weakly sclerotized and either exposed or partly overlapped by metathoracic fold ;
postnotal apophysis and postalave well developed, the latter densely reticulated distally and
occasionally with a fleshy postalary seta. Mesopostphvagma with deep emargination. Meso-
pleuron. Mesopleural ridge strong, not interrupted above coxal articulation ; pleural apophysis
and pleural wing process well developed. Basalave vestigial, incorporated into pleural wing
process, not joining the latter with episternum. Subalave small. Episteyrnum without poly-
gonal reticulation ; subepisternal ridge becoming broader ventrally, but below membranous
cleft indistinct and only marked by a band of darker sclerotization. Epimeron small. Latero-
pleurite not bounded by an extension from marginal ridge. Basisternum large, about 244 u
wide and 212 yu long, i.e. 1-67—1-78 (average 1-73) 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 numerous, consisting of 58-85 (average 74) f.s. arranged in a broad band behind the
spiracles and prosternum, with a few occurring on the episternae. Tegula small, membranous
bulge with a small weak sclerite posteriorly and with 4-8 (average 6) h.s. Third axillary wing
scleyite with a small, pointed, ventral projection at its base. Additional sclerite small, well
sclerotized. Antemetaspivacular setae : 3-7 (average 4:8) f.s.

Metathovax. Metanotum with thickening of posterior margin usually desclerotized medially;
suspensorial sclevites absent. Postnotum consisting of a small subtriangular sclerite on each
side. Metatergal setae : one h.s. and usually one (0-2, average 0-9) f.s. on each side. Pleural
vidge considerably reduced, extending only a short distance above coxal articulation. Epi-
sternum reduced to a small subtriangular plate ; epimeron produced posteriorly. Metathoracic
spivacle similar to mesothoracic one. Dorsospivaculay setae : 10-19 (average 15) f.s. Post-
metaspivacular setae : 19-32 (average 25) f.s. and o—2 (average 1) h.s. Metasternal plate weak

\

‘ | \ ,
C= yy” | HN \\ 0 | Zé
ey a | ey
OG ee | TS

400p

Fic. 36. Pulvinaria ?betulae (L.), lateral view; also referable for lateral view ot P. acericola
(Walsh & Riley).

OF THE FAMILY COCCIDAE 103

and irregular, but somewhat more heavily sclerotized near anterior margin. Anterior meta-
sternal setae : 50-79 (average 68) f.s. and o—4 (average 1-6) h.s. ; posterior metasternal setae :
28-39 (average 32) f.s. and occasionally 1-2 h:s.

Wings hyaline ; of medium length (1320-1470, average 1402 u), but comparatively broad
(width 580-700, average 637 yu), the ratio width to length being 1 : 2:10-2:27 (average 2:20) ;
alar lobe and alar setae absent. Halteres absent.

Legs short and slender, with middle pair usually shortest and hind pair longest ; ratio length
of hind leg to body length is 1 : 1-94-2-03 (average 1-98). Length of segments (in pu) :

Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 61-70 80-101 201-243 346-422 137-152 27-29 827-1014
(65) (90) (227) (389) (144) (28) (942)
Il 72-89 91-99 182-213 357-410 133-148 26-28 870-977
(83) (96) (198) (383) (139) (27) (926)
Il 76-91 QI-I12 198-217 376-437 137-160 27-30 918-1034
(83) (103) (206) (403) (149) (29) (972)

Coxae : 14-24 (average 21 f.s. on the fore and 25-36 on the middle and hind coxa, and each
with 6-11 h.s. ; fore coxa with 2—4 (average 2:7) capitate coxal bristles ; apical seta about $ as
long as trochanter. Tyvochanters 26-31 uw wide ; with 6 oval sensilla, 11-22 f.s. and 6-8 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 1-5—2-4 (average 2-1) times as long as width of
trochanter. Femora of medium width (39-49 w), ratio width to length of hind femur being
I : 4:5-4°8 (average 4-6) ; with 28-48 f.s. and 7-14 h.s. Tuibiae 21-27 » wide, ratio width to
length of hind tibia being 1 : 16-5-17-7 (average 17:0) ; each with 70-115 setae of which
16-31 are h.s. and 54-89 f.s., the latter about 1? to 2 times longer than width of tibia ; apical
spur about the same size on all tibiae. Tarsi 21-23 u wide, hind tarsus 6—7 (average 6-5) times
longer than wide ; each with 23-28 f.s. and 10-18 h.s. ; tarsal digitules subequal, slightly longer
than claw. Claws of medium length, a little longer than width of tarsus ; slightly curved, with
small denticle near tip ; wngual digitules subequal, about as long as claw, with small apical
knob.

Abdomen 570-600 (average 594) uw long and 350-450 (average 410) p wide.

Segments I-VII : tergites represented by 3 small sclerites, one medially and one on each side,
on anterior margin of segments II-III, and a weak transverse plate on VII ; sterynites usually
present on all segments, represented by a weak transverse plate on segments II, III and VII,
and a weak sclerite on each side on IV-VI. Caudal extension of segment VII extending
beyond the level of posterior margin of segment VIII, tapering, weakly sclerotized latero-
ventrally. Dorsal setae : o-2 (average 0-7), o-1 (average 0-6), o-1 (average 0-3), 0-3 (average
0-9), O-I (average 0-3) and o~-2 (average 0-3) f.s. on segments I-VI respectively ; one h.s. on
each side on I and each of segments [V—VII, and very rarely a single setaon Il and III. Pleural
setae consisting of dorsopleural setae : 10-16 (average 13), 2-6 (average 4-1), 2-8 (average 4:2),
2-6 (average 4:3), 0-4 (average 2-7) and o-4 (average 2-1) f.s. on I-VI and o-1 (average o-r),
o-3 (average 1:2), 2-3 (average 2-6), 2-5 (average 3:4) and 2-6 (average 3-5) h.s. on segments
II-VI respectively, and of ventropleural setae : o-2 (average 0-6) f.s. on VI and usually one h:s.
on each of segments III-VI. Segment VII with 12-24 (average 20) f.s. and 4-12 (average 7-3)
h.s. ; some of the posterior h.s. somewhat longer than the rest. Ventral setae : 26-42 (average
36), 22-33 (average 27), 11-18 (average 15), 9-15 (average 12), 9-16 (average 12) and 9-12
(average 11) f.s. on segments II—VII respectively ; usually one h.s. on each side on II-IV, and
4 (range 3-6) on segments V—VII.

Segment VIII with weak tergite and transverse sternite, the latter carrying o—2 (average 0-7)
f.s. ; caudal extension forming a mammillate lobe, with a small, membranous cicatrix laterally;
glandular pouch with 2 long, pointed setae, whose protruding part is 24-3 times as long as
section within pouch. Small IXth tergite present. Ante-anal setae 2-7 (average 4-4) f.s. and
2 strong h.s. Posterior margin with 2-3 (average 2-9) h.s. and an occasional f.s. on each side.

104 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Genital segment. Penial sheath of medium length, about + total body length (ratio 1 : 4-9-
5°3, average 5°I), 365-388 (average 375) uw long and 40-46 (average 44) wu wide ; lateral sclero-
tizations narrowly joined anterior to anus ; length of basal vod about equal to 1} times that of
aedeagus, extending anteriorly from the base of the aedeagus for 4 to } of the distance to the
apex of the basal membranous area ; apex of sheath with a very small membranous extension.
The area from base of sheath to tip of aedeagus with 22-31 (average 25) sensilla ; a cluster of
small sensilla occurring ventrally near apex of sheath. Aedeagus short (72—-82, average 75 u
long), penial sheath and basisternum longer, the ratios being I : 4-:9-5-3 (average 5) and
I : 2:56-3:08 (average 2-83) respectively.

Material examined : 7 specimens, collected by D. J. Williams on Salix sp. in
Newcastle-on-Tyne, England during 1950.

In addition 10 specimens, received from Z. Kawecki (collected on Rzbes sp. during
August, 1962 in Warsaw, Poland) as Pulvinaria Pribesiae Sign., were examined.

There are no distinct structural differences between the males of these two samples,
and the various measurements, ratios and setal counts all overlapped. The descrip-
tion given above is based on the specimens from England. The characteristics of
the specimens from Poland can be summarized as follows:

Total body length 1760-2160 (average 1963 uw). Head: Dorsal head setae 26-51 (average
34) f.s. and 5-10 (average 7:9) h.s. ; dorsal ocular setae 0-7 (average 3-6) f.s. and 0-3 (average
0-9) h.s. ; ventral head setae 54-108 (average 84) f.s. and 5-7 (average 6-1) h.s. of which 5-9
(average 6:6) f.s. occur in the area anterior to the ocular sclerite and 17-25 (average 21) between
and behind the ventral eyes ; genal setae 18-30 (average 23) f.s. and 2-9 (average 4:6) hs. ;
antennae 809-999 (average 887) u long, length in relation to body length 1 : 2:08-2:37 (average
2°23).

Thorax. Medial pronotal setae absent ; post-tergital setae o—8 (average 2-3) f.s. on each side;
prosternal setae 22-50 (average 33) f.s. and 1-3 (average 2-1) h.s. ; median membranous area
of scutum 122-141 (average 128) uw long and 205-239 (average 223) uw wide (ratio I : 1-62-1-88,
average 1°74), with 14-30 (average 21) h.s. ; fleshy postmesospiracular setae 67-84 (average
76) ; antemetaspiracular setae 3-9 (average 5:2) f.s. ; dorsospiracular setae 9-19 (average 13)
f.s. ; postmetaspiracular setae 16-28 (average 23) fs. and o-4 (average 1-8) h.s. ; anterior
and posterior metasternal setae 53-89 (average 79) and 37-54 (average 44) f.s. respectively ;
wings 1340-1440 (average 1410) w long and 610-670 (average 642) w wide ; front coxa with
2-4 (average 3) knobbed coxal bristles. Abdomen. Pleural setae on segment VII 13-29
(average 22) f.s. and 3-8 (average 5:6) h.s. ; fleshy ventral setae on segments II-VII 33-56
(average 41), 23-49 (average 35), 18-26 (average 20), 13-27 (average 17) 10-17 (average 14)
and 5-16 (average 11) respectively ; penial sheath 353-384 (average 367) u long, length in
relation to body length 1 : 5:15-5:63 (average 5-41) ; aedeagus 72-82 (average 77) u long.

Pulvinaria acericola (Walsh & Riley)
(Text-figs. 37 and 36)

A moderately long, robust species with comparatively short antennae and legs; with
numerous setae covering the body and appendages ; wings with a faint purplish tinge between
the anterior margin and first wing vein. When mounted, total body length 1670-1890 (average
1786) uw; width at mesothorax 420-480 (average 458) u. Wing expanse 2840-3140 (average
2978) wu.

Head subconical in dorsal view ; in lateral view obliquely elongated, with anterodorsal bulge
pronounced ; length from apex to pronotal ridge 228-293 (average 260) p, width across genae
251-266 (average 257) wu. Median crest sclerotized and distinctly polygonally reticulated ;
with numerous (19-43, average 30) fleshy dorsal head setae and 1-7 (average 5-2) hair-like ones.

OF THE FAMILY COC€CIDAE 105

Midcranial ridge dorsally weak and irregular ; ventrally narrow, but well defined, reaching
ocular sclerite posteriorly, area surrounding ventral part showing weak polygonal reticulation.
Genae large, weakly sclerotized, but showing distinct polygonal reticulation ; each with 16-31
(average 23) fleshy and 1-9 (average 4:2) hair-like genal setae. Eyes : two pairs, subequal ;
corneae of dorsal eyes 27-34 (average 31) uw in diameter and 1-7—2-4 (average 1-9) times as much
apart ; those of the ventral eyes 27-34 (average 30) » in diameter and 0-9-1-4 (average 1-1)
times as much apart. Ocellus small. Ocular sclerite well sclerotized dorsally, but weak ven-
trally ; polygonally reticulated throughout. Preoculay ridges extending only a very short
distance below articular process. Postoculay ridge well developed throughout ; below ocellus
the ridge splits up with the anterior branch partly surrounding the ocellus. Interocular ridge
absent. Dorsal ocular setae : o—6 (average 2-3) f.s. and o—2 (average 0-5) h.s. on each side.
Ventral head setae : 70-90 (average 80) f.s. and 4~—7 (average 4:8) h.s., scattered over the ocular
sclerite, always with some (4-7, average 4°8) f.s. occurring in front of this sclerite and with
10-15 (average 12) f.s. between and behind the eyes. Preorval ridge present. Tendon-like
apodeme long. Cyranial apophysis long ; apex deeply bifurcate, extending to around the anterior
margin of the ventral eyes. Mouth opening irregular. Anterior tentorial pits absent.

Antennae 1o-segmented, filiform ; 733-828 (average 794) uv long, i.e. shorter than half body
length (ratio 1 : 2:08-2-45, average 2-24), shorter than posterior leg (ratio 1 : 0-78-o-81,
average 0-80) and longer than penial sheath (ratio 1 : 2-14-2-35, average 2:20). Scape 46-53
(average 49) vp long and 34-46 (average 41) uw wide, with 3 h.s. Pedicel with distinct, polygonal,
dorsal reticulation ; 53-65 (average 61) uw long and 38-46 (average 43) w wide ; with 1o-15
(average 12) f.s., 3-5 (average 4) h.s. and a sensillum placodeum. Segment III somewhat
barrel-shaped, 2-1-2-3 (average 2-2) times longer than wide (57-72, average 64 w long and
23-29, average 26 uw wide) ; with 9-19 (average 12) f.s. of medium length, 1-6—1-8 (average
1-7) times longer than width of segment ; with 1 to 3 usual sensilla basiconica. Segments [V-IX

: z AY:
PSP ak... My WY

¢ 4 ° wy b,
\ } 4

iy

Fic. 37. Pulvinaria acericola (Walsh & Riley), dorsal and ventral view.

106 MORPHOLOGY AND TAXONOMY OF ADULT MALES

cylindrical ; lengths of these segments (in w) 110-133 (average 122), 95-133 (average 114),
87-118 (average 106), 80-122 (average 94), 57-72 (average 66) and 53-65 (average 59) respec-
tively, widths varying from 21-29 p, with distal segments slightly wider than proximal ones ;
with 19-31 (average 25), 23-36 (average 29), 20-35 (average 27), 21-27 (average 24), 17-25
(average 22) and 14-21 (average 17) f.s. respectively, but no h.s. ; antennal bristles on segments
VIII-IX distinctly larger than f.s. Segment X: terminal part not constricted ; 53-68
(average 59) uw long and 22-29 (average 23) uw wide ; carrying 6—13 (average Io) 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 4 as long as the f.s. ; with two sensilla basiconica ventrally, one
near apex and the other more proximal.

Thorax 635-741 (average 687) u long.

Prothovax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, without setae. Medial pronotal setae and pores absent. Post-tergites
medium-sized, with irregular wavy striations and usually with some o-7, (average 3-9) fleshy
post-tergital setae occurring on or behind the sclerite on each side. Pleural structures typical of
family. Steynum with strong transverse ridge, weak and interrupted median ridge, and a small
triangular sclerite. Anteprosternal setae : 2-7 (average 3-6) f.s. on each side ; prosternal
setae : 7-19 (average 14) f.s. and occasionally one h.s., scattered over the sternal area and
spreading into the area anterior to the spiracles.

Mesothovax. Mesoprephragma with shallow emargination. Pyrescutuwm about 1? times as
wide as long (average 218 uw and 119 w respectively) ; anterior margin strongly curved ; laterally
bounded by the prescutal ridges and posteriorly by the prescutal suture ; slightly more heavily
sclerotized medially ; showing polygonal reticulation, which sometimes tends to be irregular.
Scutum. Median membranous avea subrectangular ; 99-139 (average 124) w long and 1-53-
2-04 (average 1-76) times as wide (width 201-228, average 212 u) ; with 6-20 (average 14)
h.s. and rarely 2 f.s. Scutellum 65-72 (average 69) p long and 213-247 (average 231) w wide,
ratio being 1 : 3:0-3:8 (average 3-4) ; tubular, with moderately large ventral foramen ;
without setae. Postnotum with anterior margin irregular, weakly sclerotized and either exposed
or partly overlapped by metathoracic fold ; postnotal apophysis and postalarve well developed,
the latter densely reticulated distally and without setae. Mesopostphragma with deep emargina-
tion. Mesopleuron. Mesopleural ridge strong, not interrupted above coxal articulation ;
pleural apophysis and pleural wing process well developed. Basalare vestigial, incorporated
into pleural wing process, not joining the latter with episternum. Swbalave small. Episternum
usually showing weak polygonal reticulation dorsally ; subepisteynal ridge well developed, but
below membranous cleft indistinct and only marked by a band of dark sclerotization.
Epimeron small. Lateropleurite not bounded anteriorly by an extension from marginal ridge.
Basisternum large, about 287 u. wide and 214 yu long, i.e. 1-45—-2-00 (average 1-74) 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 spivacle with well
developed peritreme ; postmesospivaculay setae numerous, consisting of 55-79 (average 66) f.s.,
arranged in a broad band behind the spiracles and prosternum, with a few occurring on the
episternae. Tegula small, membranous bulge with a small weak sclerite posteriorly and with
2-7 (average 5) h.s. Third axillary wing sclerite with a small, pointed, ventral projection at its
base. Additional sclerite small, wellsclerotized. Antemetaspivacular setae : 1-7 (average 2-9) f.s.

Metathovax. Metanotum with thickening of posterior margin usually desclerotized medially;
suspensorial sclerites absent. Postnotum consisting of a small transverse sclerite on each side.
Metatergal setae : one h.s. and usually a few (0-6, average 2-4) f.s. on each side. Pleural ridge
considerably reduced, extending only a short distance above coxal articulation. Episternum
reduced to a small subtriangular plate ; epimevon produced posteriorly. Metathoracic spiracle
similar to mesothoracic one. Doyrsospivacular setae : 10-23 (average 16) f.s. Postmetaspivacular
setae : 12-25 (average 16) f.s. and o-2 (average 0-8) h.s. Metasteynal plate weak and irregular,
but more heavily sclerotized anteriorly. Anterioy and posterioy metasternal setae consisting of
58-94 (average 77) and 34-49 (average 38) f.s. respectively.

OF THE FAMILY COCCIDAE 107

Wings : hyaline ; of medium length (1230-1350, average 1292 uw) but comparatively broad
width (660-690, average 670 yw), ratio width to length being 1 : 1:86-1:97 (average 1-93) ; alar
lobe and alay setae absent. Halteres absent.

Legs moderately long and slender, with the fore pair usually shortest and the hind pair
longest ; the ratio length of hind leg to body length is 1 : 1-76-1-85 (average 1-79). Length
of segments (in p) :

Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 61-72 87-99 205-243 369-422 118-133 19-25 864-998
(65) (95) (227) (391) (126) (24) (928)
II 80-91 99-114 194-220 372-418 118-133 23-25 890-991
(86) (108) (208) (391) (125) (24) (941)
IIL 84-91 106-116 193-228 399-450 125-144 25-30 942-1056
(87) (112) (211) (422) (135) (27) (995)

Cowae : 11-24 (average 18), 18-29 (average 23) and 22-30 (average 26) f.s. on the fore, middle
and hind coxa respectively, and each with 7-11 h.s. ; fore coxa with 3—5 (average 4) capitate
coxal bristles ; apical seta about $ as long as trochanter. Tyvochanters 30-38 uw wide ; with 6
oval sensilla ; with 11-16 (average 13) f.s. on the fore and ro—22 on the middle and hind tro-
chanters ; each with 5-8 h.s., the latter including 2 or 3 minute setae near basal ridge, one small
seta on outer margin and a long apical seta which, on the fore trochanter, is 1-6—1-8 (average
1-7) times as long as the width of trochanter. Femorva of medium width (41—53 w), the ratio
width to length of hind femur being 1 : 3:9-4-6 (average 4:2) ; each with 28-47 f.s. and 8-16
h.s. Tuibiae 24-30 uw wide, the ratio width to length of hind tibia being 1 : 13-5-16-6 (average
14°9) ; each with 90-118 setae of which 14~—22 are h.s. and 72-100 f.s., the latter about 14-2
times longer than width of tibia ; apical spur about the same size on all tibiae. Tarsi 21-27 uv
wide, hind tarsus 4:3—5-:2 (average 5) times longer than wide ; each with 22-32 f.s. and 5-11
h.s. ; tarsal digitules subequal, slightly longer than claw. Claws of medium length, length
about equal to width of tarsus ; slightly curved, with small denticle near tip ; ungual digitules
subequal, about as long as claw.

Abdomen 450-570 (average 538) vp long and 400-450 (average 427) p wide.

Segments I-VII : tergites on segmenst II-III represented by 3 small sclerites on anterior
margin—one medially and one on each side, on segment IV by a small sclerite on each side, and
on VII by a weak transverse plate ; sternites represented by a weak transverse plate on segments
II, 111 and VII, and sometimes by a small weak sclerite on each side on [LV and VI. Caudal
extension of segment VII large, extending beyond level of posterior margin of segment VIII,
tapering, weakly sclerotized lateroventrally. Dorsal setae : a few (up to 4) f.s. occasionally
present ; usually one h.s. on each side on segment I and each of [V—VII but none on II and III.
Pleural setae consisting of dorsopleural setae : o-8 (average 4-4), 1-8 (average 3°5), 0-9 (average
3°3), 0-9 (average 3-6), 4-13 (average 6-4) and 1-9 (average 3-9) f.s. on segments I-VI respec-
tively and o-2 (average 0-3), 0-2 (average 1), 1-5 (average 2-8), o-3 (average 2-4) and 1-5
(average 2-6) h.s. on II-VI and of ventropleural setae : one or two f.s. occasionally present on
segments III-IV, and 0-6 (average 2) and 0-6 (average 2-4) on V—VI respectively, with usually
one h.s. on each of segments [V-VI. Segment VII with 9-24 (average 15) f.s. and 4—6 (average
4°8) h.s. ; some of the posterior h.s. somewhat longer than rest. Ventral setae : 22-42 (average
33), 20-37 (average 28), 8-22 (average 16), 5-19 (average 12), 5-16 (average 10), 3-6 (average
4°6) f.s. on segments II—VII respectively ; usually no h.s. on II, one on each side on III-IV and
4 (range 3-6) on each of segments V—VII.

Segment VIII with weak tergite and transverse sternite ; caudal extension forming a mam-
millate lobe with a small, membranous cicatrix laterally ; glandular pouch with 2 long, pointed
setae, whose protruding part is 2-3 times as long as section within pouch. Small IXth tergite
present. Ante-anal setae : usually a few (0-5, average 1-2) f.s. and 2 (0-2, average 1-3) strong
h.s. present. Posterior margin with 2-3 (average 2-7) h.s. on each side.

Genital segment. Penial sheath of medium length, about + total body length (ratio 1 : 4-8-
5°2, average 5), 342-369 (average 357) uv long and 38-53 (average 44) u wide ; lateral sclerotiza-

108 MORPHOLOGY AND TAXONOMY OF ADULT MALES

tions narrowly joined anterior to anus ; length of basal vod equal to about 1} times that of
aedeagus, extending anteriorly from the base of the aedeagus for }—? of the distance to the apex
of the basal membranous area ; apex of sheath with a very small membranous extension.
Area from base of sheath to tip of aedeagus with 24-35 (average 28) small setae ; a cluster of
small sensilla occurring ventrally near apex of sheath. Aedeagus short (76-95, average 85 wu
long), penial sheath and basisternum longer, the ratios being 1 : 3-6—-4:8 (average 4-3) and
I : 2:26-2:95 (average 2°59) respectively.

Material examined : 9g specimens ; collected by T. E. Tabor on Acer saccharinum
L. in Hillsville, Virginia, U.S.A. on 6.v.63 ; received from M. Kosztarab, identified
by K. Boratynski.

P. acericola can readily be separated from P. ?betulae by the 2 short antennal
bristles on antennal segment X and also by the distinct polygonal reticulation on
the pedicel, gena and mesoprescutum.

PARTHENOLECANIUM
Parthenolecanium corni (Bouché)
(Text-figs. 38 and 40)

A medium-sized and moderately robust species, with comparatively short antennae and long
legs ; with numerous setae on the body and appendages. When mounted, total body length
1880-2110 (average 1980) » ; width at mesothorax 390-500 (average 446) u. Wing expanse
3030-3400 (average 3283) uw.

Head subconical in dorsal view ; in lateral view obliquely elongated dorsoventrally, with
anterodorsal bulge pronounced ; length from apex to pronotal ridge 239-293 (average 273) u,
width across genae 258-289 (average 275) uw. Median crest sclerotized and distinctly poly-
gonally reticulated ; with 23-37 (average 28) fleshy and 16-19 (average 17) hair-like dorsal head
setae. Muidcranial ridge dorsally absent ; ventrally narrow but well defined, reaching ocular
sclerite posteriorly, surrounding area showing distinct polygonal reticulation. Genae large,
sclerotized, polygonally reticulated ; each with 17-30 (average 23) fleshy and 7-13 (average
g‘1) hair-like genal setae. Eyes : two pairs, subequal ; corneae of dorsal eyes 34—42 (average
38) w in diameter and 1-2-1-7 (average 1:5) times as much apart ; those of the ventral eyes
34-42 (average 38) u in diameter and 0-5—-0°8 (average 0:7) times as much apart. Ocellus small.
Ocular sclerite well sclerotized and polygonally reticulated throughout. Preoculay ridge extend-
ing only a short distance below articular process. Postoculay ridge usually well developed
throughout, usually with a weak anterior branch below ocellus. Intevoculay ridge absent.
Dorsal ocular setae : 1-7 (average 4:1) f.s. and o—4 (average 2) h.s. on each side. Ventval head
setae : 65-83 (average 70) f.s. and 7-12 (average 9) h.s., scattered over the ocular sclerite,
occasionally with one or two f.s. occurring in front of the sclerite, and with 12-17 (average 15)
f.s. and o-2 (average 0-7) h.s. between and behind the eyes. Preoval ridge present. Tendon-
like apodeme long. Cranial apophysis of medium length ; apex bifurcate, without central bulge,
not reaching the level of anterior margin of ventral eyes. Mouth opening irregular. Anterior
tentorial pits apparently absent.

Antennae to-segmented, filiform ; 825-998 (average 895) uw long, i.e. shorter than half body
length (ratio I : 2-06—2-41, average 2-23), shorter than posterior leg (ratio 1 : 0-77—0-89, average
0-82) and longer than penial sheath (ratio 1 : 1:89-2-10, average 1:95). Scape 49-61 (average
55) wu long and 42-53 (average 48) uw wide, with 3 h.s. Pedicel with distinct, polygonal, dorsal
reticulation ; 61-68 (average 64) uw long and 42-49 (average 46) uw wide ; with 7-11 (average
9°3) f.s., 5-8 (average 6) h.s. and a sensillum placodeum. Segment III club-shaped, 2-1-2-4
(average 2-3) times longer than wide (61-84, average 66 » long and 27-34, average 28 wu wide) ;

OF THE FAMILY COCCIDAE 109

with 10-15 (average 13) f.s. of medium length, 1-1-1-4 (average 1-2) times longer than width of
segment ; with 1-4 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths of these
segments (in uw) 171-220 (average 192), 103-141 (average 114), 99-113 (average 107), 80-103
(average 90), 65-84 (average 73) and 53-68 (average 59) respectively, all of about the same
width, varying from 21 to 29 uw ; with 38-67 (average 53), 31-46 (average 36), 32-54 (average
36), 27-38 (average 32), 22-30 (average 25) and 17-25 (average 20) f.s., but no h.s. ; antennal
bristles on segments VIII-IX distinctly larger than f.s. Segment X : terminal 4+ somewhat
constricted ; carrying 7-13 (average 9-4) 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 somewhat
shorter and thicker than the f.s. ; with 2 sensilla basiconica ventrally, one near apex and the
other more proximal.

Thorax 669-790 (average 733) u long.

Prothovax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, occasionally with one fleshy lateral pronotal seta on each side. Medial
pronotal setae absent. Post-tergites comparatively large, with irregular, wavy striations ; post-
tergital setae absent. Pleural structures typical of the family. Steynum with strong transverse
ridge, interrupted median ridge and small triangular sclerite. Anteprosternal setae: 1-3
(average 2-1) f.s. on each side ; prosternal setae 19-30 (average 26) f.s., scattered over the sternal
area and spreading into the area anterior to the spiracles.

Mesothorax. Mesoprephragma with shallow emargination. Prescutum less than twice as wide
as long (average 223 and 127 yp respectively) ; anterior margin strongly curved ; laterally
bounded by the prescutal ridges and posteriorly by the prescutal suture ; slightly more heavily

ADEHM
DEFGIQ.
JKL__

Fic. 38. Parthenolecanium corni (Bouché), dorsal and ventral view.

Tee) MORPHOLOGY AND TAXONOMY OF ADULT MALES

sclerotized medially ; polygonally reticulated. Scutwm. Median membranous area sub-
rectangular ; 118-133 (average 128) u long and 1-53~1-82 (average 1-61) times as wide (width
182-236, average 206 u) ; with 11-26 (average 20) h.s., but nof.s. Scutellum 65-87 (average
74) w long and 198-247 (average 220) uw wide, ratio being I : 2-9-3-3 (average 3) ; tubular ;
ventral foramen small, its length usually less than half that of scutellum ; without setae.
Postnotum with anterior margin weakly sclerotized, irregular, and usually not overlapped by
metathoracic fold ; polygonally reticulated dorsally ; postnotal apophysis and postalare well
developed, the latter densely reticulated distally and without setae. Mesopostphragma with
deep emargination. Mesopleuron. Mesopleural vidge strong, not interrupted above coxal
articulation ; pleural apophysis and pleural wing process well developed. Basalare vestigial,
incorporated into pleural wing process, not joining the latter with episternum. Subulave small,
Episternum showing polygonal reticulation dorsally ; subepisternal ridge well developed, but
below membranous cleft indistinct and only marked by a band of dark sclerotization.
Epimeron small. Lateropleurite not bounded anteriorly by an extension from marginal ridge.
Basisternum : large, about 270 w wide and 214 up long, i.e. 1-50-1-82 (average 1-68) 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 numerous, consisting of 71-98 (average 89) f.s.
and o-2 (average o-g) h.s., arranged in a broad band behind the spiracles and prosternum.
Tegula small, membranous bulge with a small weak sclerite posteriorly and with 6-8 (average
7:2) hs. Third axillary wing sclerite with a small ventral projection at its base. Additional
sclerite small, well sclerotized. Antemetaspivacular setae : 4-12 (average 6:5) f.s.

Metathovax. Metanotum with posterior margin desclerotized medially ; suspensorial sclerites
absent ; a small, additional sclerite sometimes present anterior to postnotum. Postnotum
consisting of a small, transverse sclerite on each side. Metatergal setae : o—2 (average 0-8) f:s.
and o-2 (average 1) h.s. on each side. Pleural ridge considerably reduced, extending only a
short distance above coxal articulation. Vestigial precoxal ridge usually present. Episternum
reduced to a small plate ; epimeron produced posteriorly. Metathoracic spivacle similar to
mesothoracic one. Dorsospivaculay setae: 5-14 (average 10) f.s. Postmetaspivacular setae :
21-35 (average 30) f.s. and 0-2 (average 0-9) h.s. Metasternal sclerite represented by a weak
transverse plate. Awnterioy and posterioy metasternal setae : 76-94 (average 86) and 31-60
(average 42) f.s. respectively.

Wings hyaline ; short (length 1330-1510, average 1439 w) but comparatively broad (width
620-740, average 686 yu), the ratio width to length being 1 : 2:04—2-15 (average 2-10) ; alar
lobe and alay setae absent. Halteves absent.

Legs long and slender, with fore pair shortest and hind pair longest ; ratio length of hind leg
to body length is 1 : 1-74—1-84 (average 1-81). Length of segments (in uy) :

Leg Coxa Trochanter Femur Tibia Tarsus Claw Total

I 72-80 99-118 247-285 395-452 125-141 24-29 984-1091
(76) (108) (271) (429) (132) (26) (1042)

Il 87-95 106-118 224-262 418-471 137-143 27-30 1003-1113
(92) (112) (244) (444) (140) (28) (1062)

Ill 95-107 106-122 232-266 441-479 137-152 27-32 1051-1146
(102) (117) (251) (465) (144) (29) (1109)

Coxae : 17-24 (average 20) f.s. on the fore and 23-43 on the middle and hind coxa, and each
with 10-19 h.s. ; fore coxa with 3-4 (average 3-5) capitate coxal bristles ; apical seta about
4-3 as long as trochanter. Tvochanters 29-34 up. wide ; with 6 oval sensilla, 12-21 f.s. and 8-11
h.s., the latter including 2 minute setae near basal ridge, one small seta on outer margin and one
long apical seta which, on the fore trochanter, is 2-6—3-4 (average 3) times as long as width of
trochanter. Femova of medium width (42-53 uw), ratio width to length of hind femur being
I : 4°8-5°7 (average 5:2) ; with 36-54 f.s. and 15-27 h.s. Tuibiae 23-30 u wide, ratio width to
length of hind tibia being 1 : 15:0-19-1 (average 16-4) ; each with 107-153 setae, of which
30-47 are h.s. and 76-110 f.s., the latter about as long as width of tibia ; apical spur about the

OF THE FAMILY COCCIDAE II

same size on all tibiae. Tarsi 21-27 uw wide, with 13-28 f.s. and 16—26 his. ; tarsal digitules
subequal, somewhat longer than claw. Claws of medium length, a little longer than width of
tarsus ; slightly curved, with small denticle near tip ; wngual digitules subequal, about as long
as claw.

Abdomen 520-600 (average 570) uv. long and 39g0—500 (average 425) vu 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 ; sternites represented by a weak transverse
sclerite on segments II, III, sometimes VI, and VII. Caudal extension of segment VII large,
reaching or extending beyond the level of posterior margin of segment VIII, tapering, weakly
sclerotized. Dorsal setae : occasionally one f.s. on segments I-II ; usually one h.s. on each
side on I and each of segments IV-VII. Pleural setae consisting of dorsopleural setae : 1-7
(average 3-4), 1-6 (average 2-5), I—4 (average 2-2), 2-6 (average 3-9), I-6 (average 3-1) and 0-3
(average 1-4) f.s., and 0-2 (average 0-9), o-1 (average 0-1), O-1 (average 0-3), I-4 (average 1-8),
0-5 (average 3-3), and 2-5 (average 3-9) h.s. on segments I-VI respectively, and of ventropleural
setae : O-3 (average 1-1) f.s. on VI and usually one h.s. on each of segments [V-VI. Segment
VII with 10-17 (average 15) f.s. and 5~12 (average 7-3) h.s. ; some of the posterior h.s. usually
longer than the rest. Ventral setae : 30-38 (average 34), 13-23 (average 20), 8-18 (average 15),
5-11 (average 7-6), 5-11 (average 7-8) and 4-9 (average 6-5) f.s. on segments II—VII respectively ;
usually one h.s. on each side on segments III-IV and 4 on each of V-VII.

Segment VIII with weak tergite and transverse sternite ; caudal extension forming a small,
sclerotized, papilla-shaped lobe with a small, membranous cicatrix laterally ; glandular pouch
with 2 long, pointed setae, whose protruding part is 2}—3 times as long as section within pouch.
Small IXth tergite present. Ante-anal setae: 2 strong h.s. which are sometimes forked.
Posterior margin with 3 h.s. on each side.

Genital segment. Penial sheath long, about } total body length (ratio 1 : 4-19-4°44, average
4°38), 456-486 (average 470) uw long and 45~49 (average 46) uw wide at base of aedeagus ; lateral
sclerotizations not joined anterior to anus ; length of basal rod about % or equal that of aedeagus,
extending anteriorly from base of the aedeagus for about 4 of the distance to apex of the basal
membranous area ; apex of sheath without membranous extension. Area from base of sheath
to tip of aedeagus with 20-30 (average 25) small setae ; a cluster of small sensilla occurring near
apex of sheath. Aedeagus short (106-125, average 114 u long), penial sheath and basisternum
longer, the ratios being 1: 3:9-4:3 (average 4:1) and 1: 1:79-2:67 (average 1-99) re-
spectively.

Material examined : 8 specimens, collected by Z. Kawecki on Ribes aurewm
Pursh. and Ribes sp. in Warsaw, Poland during January, 1962.

In addition, ten specimens were examined from Belgrade, Yugoslavia (received
from N. Miti¢é-MuZina, collected on an unknown host during 1961) and two from
Alma-Ata, Kazakhstan, USSR (received from G. Matesova, collected in fruit
orchards on 20.iv.51). These specimens were all smaller than the specimens from
Warsaw, consequently the measurements of the various structures and the numbers
of setae are somewhat reduced ; otherwise they are practically identical. On the
other hand, the specimens that Habib (1956) described from Cotoneaster microphylla
Lindl., collected in Wisley, England, were bigger (2100—2300, average 2200 u long).
It is worthy of note that the females of this species also show considerable size
variation, due mainly to the influence of the host plant (Habib, 1953 and Kawecki,

1958a).

The variation in the specimens from Poland, Yugoslavia and the USSR can be
summarized as follows (all measurements in uy) :

112 MORPHOLOGY AND TAXONOMY OF ADULT MALES

VARIATION IN P. coyni FROM THREE LOCALITIES

Characters Localities
Warsaw Belgrade Alma-Ata
Total length 1880-2110 1470-1770 1650-1850
(1980) (1660) (1750)
Wing expanse 3030-3400 2470-3110 2950-3130
(3283) (2856) (3040)
HEAD:
Width across genae 258-289 217-274 251-258
(275) (250) (255)
Dorsal head setae: fleshy 23-37 19-46 18-23
(28) (29) (21)
Dorsal head setae: hair-like 16-19 10-22 19-21
(17) (15) (20)
Genal setae: fleshy 17-30 16-30 21-25
(23) (22) (23)
Genal setae: hair-like 7-13 7-12 IO-II
(9) (9)
Diameter of dorsal eyes 34-42 23-34 30-31
(38) (28)
Diameter of ventral eyes 34-42 23-36 34
(38) (29)
Ventral head setae: fleshy 65-83 57-92 69-76
(70) (72) (73)
Ventral head setae: hair-like 7-12 3-7 4
(9) (5)
Antennal length 825-998 659-827 906
(895) (740)
Antennal length in relation to body
length I :2:06-2-41 I : 2:12-2°33 E 32-04
(2-23) (2-24)
THORAX
Prosternal setae: fleshy 22-39 18-40 13-28
(31) (28) (20)
Median membranous area of scutum:
length: 118-133 84-118 103
(128) (ror)
Median membranous area of scutum:
width: 182-236 135-194 179-190
(206) (173) (184)
scutal setae (hair-like) 11-26 10-22 13-16
(20) (15) (15)
Scutellum: length 65-87 46-65 65-76
(74) (57) (70)
Scutellum: width 198-247 144-205 182-205
(220) (184) (194)
Basisternum: length 194-236 156-201 213
(214) (182)

OF THE FAMILY COCCIDAE 113

Characters Localities
Warsaw Belgrade Alma-Ata
Basisternum: width 233-304 194-262 266-277
(270) (237) (272)
Postmesospiracular setae (fleshy) 71-98 44-85 73-80
(89) (63) (77)
Tegular setae (hair-like) 6-8 4-9 7-9
(7:2) (5-7) (8-0)
Antemetaspiracular setae (fleshy) 4-12 2-5 5
(6°5) (3-1)
Dorsospiracular setae (fleshy) 5-14 5-15 7
(10) (10)
Postmetaspiracular setae (fleshy) 21-35 15-34 18
(39) (19)
Anterior metasternal setae: fleshy 76-94 40-82 76
(86) (57)
Posterior metasternal setae: fleshy 31-60 18-45 39-50
(42) (29) (45)
Wing length 1330-1510 1100-1380 1310-1400
(1439) (1260) (1355)
Wing width 620-740 480-650 650-710
(686) (598) (680)
Ratio width to length I: 2°04-2°15 1: 1°10-2°31_ I : 1:97-2:02
(2-10) (2-18) (2-00)
Hind leg: length 1051-1146 832-1098 1064-1139
(1109) (973) (1102)
length in relation to body length 1 :1-74-1-84 _ I : 1:72-1-77 I : 1°62
(1-81) (1-75)
Tibial setae 107-153 81-123 IOI—127
ABDOMEN
Pleural setae on segment VII: fleshy 10-17 II-25 II-I4
(15) (20) (12)
Ventral setae on segment II: fleshy 30-38 13-42 26-31
(34) (25) (29)
Penial sheath: length 456-486 361-410 410-429
(479) (394) (420)
length in relation to body length 1 : 4:19-4:44 =I : 3°91-4°39_—siI_ :: 3°84-4°51
(4°38) (4°23) (4°28)
Aedeagus: length 106-125 QI-I14 114
(114) (100)

Parthenolecanium pomeranicum (Kawecki)
(Text-figs. 39 and 40)
A medium-sized and moderately robust species, with comparatively short antennae and long
legs ; with numerous setae on the body and appendages. When mounted, total body length

1610-1800 (average 1690) uw ; width at mesothorax 405-450 (average 418) uw. Wing expanse
2590-2910 (average 2750) uw.

114 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Head subconical in dorsal view ; in lateral view obliquely elongated dorsoventrally, with
anterodorsal bulge pronounced ; length from apex to pronotal ridge 243-277 (average 261) u,
width across genae 247-281 (average 265) wu. Median crest well sclerotized and distinctly poly-
gonally reticulated ; with 15-32 (average 24) fleshy and 10-17 (average 13) hair-like dorsal head
setae. Muidcranial ridge dorsally absent ; ventrally narrow but well defined, reaching ocular
sclerite posteriorly, surrounding area showing distinct polygonal reticulation. Genae large,
sclerotized, polygonally reticulated ; each with 14-24 (average 19) fleshy and 6-13 (average
6-9) hair-like genal setae. Eyes : two pairs, subequal ; corneae of dorsal eyes 23-27 (average
25) win diameter and 2-7—3-3 (average 3) times as much apart ; those of the ventral eyes 23-30
(average 26) pw in diameter and 1-I-1-5 (average 1-3) times as much apart. Ocellus small.
Ocular sclerite well sclerotized and polygonally reticulated throughout. Pyreoculay ridge extend-
ing only a short distance below articular process. Postoculay ridge well developed dorsally and
lateroventrally, but usually weak posteromedially ; below ocellus the ridge usually has a weak
anterior branch. Interoculary ridge absent. Dorsal ocular setae : 2-8 (average 4-2) f.s. and
o-4 (average 1-6) h.s. on each side. Ventral head setae: 44-60 (average 51) f.s. and 8-15
(average 10) h.s., scattered over the ocular sclerite, occasionally with a fleshy seta occurring in
front of the sclerite, and with 9-16 (average 13) f.s. and o-1 (average 0-6) h.s. between and
behind the eyes. Pveoval ridge present. Tendon-like apodeme long. Cranial apophysis of
medium length ; apex bifurcate, sometimes with central bulge, not reaching level of anterior
margin of ventral eyes. Mouth opening irregular. Anterior tentorial pits absent.

ADEHM
BCFGIQ
JKL

Fic. 39. Parthenolecanium pomeranicum (Kaw.), dorsal and ventral view.

OF THE FAMILY COCCIDAE 115

Antennae 1o-segmented, filiform ; 686-758 (average 730) u long, i.e. shorter than half body
length (ratio 1 : 2:20-2:54, average 2-39), shorter than posterior leg (ratio I : 0-72—0-74, average
0-73) and longer than penial sheath (ratio 1 : 1-92—2-09, average 2-01). Scape 49-57 (average
53) w long and 40-44 (average 42) w wide, with 3 h.s. Pedicel with distinct, polygonal, dorsal
reticulation ; 49-57 (average 53) wu long and 42-46 (average 43) uw wide ; with 3-8 (average
51) f.s., 5-9 (average 6-9) h.s. and a sensillum placodeum. Segment III : club-shaped, 2-0—2-9
(average 2-3) times longer than wide (53-63, average 59 u long and 21~—27, average 26 wu wide) ;
with 8-15 (average 13) f.s. of medium length, 1-o—-1-4 (average 1-2) times longer than width of
segment ; with 1-4 usual sensilla basiconica. Segments IV-IX : cylindrical ; lengths of these
segments (in wu) 95-122 (average 113), 84-114 (average 97), 93-99 (average 95), 65-91 (average
75), 57-78 (average 66) and 46-57 (average 52) respectively, all of about the same width, varying
from 19 to 27 w; with 21-33 (average 30), 22-36 (average 32), 28-43 (average 36), 23-37
(average 27), 19-30 (average 21) and 16—23 (average 19) f.s. respectively, but no h.s. ; antennal
bristles on segments VIII-IX distinctly larger than the fleshy setae. Segment X : terminal
4 somewhat constricted ; carrying 10-15 (average 12) f.s., occasionally one h.s., 3 capitate
subapical setae and 5 antennal bristles of which the 3 long ones are just more than half as long as
the segment and the 2 shorter ones somewhat shorter and thicker than the f.s. ; with 2 sensilla
basiconica ventrally, one near apex and the other more proximal.

Thorax 638-688 (average 660) u long.

Prothovax. Pyronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, with o-3 (average 1-1) fleshy and o-—3 (average 0-5) hair-like lateral
pronotal setae on each side. Medial pronotal setae : usually one h.s. and occasionally one f.s.
on each side. Post-tergites comparatively large, with irregular wavy striations ; post-tergital
setae absent. Pleural structures typical of family. Steynwm with transverse ridge strong,
median ridge weak and only represented anteriorly, and a well sclerotized triangular sclerite.
Anteprosternal setae : 0-4 (average 1-3) f.s. on each side ; prosternal setae : 13-21 (average 15)
f.s. and sometimes one h.s., scattered over the sternal area and spreading into the area anterior
to the spiracles.

Mesothorvax. Mesoprephragma with shallow emargination. Prescutum less than twice as
wide as long (average 115 and 201 yu respectively) ; anterior margin strongly curved ; laterally
bounded by the prescutal ridges and posteriorly by the prescutal suture ; slightly more heavily
sclerotized medially ; polygonally reticulated. Scutwm. Median membranous area subrectan-
gular ; 103-118 (average 111) w long and 1-50~-1-78 (average 1-60) times as wide (width 175-—
182, average 178 w) ; with 14-22 (average 19) h.s., but no f.s. Scutellum 53-61 (average 58) u
long and 175-182 (average 178) u wide, ratio being 1 : 2-9—3°8 (average 3-3) ; tubular, witha
large ventral foramen of which the length is half or more than half that of the scutellum ;
without setae. Postnotum with anterior margin weakly sclerotized, irregular, and partly
overlapped by the metathoracic fold ; postnotal apophysis and postalare well developed, the
latter densely reticulated distally and without setae. Mesopostphragma with deep emargination.
Mesopleuron. Mesopleural ridge strong, not interrupted above coxal articulation ; pleural

Fic. 40. Parthenolecanium pomeranicum (Kaw.), lateral view; also referable for lateral
view of P. corni (Bouché).

116 MORPHOLOGY AND TAXONOMY OF ADULT MALES

apophysis and pleural wing process well developed. Basalare vestigial, incorporated into pleural
wing process, not joining the latter with episternum. Subalave small. Episteynum showing
polygonal reticulation dorsally ; subepisternal ridge well developed, but below membranous
cleft indistinct and only marked by a band of dark sclerotization. Epimevon small. Latero-
pleunite not bounded anteriorly by an extension from marginal ridge. Basisteynum large, about
240 pv. wide and 179 yu long, i.e. 1-48-1-74 (average 1-61) times longer than membranous area of
scutum ; with strong median ridge and bounded by strong marginal and precoxal ridges ;
without setae. Fuvca well developed. Mesothovacic spivacle with well developed peritreme ;
postmesospivaculay setae numerous, consisting of 54-64 (average 57) f.s. and 2-5 (average 3:8)
h.s., arranged in a broad band behind the spiracles and prosternum. Tegula small, membranous
bulge with a small weak sclerite posteriorly and with 3-9 (average 5-4) h.s. Third axillary wing
sclevite with a small ventral projection at its base. Additional sclerite small, well sclerotized.
Antemetaspivacular setae : 2-8 (average 5-3) f.s.

Metathovax. Metanotum with posterior margin desclerotized medially ; suspensorial sclerites
absent. Postnotum consisting of a small transverse sclerite on each side. Metatergal setae :
o-3 (average 0-6) f.s. and 1-3 (average 1-3) h.s. on each side. Pleural ridge considerably
reduced, extending only a short distance above coxal articulation. Vestigial precoxval ridge
usually present. Episteyrnum reduced to a small plate ; epimevon produced posteriorly.
Metathovacic spivacle similar to mesothoracic one. Dorsospivacular setae : 3-11 (average 7-8)
f.s. and o-3 (average 1) h.s. Postmetaspivacular setae : 14-24 (average 18) f.s. and 1-4 (average
2:2) h.s. Metasternal sclervite represented by a weak, transverse plate. Anterior and posterior
metasteynal setae : 56-74 (average 64) and 20—32 (average 28) f.s. respectively.

Wings hyaline; short (length 1130-1290, average 1210 uw) but comparatively broad (width
520-590, average 557 uw), ratio width to length being I : 2:15-2:20 (average 2-17) ; alar lobe
and alar setae absent. Halteves absent.

Legs long and slender, with fore pair shortest and hind pair longest ; ratio length of hind leg
to body length is 1 : 1:69-1:84 (average 1-77). Length of segments (in y) :

Leg Coxa Trochanter Femur Tibia Tarsus Claw Total
I 61-72 76-87 222-241 338-372 118-141 24-27 868-918
(67) (81) (231) (355) (130) (25) (890)
Il 76-87 84-91 201-224 353-380 133-148 25-27 882-948
(81) (87) (209) (366) (140) (26) (972)
Ill 76-95 80-99 209-228 372-410 141-163 26-29 934-1017
(87) (92) (216) (391) (150) (27) (963)

Cowxae : 12-16 (average 13) f.s. on the fore and 24—32 on the middle and hind coxa, and each
with 13-20 h.s. ; fore coxa with 2—5 (average 3-8) coxal bristles ; apical seta about ? as long as
trochanter. Tvochanteys 28-32 » wide ; with 6 oval sensilla ; with 9-12 (average 11) f.s. on
the fore and 14-18 on the middle and hind trochanter, and each with 8-14 h.s., the latter
including 2 minute setae near basal ridge, one small seta on outer margin and one long apical
seta which, on the fore trochanter, is 2:2—2-8 (average 2-5) times as long as width of trochanter.
Femova of medium width (42-49 wu), ratio width to length of hind femur being I : 4-5—4:9
(average 4:6) ; each with 26—42 f.s. and 20-37 h.s. Tzbiae 23-30 wu wide, ratio width to length
of hind tibia being 1 : 12:3-16-3 (average 14:2) ; each with 120-150 setae of which 44—64 are
h.s. and 68-92 f.s., the latter about as long as width of tibia ; apical spur about the same size
on all tibiae. Tarsi 19-25 w wide, hind tarsus 6:2—-8-6 (average 7) times longer than wide ;
each with 8-19 f.s. and 27-37 h.s. ; tarsal digitules subequal, about as long as 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 440-550 (average 491) uw long and 410-530 (average 460) uw wide.

Segments I-VII : tergites represented by a small sclerite on each side on anterior margin of
segments II-IIland byaweak transverse plateon VII; stevnites represented by a weak transverse
sclerite on segments II, III and VII. Caudal extension of segment VII large, reaching or

OF THE FAMILY COCCIDAE 117

extending beyond the level of posterior margin of segment VIII, tapering, weakly sclerotized.
Dorsal setae : occasionally one or two f.s. on the 1st segment ; usually one h.s. on each side on
I and each of segments IV-VII. Pleural setae consisting of dorsopleural setae : 1-5 (average
2:1), 0-4 (average 1°5), O-5 (average 1-7), 1-4 (average 2-6), o-3 (average 2) and o-4 (average
1-4) f.s., and o-1 (average 0-2), 0-2 (average 0-9), 0-3 (average 1-5), I-4 (average 2-8), I-3
(average 2-1) and o~—2 (average 1-6) h.s. on segments I-VI respectively, and ventropleural setae :
occasionally one f.s. on segments V—VI, sometimes one h.s. on IV and usually one on each of
segments V-VI. Segment VII with 4-8 (average 5-4) f.s. and 4—8 (average 5-9) h.s. ; some of
the posterior h.s. usually longer than the rest. Ventral setae : 9-18 (average 13), 6-16 (average
II), 9-14 (average I1), 5-I1 (average 7-3), 4-11 (average 7:1) and 1-4 (average 2-7) f.s. on
segments II—-VII respectively ; usually one h.s. on each side on segments III-IV and 4 on each
of V-VII.

Segment VIII with weak tergite and transverse sternite ; caudal extension forming a large,
papilla-shaped lobe with a small, membranous cicatrix laterally ; glandular pouch with 2 long,
pointed setae, whose protruding part is about twice as long as section within pouch. Small IXth
tergite sometimes present. Ante-anal setae: 2 strong h.s., which are sometimes forked.
Posterior margin with 2—3 (average 2-8) h.s. on each side.

Genital segment. Penial sheath of medium-length, about 4 total body length (ratio 1 : 4-40—
5°09, average 4°77), 342-361 (average 353) uw long and 36-42 (average 38) uw wide ; lateral
sclerotizations apparently not joined anterior to anus ; basal rod about 4—% as long as aedeagus,
extending anteriorly from the base of the aedeagus for about }—} of distance to apex of the basal
membranous area ; apex of sheath without membranous extension. Area from base of sheath to
tip of aedeagus with 20~30 (average 25) sensilla ; a cluster of small sensilla occurring ventrally
near apex of sheath. Aedeagus short (76-95, average 84 u long), penial sheath and basisternum
longer, the ratios being 1 : 3-8—4-5 (average 4-2) and 1 : 1-84—2-25 (average 2-10) respectively.

Material examined : 10 specimens, collected by M. S. K. Ghauri on Taxus baccata
L. at Imperial College Field Station, Silwood Park, Sunninghill, Berks., on 5.v.57.

This species is very closely related to P. corni (Bouché). The two species can be
separated by the number of fleshy pleural setae on the caudal extension of segment
VII. InP. pomeranicum these setae vary from 4-8 (average 5) and in P. corni they
vary from II-I4 (average 12), 10-17 (average 15) and I1-—25 (average 20) in the
specimens from Kazakstan, Warsaw and Belgrade respectively. In addition, the
pair of hair-like medial pronotal setae was absent in only one specimen of P.
pomeranicum, whereas only one medial pronotal seta was present in one specimen
of P. corn (from Warsaw). The ventral foramen of the scutellum is large in P.
pomeranicum, its length being half or more than half that of the scutellum, while in
P. corm it is usually very small, although it was more than half as long as the
scutellum in one specimen from Belgrade. The posteromedian part of the post-
ocular ridge is usually weak in P. pomeranicum, but strong in P. cornt.

CEROPLASTES
Ceroplastes berliniae (Hall)
(Text-figs. 41 and 42)
A short, robust species, with comparatively short antennae and moderately long legs ; with
numerous setae covering the body and appendages. When mounted, total body length 1450—

1610 (average 1529) uw; width at mesothorax 360-400 (average 372) u. Wing expanse 2170-
2310 (average 2277) wu.

118 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Head subconical in dorsal view ; in lateral view dorsoventrally elongated, with anterodorsal
bulge not pronounced ; length from apex to pronotal ridge 201-232 (average 217) u, width
across genae 201-224 (average 213) uw. Median crest sclerotized, showing distinct polygonal
reticulation which enclose irregular striations ; with numerous (15-27, average 21) fleshy
dorsal head setae and 4-10 (average 8-1) hair-like ones. Mudcranial ridge dorsally absent ;
ventrally strong, reaching ocular sclerite posteriorly, surrounding area weakly polygonally
reticulated. Genae large, sclerotized, with polygonal reticulation ; each with 9-18 (average
14) fleshy and 2-7 (average 4-4) hair-like genal setae. Eyes : two pairs, subequal ; corneae of
dorsal eyes 23-27 (average 25) uw in diameter and 1-8—2-7 (average 2-2) times as much apart ;
those of the ventral eyes 25-27 (average 26) up in diameter and o-6—1-o (average 0-7) times as
much apart. Ocellus small. Ocular sclerite well sclerotized and polygonally reticulated
throughout. Pyreoculay vidge extending only a very short distance below articular process.
Postoculay ridge well developed throughout ; dorsal and ventral to ocellus the ridge usually
splits up with the anterior branch partly surrounding ocellus. Intevoculay ridge absent.
Dorsal ocular setae : o-4 (average 2) f.s. and occasionally one h.s. on each side. Ventral head
setae consisting of 47-67 (average 53) f.s. and 3-8 (average 4-4) h.s., scattered over the ocular
sclerite, always with some (2-7, average 4-8) f.s. occurring in front of this sclerite and with

5 ANS th
OH

ee Wage Ne Ss
SASS r >
hay”
AW i
hy

Fic. 41. Cevoplastes berlimae Hall, dorsal and ventral view.

OF THE FAMILY COCCIDAE 119

5-9 (average 7-1) between and behind the eyes. Preoral ridge present. Tendon-like apodeme
long. Cranial apophysis of medium length ; apex deeply bifurcate, extending to around level
of posterior margin of ventral eyes. Mouth opening irregular. Anterior tentorial pits absent.

Antennae to-segmented, filiform ; 541-673 (average 622) wu long, i.e. shorter than half body
length (ratio I : 2:39-2:54, average 2-46), shorter than posterior leg (ratio I : 0-68-0-74,
average 0-71) and longer than penial sheath (ratio 1 : 1-43-1-64, average 1°55). Scape 38-42
(average 40) » long and 34~—40 (average 37) » wide, with 3 h.s. Pedicel with distinct, polygonal,
dorsal reticulation ; 48-53 (average 50) uw long and 27-30 (average 29) wp wide; with 6-8
(average 7-1) f.s., 3-5 (average 4-3) h.s. and a sensillum placodeum. Segment III somewhat
club-shaped, 2:2—2-8 (average 2-5) times longer than wide (44-53, average 49 u long and 19-23,
average 20 uw wide) ; with 4-8 (average 6-2) f.s. of medium length, 1-o-1-4 times longer than
width of segment ; with 1-4 usual sensilla basiconica. Segments IV-IX cylindrical ; length of
these segments (in yw) 80-103 (average 90), 68-89 (average 78), 46-65 (average 57), 60-76
(average 65), 53-63 (average 58) and 46-57 (average 52) respectively, widths varying from 15 to
23 uw, with distal segments slightly wider than proximal ones ; with 11-19 (average 15), 13-22
(average 17), 9-17 (average 13), 12-20 (average 15), 12-14 (average 13), 11-15 (average 13) f.s.
respectively, but no h.s. ; antennal bristles on segments VIII-IX distinctly larger than f.s.
Segment X : terminal } constricted ; 74-93 (average 82) u long and 20-23 (average 21) wp wide
(near base) ; carrying 5-7 (average 5-8) 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. (but stouter) ; with 2 sensilla basiconica ventrally, one near apex and the other more
proximal.

Thorax 498-555 (average 540) u long.

Prothovax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, without setae. Medial pronotal setae : occasionally one f.s. present
behind pronotal sclerite. Post-tergites relatively large, with irregular striations and without
setae. Pleural structures typical of family. Steynum with strong transverse ridge, median ridge
represented by a weak basal stalk, and triangular sclerite narrow and very weakly sclerotized.
Anteprosternal setae : 2-7 (average 4:3) f.s. on each side, with usually one f.s. situated dorsal to
proepisternum + cervical sclerite ; prosternal setae : 27-39 (average 32) f.s., scattered over the
sternal area and often spreading into the area anterior to the spiracles.

Mesothovax. Mesoprephragma with moderately deep emargination. Pyrescutwm more than
twice as wide as long (average 197 and 85 u respectively) ; anterior margin curved ; laterally
bounded by the prescutal ridges and posteriorly by the prescutal suture ; medially with slightly
more heavy sclerotization becoming ridge-like anteroventrally ; with distinct polygonal reticul-
tion. Scutum. Median membranous area subrectangular ; 99-112 (average 103) yu long and
1-69-1'94 (average 1-80) times as wide (width 175-199, average 188 uw) ; with 10-20 (average
14) fs. and 4-14 (average 10) h.s. Scutellum 53-65 (average 60) u long and 190-209 (average
200) uw wide, the ratio being 1 : 3-0-3°8 (average 3:4) ; tubular, with small ventral foramen ;
without setae. Postnotum reticulated, with anterior margin irregular and exposed ; postnotal
apophysis and postalare well developed, the latter densely reticulated distally and with 1-3

Fic. 42. Cevoplastes berliniae Hall, lateral view; also referable for lateral view of Ceroplastes sp.

120 MORPHOLOGY AND TAXONOMY OF ADULT MALES

(average 1-7) fleshy postalavy setae occurring on or immediately posterior to it. Mesopost-
phragma with deep emargination. Mesopleuron. Mesopleural ridge strong, not interrupted
above coxal articulation ; pleural apophysis and pleural wing process well developed. Basalare
absent. Subalave small. Episteynum distinctly polygonally reticulated ; subepisternal ridge
well developed, but below membranous cleft indistinct and only marked by a band of dark
sclerotization. Epimevon small. Lateropleurite partly bounded anteriorly by an extension
from marginal ridge. Basisternum large, about 223 uw wide and 171 yw long, i.e. 1-48-1-77
(average 1:65) 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 spivacle with well developed peritreme ; postmesospirvacular setae : 18-29 (average
24) f.s., arranged in a band behind the spiracles and prosternum. Tegula small, membranous
bulge with a small weak sclerite posteriorly and with 4-8 (average 6-3) h.s. Third axillary wing
scleyite with a pronounced ventral projection at its base. Additional sclerite small. Antemeta-
spivaculay setae : 1-5 (average 2-7) f:s.

Metathorax. Metanotum with thickening of posterior margin desclerotized medially ;
suspensorial sclevites absent. Postnotum consisting of two small sclerites, one on each side.
Metatergal setae : 2-7 (average 3-6) f.s. on each side but no h.s. Pleural ridge considerably
reduced, extending only a short distance above coxal articulation. Vestigial precoval ridge
present. Episteynum reduced to a small subtriangular plate ; epimevon produced posteriorly.
Metathoracic spivacle similar to mesothoracic one. Dorsospivaculay setae : 3-9 (average 5-5)
f.s. Postmetaspivacular setae : 12-16 (average 14) f.s. and o—2 (average 0-6) h.s. Metasternal
plate weak and irregular, but more heavily sclerotized anteriorly. Anterioy and posterior
metasternal setae : 36-48 (average 41) and 18—29 (average 23) f.s. respectively.

Wings hyaline; short (length 920-980, average 968 uw) and comparatively broad (width 450—
480, average 465 w), the ratio width to length being 1 : 2-00—2-18 (average 2-08) ; alar lobe and
alay setae absent. Halteres absent.

Legs moderately long, and slender, with fore pair shortest and hind pair longest ; ratio length
of hind leg to body length is 1 : 1-75-1-86 (average 1-79). Length of segments (in yp) :

Leg Coxa Trochanter Femur Tibia Tarsus Claw Total

i 49-61 68-76 186-205 289-325 106-122 21-23 724-798
(54) (73) (197) (310) (113) (22) (770)

II 65-74 80-89 158-175 304-338 112-125 20-23 744-813
(70) (84) (166) (319) (119) (22) (780)

Ill 76-84 89-95 175-186 338-365 122-133 22-24 825-876
(80) (92) (180) (352) (127) (23) (854)

Coxae : with 8-13 (average 11), 13-22 (average 16) and 20—29 (average 24) f.s., and 6—10
(average 8-2), 9-14 (average 11) and 14-17 (average 15) h.s. on the fore, middle and hind coxa
respectively ; fore coxa without coxal bristles ; apical seta about # as long as trochanter.
Trochanters 23-34 p. wide ; with 6 oval sensilla ; with 9-14 f.s. on the fore and middle trochanter
and 15-24 (average 19) on the hind one, and each with 6-10 h.s., the latter including 2 minute
setae near basal ridge, one small seta on the outer margin and a long apical seta which, on the
fore trochanter, is 1:32-1:77 (average I-52) times as long as the width of trochanter. Femora of
medium width (37-45 uw), with ratio width to length of hind femur I : 4-1-4:4 (average 4:2) ;
with 24-35 f.s.and 8-16h.s. Tibiae 19-27 up wide ; ratio width to length of hind tibia 1 : 12-8—
15-9 (average I14°3) ; each with 81-111 setae of which 21-30 are h.s. and 58-83 f.s., the latter
about 14 times longer than width of tibia ; apical spur about the same size on all tibiae. Tarsi
17-21 u wide, hind tarsus 6-4—6:6 (average 6-5) times longer than wide ; each with 17-27 f.s.
and 6-11 hs. ; tarsal digitules subequal, slightly 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 380-450 (average 409) wu long and 330-380 (average 345) uw wide.

Segments I-VII : tergites represented by a small transverse sclerite on each side on anterior
margin of segments II-III and a weak transverse plate on VI-VII ; stervnites represented by a

OF THE FAMILY COCCIDAE 121

weak transverse plate on segments II, III, Vl and VII. Caudal extension of segment VII large,
reaching the level of the posterior margin of segment VIII, tapering and weakly sclerotized.
Dorsal setae : up to 4, 5, 6, 3, 2 and 1 f.s. on segments I-VI respectively, but none on VII ;
occasionally a single h.s. present on segments I-III, and usually one on each side on each of
segments IV-VII. Pleural setae consisting of dorsopleural setae: o-3 (average 0-7), 0-3
(average 1-3), 0-3 (average 1-7), I-3 (average 2), o-3 (average 1-6) and o-4 (average 1) f.s. on
I-VI, and o~2 (average 0-9), 2, o-2 (average 1-6) and 1-2 (average 1-7) h.s. on segments III-VI
respectively, and of ventropleural setae : occasionally one (range o-2) f.s. on II, usually 2
(range 0-3) on III, and 3 (range 1~5) on each of segments IV—VI, and usually one h.s. on each of
segments IV-VI. Segment VII with 9-15 (average 12) f.s. and 3—4 (average 3-2) h.s. ; some of
the posterior h.s. usually longer than rest. Ventral setae : 21-28 (average 25), 17-25 (average
21), 11-18 (average 14), 7-12 (average 10), 5-8 (average 7) and 1-5 (average 3) f.s. on the
segments II—VII respectively ; usually one h.s. on each side on III—-VI and 4 on VII.

Segment VIII with weak tergite and transverse sternite ; caudal extension forming a
prominent, weakly sclerotized, semi-circular lobe with a large cicatrix posterodorsally, the latter
weakly sclerotized and reticulated in the middle ; glandular pouch with 2 long, pointed setae,
whose protruding part is 3—4 times as long as section within pouch. Small [Xth tergite present.
Ante-anal setae : o-3 (average 1-9) f.s. Posterior margin with 2-6 (average 3-8) f.s. and 2-3
(average 2-9) h.s. on each side.

Genital segment. Penial sheath long, about 3—} total body length (ratio 1 : 3-6—4-1, average
3°8), 395-410 (average 4:2) uw long and 38-42 (average 40) uw wide ; lateral sclerotizations
narrowly joined anterior to anus ; length of basal rod about equal to that of aedeagus, extending
anteriorly from base of the aedeagus for about % of the distance to apex of the basal membranous
area ; apex of sheath with a distinct, finger-like membranous extension. Area from base of
sheath to tip of aedeagus with 17-22 (average 20) small setae ; a cluster of small sensilla occur-
ring ventrally near apex of sheath. Aedeagus of medium length (118-137, average 125 u long),
penial sheath and basisternum longer, the ratios being 1 : 3-0-3-4 (average 3-1) and 1 : 1-28—
1°52 (average 1-39) respectively.

Material examined : 10 specimens, collected on Brachystegia tamarindoides Welw.

by R. Boulton in the Salisbury district, [Southern] Rhodesia on 11.xii.62 ; received
from J. Munting ; identified by D. J. Williams.

Ceroplastes sp.
(Text-figs. 43 and 42)

A short, robust species, with comparatively short antennae and long legs ; with numerous
setae covering the body and appendages. When mounted, total body length 1430-1670
(average 1529) uw; width at mesothorax 370-410 (average 386) uy. Wing expanse 2320-2500
(average 2417) wu.

Head subconical in dorsal view ; in lateral view dorsoventrally elongated, with anterodorsal
bulge not pronounced ; length from apex to pronotal ridge 205-247 (average 225) wu, width
across genae 213-251 (average 231) uw. Median crest sclerotized, showing distinct, polygonal
reticulation which does not enclose irregular striations ; with numerous (16-27, average 22)
fleshy dorsal head setae and 7-11 (average 8-1) hair-like ones. Midcranial ridge dorsally repre-
sented by a short, vestigial ridge on anterior margin of head ; ventrally strong, reaching ocular
sclerite posteriorly, surrounding area weakly polygonally reticulated. Genae large, sclerotized,
with polygonal reticulation ; each with 11-19 (average 15) fleshy and 2-7 (average 4-3) hair-like
genal setae. Eyes: two pairs, subequal ; corneae of dorsal eyes 27-30 (average 29) w in
diameter and 1-6—2-6 (average 2-1) times as much apart ; those of the ventral eyes 29-32
(average 30) uw in diameter and o-6—1-o (average o-8) times as much apart. Ocellus small.
Ocular sclerite well sclerotized, polygonally reticulated throughout. Preoculay ridge extending

122 MORPHOLOGY AND TAXONOMY OF ADULT MALES

a very short distance below articular process. Postoculay ridge well developed throughout ;
ventral to ocellus the ridge usually splits up, with the anterior branch partly surrounding
ocellus. Interoculay ridge absent. Dorsal ocular setae : 1-4 (average 2-5) f.s. and 1-5 (average
2:8) h.s. Ventral head setae : 55-84 (average 71) f.s. and 4-8 (average 6-3) h.s., scattered over
the ocular sclerite, always with some (3-6, average 2-3) f.s. occurring in front of the sclerite and
with 11-21 (average 16) between and behind the eyes. Preoval ridge present. Tendon-like
apodeme long. Cranial apophysis of medium length ; apex bifurcate, extending to around
level of posterior margin of ventral eyes. Mouth opening irregular. Anterior tentorial pits
absent.

Antennae 1o-segmented, filiform ; 678-770 (average 727) uv long, i.e. shorter than half body
length (ratio I : 2:02-2:24, average 2:10), shorter than posterior leg (ratio 1 : 0:72-0:78,
average 0-75) and longer than penial sheath (ratio 1 : 1-59-1-80, average 1-71). Scape 42-46
(average 43) uw long and 38—42 (average 39) wu wide, with 3 h.s. Pedicel occasionally with poly-
gonal reticulation dorsally, but usually with only wavy striations ; 46-53 (average 50) u long
and 34-47 (average 38) uw wide ; with 8-17 (average 12) f.s., 4-6 (average 5-4) h.s. and a sen-
sillum placodeum. Segment II] somewhat club-shaped, 2-6—3-2 (average 3) times longer than
wide (59-72, average 69 u long and 21-25, average 23 u wide) ; with 8-15 f.s. of medium length,
1:2-1'4 (average 1:3) times longer than width of segment ; with 1-3 usual sensilla basiconica.
Segments IV-IX cylindrical ; lengths of these segments (in wu) 106-137 (average 120), 9I-105
(average 98), 68-80 (average 74), 68-87 (average 79), 49-65 (average 58) and 53-65 (average
59) respectively, widths varying from 15 to 23 wu, with distal segments slightly wider than

2 ( Ze CG }
Sg A
a “Ss 1
Ge
we JS ~ [i
\ Se ; i
3 al ~ Rhy
” + SH A
aan
OE,
Weck df
Q
Ye

ADEHM.
BCFGIQ

J K L ie

JKL

Fic. 43. Cevoplastes sp., dorsal and ventral view.

OF THE FAMILY COCCIDAE 123

proximal ones ; with 23-29 (average 26), 20-27 (average 23), 13-24 (average 19), 17-23
(average 21), 14-20 (average 17) and 12-17 (average 15) f.s. respectively, but no h.s. ; antennal
bristles on segments VIII-IX distinctly larger than f.s. Segment X : terminal 4 constricted ;
65-84 (average 78) uw long and 19-23 (average 21) uw wide (near base) ; carrying 2-11 (average
7:8) 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 f.s. (but stouter) ; with 2 sensilla
basiconica ventrally, one near apex and the other more proximal.

Thorax 498-593 (average 531) pv long.

Prothovax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral
pronotal sclerites small, without setae. Medial pronotal setae : occasionally one f.s. behind
pronotal sclerites. Post-tergites relatively large, with irregular striations, without setae.
Pleural structures typical of family. Steynum with transverse ridge strong, median ridge
represented by a weak basal stalk, and triangular sclerite narrow and weakly sclerotized.
Anteprosternal setae : 2-5 (average 3:5) f.s. on each side, with no setae occurring dorsal to
proepisternum + cervical sclerite ; prosternal setae : 14-24 (average 19) f.s. and occasionally
one h.s., scattered over the sternal area but not spreading into the area anterior to the spiracles.

Mesothovax. Mesoprephragma with moderately deep emargination. Prescutum about twice
as wide as long (average 196 and 96 u respectively) ; anterior margin curved ; laterally bounded
by the prescutal ridges and posteriorly by the prescutal suture ; medially with somewhat more
heavy sclerotization becoming ridge-like anteroventrally ; with distinct polygonal reticulation.
Scutum. Median membranous area subrectangular ; 80-99 (average 91) u long and 1:75-2:14
(average 1-99) times as wide (width 160-209, average 180 1), with 2-11 (average 6-7) f.s. and g-12
(average 11) h.s. Scutellum 46-65 (average 55) uw long and 175-198 (average 188) uw wide, the
ratio being 1 : 3:1-4:1 (average 3:5) ; tubular, with small ventral foramen ; without setae.
Postnotum reticulated, with anterior margin irregular and exposed ; postnotal apophysis and
postalave well developed, the latter densely reticulated distally and with 1-6 (average 3-3)
fleshy postalary setae occurring on or immediately posterior to it. Mesopostphragma with
moderately deep emargination. Mesopleuron. Mesopleural ridge strong, not interrupted above
coxal articulation ; pleural apophysis and pleural wing process well developed. Basalare
absent. Subalare small. Episternum distinctly polygonally reticulated ; subepisternal ridge
well developed, but below membranous cleft indistinct and only marked by a band of darker
sclerotization. Epimeron small. Lateropleurite partly bounded anteriorly by an extension
from marginal ridge. Basisteyrnum large, about 232 uw wide and 158 yw long, i.e. 1-60-1-88
(average 1-74) 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 spivacle with well developed peritreme ; postmesospivacular setae : 14-29 (average
21) f.s. and occasionally one h.s., arranged in a group behind each spiracle and a small number
posterior to prosternum. Tegula small, membranous bulge with a small weak sclerite posteriorly
and with 3-10 (average 5-7) h.s. Third axillary wing sclerite with a pronounced ventral projec-
tion at its base. Antemetaspiracular setae : 2-7 (average 4:1) f.s.

Metathorax. Metanotum with thickening of posterior margin desclerotized medially ;
suspensorial sclerites absent. Postnotum consisting of two small sclerites, one on each side.
Metatergal setae : 1-6 (average 2-9) f.s. and sometimes up to 3 (0-3, average 0-8) h.s. of which one
is occasionally situated more medially than the others. Pleural ridge considerably reduced,
extending only a short distance above coxal articulation. Vestigial precoxal ridge present.
Episteyrnum reduced to a small subtriangular plate ; epimeron produced posteriorly. Meta-
thoracic spivacle similar to mesothoracic one. Dorsospivaculay setae : 9-18 (average 15) f.s.
Postmetaspivacular setae : 13-19 (average 16) f.s. and 1-3 (average 1-6) h.s. Metasternal plate
weak and irregular, but more heavily sclerotized anteriorly. Anterior and posterior meta-
sternal setae : 41-66 (average 54) and 20-46 (average 33) f.s. respectively.

Wings hyaline ; short and comparatively broad : 970-1060 (average 1018) uw long and 445~—500
(average 483) u wide, the ratio width to length being 1 : 2-04—2-18 (average 2:11) ; alay lobe and
alay setae absent. Halteres absent.

124 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Legs long and slender, with the fore pair usually shortest and the hind pair longest ; the ratio
length of hind leg to body length is 1 : 1-54-1-69 (average 1-61). Length of segments (in py) :

Leg Coxa Trochanter Femur Tibia Tarsus Claw Total

I 57-65 70-78 220-247 323-376 124-131 24-27 827-928
(62) (76) (231) (347) (129) (25) (870)

II 68-78 84-95 194—209 346-380 125-137 25-28 849-927
(75) (89) (201) (363) (132) (27) (885)

TEE 80-87 QI-I101 201-220 305-418 133-137 25-27 903-989
(83) (98) (213) (394) (136) (26) (950)

Coxae : with 14-16 (average 15), 22-30 (average 26) and 30-41 (average 37) f.s. and 7-9
(average 8), 7-11 (average 9) and 12-17 (average 13) h.s. on the fore, middle and hind coxa
respectively ; fore coxa with 2—4 (average 3-2) capitate coxal bristles ; apical seta about $ as
long as trochanter. Tvochanterys 23-30 u wide ; with 6 oval sensilla, 10-17 f.s. and 6-8 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 1-69-2-31 (average 2-01) times as long as width of
trochanter. Femova of medium width (38-46 yu), ratio width to length of hind femur being
I : 4:6-5-3 (average 4-9) ; each with 31-44 f.s. and 12-23 h.s. Tzbia 19-25 wu wide, ratio width
to length of hind tibia being 1 : 16-18 (average 17) ; each with 92-120 setae, of which 19-33 are
h.s. and 71-88 f.s., the latter about 14 times longer than width of tibia ; apical spur about the
same size on all tibiae. Tavs: 19-22 u wide, hind tarsus 6-0-6-6 (average 6-4) times longer than
wide ; with 20-32 f.s. and 8-16 hs. ; tarsal digitules subequal, slightly 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 330-460 (average 399) u long and 300-360 (average 333) w wide.

Segments I-VII : tervgites represented by a small sclerite on each side on anterior margin of
segments II-III, and a weak transverse plate on VI-VII ; stevnites represented by a weak
transverse plate on segments II, II] and VII. Caudal extension of segment VII large, tapering,
weakly sclerotized. Dorsal setae : 5-14 (average 9) f.s.on I, 1-5 on Il and III, and up to 3 oneach
of segments IV—VII ; occasionally a single h.s. on I-III, and usually one on each side on each of
segments IV—VII. Pleural setae consisting of dorsopleural setae: 2-10 (average 4:4), I-5
(average 2:9), I-6 (average 3:2), o—-6 (average 3-2), 2-6 (average 4-1) and 1-4 (average 2:2) f.s.
on I-VI, and o~1 (average 0-3), I-2 (average 1-6), 1-2 (average 1-7), I-3 (average 2-1) and o-2
(average 1-6) h.s. on segments [I-VI respectively, and of ventvopleural setae : up to 5 f.s. on
each of segments III-VI, and usually one (range o-2) h.s. on IV-VI. Segment VII with
12-17 (average 14) f.s. and 3—5 (average 3:4) h.s. ; some of the posterior h.s., usually longer than
the rest. Ventral setae : 23-35 (average 29), 14-31 (average 23), 12-22 (average 16), 8-14
(average I1), 5-12 (average 9) and 6-11 (average 9) f.s. on segments II—-VII respectively ;
usually one h.s. on each side on segments III—VI and 4 on VII.

Segment VIII: with weak tergite and transverse sternite ; caudal extension forming a
prominent, weakly sclerotized, semi-circular lobe, with a large cicatrix posterodorsally, the latter
weakly sclerotized and reticulated in the middle ; glandular pouch with 2 long, pointed setae,
whose protruding part is 3-34 times as long as section within pouch. Small I Xth tergite present.
Ante-anal setae : 2-5 (average 3°8) f.s., o-4 (average 1-6) small h.s. and occasionally one long
h.s. Posterior margin with 3-7 (average 4-2) f.s. and 3-4 (average 3:2) h.s. on each side.

Genital segment. Penial sheath long, about ? total body length (ratio 1 : 3-5-3-8, average
3:6), 407-445 (average 426) uw long and 38-44 (average 40) uw wide ; lateral sclerotizations
narrowly joined anterior to anus ; length of basal vod about twice that of aedeagus, extending
anteriorly from base of the aedeagus for about 2 of the distance to apex of the basal membranous
area ; apex of sheath with a distinct, finger-like membranous extension. Area from base of
sheath to tip of aedeagus with 23-29 (average 25) small sensilla ; a cluster of small sensilla
occurring ventrally near apex of sheath. Aedeagus short (87-106, average 95 u long), penial
sheath and basisternum longer, ratios being I : 4-1-4:9 (average 4:5) and I : 1-53-1-78 (average
1-66) respectively.

OF THE FAMILY COCCIDAE 125

Material examined : 10 specimens, collected by J. Munting on Muilletia sp. in
Umkomaas, South Africa, on 7.11.62 ; no adult females present with the males ;
nearby females of Ceroplastes ?mimosae Hall occurred on Abaris sp. ; identification
by D. J. Williams.

This species differs from C. berliniae in having coxal bristles on the fore coxa,
and a shorter aedeagus (measured, for example, against the basal rod or the penial
sheath) and also in a number of less striking characters, e.g. in the presence of a
vestigial dorsal part of the midcranial ridge, and in the number of f.s. between and
behind the eyes, on antennal segment IV and ventrally on abdominal segment VII.

DISCUSSION

The present study of male Coccidae has revealed a number of interesting facts
concerning the taxonomy of this family, including its relationships with other mem-
bers of the superfamily Coccoidea. Unlike the other larger families of the Coccoidea,
such as the Diaspididae or even Pseudococcidae, our knowledge of the intra-family
relationship of various forms included in the family Coccidae is very limited. Only
very few workers have attempted to classify this large family on the basis of the
customary female characters. Steinweden (1929) grouped a small number of
genera around each of the three genera Coccus L., Towmeyella Ckll., and Exaerotopus
Newst., but left most of the genera studied by him as ‘‘ ungroupable " ; Sulc (1941)
formed the tribe Eriopeltini to include the three genera Eviopeltis, Scythia and
Mohelnia (the latter is now regarded as being synonomous with Scythia (Borchsenius,
1957)). Bodenheimer (1953) divided the Coccidae of Turkey into four subfamilies,
i.e. the Coccinae (including Coccus, Eulecanium, Sphaerolecanium, Saissetia, Pul-
vinaria and Paralecanopsis), Filippiinae (including Filippia and Euphilippia) and
the Ceroplastinae and Eriopeltinae which include Ceroplastes and Eriopeltis res-
pectively. The only relatively comprehensive classification, based on a com-
paratively large number of genera (37) is that of Borchsenius (1957), who divided
the family into three subfamilies, the Filippiinae, Coccinae and Ceroplastinae,
the Coccinae being further subdivided into two tribes, the Coccini and Pulvinariini.
This classification is mainly based on a few characters of the adult female and
particular emphasis is put on the way in which the body and eggs are covered.

The male material, 19 genera (23 species) used in the present investigation, in-
cluded 14 of the 37 genera on which Borchsenius based his classification, representing
all his subfamilies and tribes. On an examination of the characters of the males it
was immediately apparent that they exhibit entirely different relationships, which
do not conform with the division of the family suggested by Borchsenius. A fresh
approach was therefore necessary. The large number of characters available (listed
in Table I) made some quantitative evaluation of these relationships possible, but
detailed statistical analyses such as mentioned by Sneath and Sokal (1962) and
fully discussed very recently in their book on numerical taxonomy (Sokal and
Sneath, 1963) are beyond the scope of the present work. Therefore genera which
appeared to resemble each other, especially in sharing distinct features such as
prominent caudal extensions, an interocular ridge, a head with a pronounced

126 MORPHOLOGY AND TAXONOMY OF ADULT MALES

anterodorsal bulge, large numbers of setae, etc., were grouped together. The validity
of these groups were then tested by calculating the number of characters shared by
the group as a whole, and by pairs of genera from different groups. For the purpose
of these calculations all characters were treated as of equal weight and importance.
It was found that the family could be divided into four groups of genera, which can
conveniently be called the EULECANIUM group, the ERIOPELTIS group, the
INGLISIA group and the COCCUS group. In the following discussion the refer-
ences to suprageneric groups refer to these groups and three aspects will be discussed:

(a) the characters which are of taxonomic importance and the levels at which
they appear to be valid,

(b) the classification and interrelationships within the family, and

(c) the relationship of this family with other subdivisions of the Coccoidea.

It must be stressed here that only a small number of species were studied. All
the statements and conclusions are therefore tentative and some of them are bound
to be altered or even abandoned as more information becomes available, particularly
because the grouping suggested from a study of the male does not conform with the
classification based on the female.

Taxonomic Significance of the Characters

All the characters which appear to be of some taxonomic importance are listed in
Tables I-IV. The characters which at this stage of research were found useful for
separating the groups of genera, genera and species are discussed below and listed
in detail in Table I.

The size and general appearance show considerable variation within the family.
Broadly speaking, the size is characteristic of species or groups of species but the
actual size may vary considerably within one species. Thus specimens of P. corn
from different localities (Poland, Russia and Yugoslavia) showed large differences,
e.g. those from Yugoslavia were about ? the size of those from Poland, the Russian
ones being intermediate. Apart from local climatic conditions, this may be due to
the effect of the host plant, which has been shown to influence the size of the females
of this species (Habib, 1953 ; Kawecki, 1958a) ; Bustshik (1958) suggested that the
effect of the host plant caused size variation in males of some Diaspididae, whereas
Ghauri (1962), having found males of distinctly different sizes on the same host
plant, mentioned genetic polymorphism as a possible cause. The size is therefore
considered to be of rather limited significance. The general “ hairy ’’ appearance
of the body can be used to separate groups of genera.

The Head.

The shape of the head is characteristic of groups of genera, thus it is flat in the
ERIOPELTIS group (Text-figs. 25, 28) and elongated dorsoventrally in all the
other groups except Genus A (Text-fig. 21) where it is rounded. The peculiar
condition where the anterodorsal bulge is pronounced and the medioventral bulge
drawn far back, is characteristic of most of the COCCUS group (Text-figs. 32-40).
The conditions of the ventral part of the midcranial ridge separate groups of genera,

OF THE FAMILY COCCIDAE 127

being reduced to a greater or lesser degree in the ERIOPELTIS and INGLISIA
groups, but complete in the COCCUS and EULECANIUM groups ; the absence
of the lateral arms separates EF. ?festucae from all other species. The dorsal part of
the midcranial ridge is frequently absent or vestigial and in the latter case it can
usually only be seen in well stained preparations, thus it is not particularly useful,
but apparently operates on the specific level. The postoccipital ridge, which pro-
vided a series of characters in the Diaspididae (Ghauri, 1962), is absent in this
family. The absence of reticulation on the median crest is peculiar to two genera of
the EULECANIUM group. The degree of development of the postocular ridge
separates groups of genera, genera and species in so far as it (i) is generally weak
and tapering dorsally, but strong in the COCCUS group and the genus Eriopeltis,
(ii) usually forks below the ocellus, except in the ERJOPELTIS group and (iii) is
reduced posteromedially in some species but not in others. A broad interocular
ridge is characteristic of the ERJIOPELTIS group. The preoral ridge is absent in
the ERIOPELTIS group and two genera of the EULECANIUM group. The
size of the cranial apophysis and the shape of its apex show some variation which
can be used at the generic and specific level. The number of simple eyes is a very
distinct and useful character. In the ERIOPELTIS and COCCUS groups the
number is constant (4), but in the EULECANIUM group the number varies from
4-10, separating genera. Thus four closely related genera, Eulecanium, Nemole-
canium, Physokermes and Rhodococcus can easily be separated in having 10, 6, 4
and 8 eyes respectively. This character was suggested to be of generic importance
by Newstead (1903), and Sule (1908) when he defined the genus Eulecanium Ckll.
and his two new genera Sphaerolecanium and Palaeolecanium. The large size of the
lateral eyes is characteristic of the JNGLIJS/A group.

The setae of the head provide a number of important characters and their absence
or presence in various regions of the head can be used to separate groups of genera.
In the EULECANIUM and ERIOPELTIS groups there are no fleshy dorsal head
setae, no setae at all between and behind the ventral simple eyes and no genal
setae ; in the EULECANIUM group there are also no fleshy ventral head setae.
In the JNGLISIA and COCCUS groups there are present fleshy dorsal head setae as
well as fleshy ventral head setae, setae between and behind the ventral eyes, and
setae on the genae. The number of setae varies individually, but the differences
in the ranges of individual variation can be used to separate some species and
genera.

The antennae provide a number of characters which can be used at the generic
and specific levels. These are (i) the length of the antenna in relation to the length
of the body, the posterior leg and the penial sheath, (ii) the width of the 2nd segment
relative to that of the rst, (iii) the length of the 3rd segment in relation to its width,
(iv) the relative lengths of the 3rd and roth segments and (v) the shape of the
terminal segment. In addition the relative length of the fleshy setae and the antennal
bristles, and also the number of subapical setae can be used to separate
genera.

128 MORPHOLOGY AND TAXONOMY OF ADULT MALES

The Thorax.

Prothorax. The absence or presence of certain groups of setae and pores can be
used to separate genera ; a reticulated prosternum is characteristic of Coccus
hesperidum and a spinose prosternum of the JNGLISIA group. The condition of
the median ridge of the prosternum varies too much to be of practical use as it may
be complete or any section of it reduced within the same species.

Mesothorax. This provides a number of important characters. The shape of
the mesoprephragma, the condition of the reticulation on the prescutwm and the size
of the membranous area of the scutwm can be used as supplementary characters to
separate genera and species. A tubular scwtellum is characteristic of the ERIJO-
PELTIS and COCCUS groups and the two closely related genera Ctenochiton and
Filippia. A basalare joins the pleural wing process to the mesepisternum in all
the groups except the COCCUS group, where this structure is vestigial or absent.
The condition of the median ridge of the basisternum separates groups of genera,
genera and species. As is the case with the head, the setae of the mesothorax
(and metathorax) provide a number of very useful characters. The number of
fleshy and hair-like scutal setae separates certain genera ; the presence of fleshy
postmesospiracular setae separates the COCCUS and INGLISIA groups from the
EULECANIUM and ERIOPELTIS groups ; the setae on the postalare and
basisternum can be used to distinguish genera.

Metathorax. The condition of the metapleural ridge and suspensorial sclerites is
correlated with the absence or presence of the halteres (g.v.). When the halteres
are absent, the metapleuron is reduced and the suspensorial sclerites absent. The
presence of antemetaspiracular and dorsospiracular setae differentiates the
INGLISIA and COCCUS groups, while the variation in the number and type of
postmetaspiracular, anterior metasternal and posterior metasternal setae can be
used to differentiate groups and genera.

The fore wings vary from being long and narrow in the ERJOPELTIS group to
short and broad in the COCCUS group ; inthe EULECANIUM group this variation .
in shape can be used to separate genera. The absence of halteres is characteristic of
the ERIOPELTIS, INGLISIA and COCCUS groups and the genera Phyllostroma
and Sphaerolecanium. Sulc (1908) used this characteristic to define genera. The
differences in the number of alar and haltere setae can be used to differentiate genera.

The characters provided by the Jegs seem to operate on all taxonomic levels.
Some characters are constant within one or two of the groups, but differentiate
genera and species in others, e.g. (i) the length of the hind leg and the length of the
body is subequal in the ERIOPELTIS and INGLISIA groups, but differences in
the ratio between these two measurements separate genera in the FULECANIUM
group, and the two species of Pulvinaria in the COCCUS group. (ii) The coxal
bristles are absent in the ERIOPELTIS group and some of the genera of the
EULECANIUM group, but they are present in other genera of the FULECANIUM
group, in the JNGLISIA group as well as in the COCCUS group (except Ceroplastes

OF THE FAMILY COCCIDAE 129

berliniae). (iii) the total number of setae on the fore tibia and the preponderance
of fleshy setae on the hind tibia separate the COCCUS and INGLISIA groups from
the ERIOPELTIS group, but only distinguish genera within the EFULECANIUM
group. Other characters are mainly significant at the generic and specific levels,
e.g. (i) the relative lengths of the apical setae on the coxa and trochanter, (ii) the
shape of the femur, (ii) the size of the apical spur of the fore tibia in relation to
those on the other tibiae, (iv) the length of the fleshy setae in relation to the width
of the tibia, (v) the number of fleshy setae relative to the number of hair-like setae
on the hind tarsus, (vi) the length of the hind tarsus in relation to its width and
(vii) the relative length of the hind claw.

The Abdomen.

The abdomen and genitalia also provide a series of important characters. The
number of tergites and sternites usually vary within the group. The absence or
degree of development of the plewral sclerotization, however, remains constant
within the groups. A prominent, tapering caudal extension on segment VII is char-
acteristic of the COCCUS group. The shape of the caudal extension of segment
VIII and the position and size of the cicatrix differentiate the genera of the COCCUS
group. The number of dorsal abdominal setae on certain segments varies somewhat
individually, but can be used as a supplementary character to separate genera and
species. The length of the setae of the glandular pouch shows considerable differ-
ences between species and genera ; in Luzulaspis luzulae the pouch itself is absent.
The presence of fleshy setae lateral to the glandular pouch is characteristic of the
genus Ceroplastes. Fleshy pleural setae occur in the ERIOPELTIS, INGLISIA
and COCCUS groups and the genus Sphaerolecanium, but only in the COCCUS
group are they present on segments I-III. Fleshy ventral setae are characteristic
of the same genera as the pleural setae, but Sphaerolecanium differs from the others
in that the fleshy setae only occur on segments II and sometimes III ; a large
number of fleshy setae on segment VIII is typical of the JNGLISIA group. The
number and arrangement of the hair-like ventral setae can be used as a supporting
character to separate genera and species.

The relative lengths of the various structures of the genital segment show consider-
able differences, usually on the generic, but also on the specific level. The following
ratios were found useful : (i) length of penial sheath to body length, (ii) length of
basal rod to length of aedeagus, (ili) length of aedeagus to length of penial sheath,
(iv) length of aedeagus to length of basisternum. A finger-like membranous exten-
sion of the apex of the penial sheath is characteristic of the genus Ceroplastes.

MALES

ADULT

MORPHOLOGY AND TAXONOMY OF

130

Ceroplastes sp.

C. berliniae

pomeranicum
acericola
P. ? betulae

12),
P.

dno13 sno009

Genus B

C. hesperidum

L. luzulae

E. ?festucae
Eriopeltis sp.
S. prunastri
Ctenochiton sp.
F. viburni

P. myrtilli

P. bituberculatu
R. spiraeae

Genus A

sijadoisg dnoi3 uimiueserng

P. piceae

N. abietis

E, tiliae

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I dTavL

131

FAMILY COCCIDAE

TH

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4

OF THE FAMILY COCCIDAIE

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ADULT MALES

AND TAXONOMY OF

MORPHOLOGY

134

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135

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MALES

ADULT

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AND 7

MORPHOLOGY

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137

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*yuasqe aejas AysaTq “gd
*yuasaid avjas Aysalgy “Wy : aevjeS TeInoyd Teulwopqy

OF THE FAMILY COCCIDAE

‘quesqy ‘qd ‘juasaid a10ow 10 auQ “YW
; u013e1 Teut-ajue ul Sadod re[noI1D

‘auo Ajaaea ‘yuasqe avjas Aysala “a
L-Z eejas AyseTy “y +: yonod 1e[npueys 0} Tes19zyeT

*ae}aS JyI[-11ey Buoy ynoyyM Ayyensy “gq
*aejas ayl[-aley Buoy Z yy ATTensyH “VW

“Juasqe Ayyensn oejes
Aysetq *q ‘“yuaserd Aqyensn aejes Aysaty “V

: avjas [eue-ajuy

ADULT MALES

OF

MORPHOLOGY AND TAXONOMY

Nn

13

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sewijg uey} a1oUI Yyyeoys [elued ‘y10yS ‘y > yyeoys

Teluad Jo yySua] 0} uo1jzeTaa ul SnBeaper jo yySsueT

"varie snoueiquioul [eseq Sulyovas
JON ‘@ ‘vere snourIquiaw [eseq Suiyorer
jsowye 10 Suyoevar ATIOl1ajuy “Wy : pod [eseg

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Soul} a10W! 10 2g ATTensn yySuey Apoq ‘y104g "Y
: yydua] Apoq 0} uolyeyaa ul Yeas Te1luad jo yySuaT

*popue}xe ATSnouvaiquioul jON ‘q
*popua}xe ATsnouriquieayw ‘y : yyeays Teluad jo xady

OF THE FAMILY COCCIDAE 139

Classification and Interrelationships of the Coccidae based on the Males
Groups of Genera.

A. The characters, which were found from an examination of Table 1 to separate
groups of genera, are listed in Table 2. The table contains 34 characters and
shows their distribution among the material studied. It will be noticed that a few
characters are included of which the alternate conditions occur within the same
group (thus only separating genera), but which are of significance in separating
other groups. The characters which are exclusive to any particular group are
indicated with an asterisk (*) and their total number is given at the end of the
table. The number of these characters will be reduced as more information about
other members of the family becomes available and it must be stressed here that
because of the limited number of species studied the diagnosis and discussion of
relationships are very tentative.

To investigate the relationship between groups the method of analysis used by
Ghauri (1962) was followed, and the following were calculated :

(a) the number of characters shared by any two groups,

(b) the number of characters exclusive to these two groups, and

(c) the number of characters by which the two groups differ from each other.

The results are given in Table 2 A. The characterization of each group is implicit
in Table 2 and given fully in the keys which follow later.

TABLE 2.
List OF CHARACTERS WHICH SEPARATE MAJOR GROUPS OF GENERA

Characters Eule- Erio- Ingli- Coccus
canium peltis sia
HEAD:
1. In lateral view: A. Flat. B. Dorso-
ventrally elongated or rounded. B A*® B B

2. Length of antenna in relation to
length of posterior leg: A. 1-05 or more
times longer. 8B. Subequal or shorter. AB A A B

3. Interocular ridge: A. Present.
B. Absent. AB A B B

4. Postocular ridge: a. Forking below

ocellus. B. Not so. A B* A A
5. Midcranial ridge, ventrally:
A. Reduced. gs. Not so. B AB A B

6. Number of simple eyes: A. 4.
B. More than 4. AB A B A

140 MORPHOLOGY AND TAXONOMY OF ADULT MALES

TABLE 2. LisT OF CHARACTERS WHICH SEPARATE MAJOR GROUPS OF GENERA
Characters Eule- Erio- Ingli- Coccus
canium peltis sia

HEAD—contd.
7. Lateral simple eyes (when present):
A. About as large as dorsal ones.
B. Distinctly smaller. —B - Ae -

8. Pre-oral ridge: a. Absent. 3B. Present. AB A B B

g. Fleshy dorsal head setae: A. Present.
B. Absent. B B A A

Io. Dorsal ocular setae: a. Present on
at least one side. B. Absent. AB B A A

11. Fleshy ventral head setae:
A. Present. B. Absent. B* A A A

12. Setae between and behind ventral

eyes. A. Present. B. Absent. B B A A

13. Genal setae: A. Present. 3B. Absent. B B A A
THORAX:

14. Prosternum: A. Spinose. B. Not so. B B AX B

15. Scutellum: a. Tubular. Bs. Not so. AB A B A

16. Scutellum: A. 3-4 times as wide as
long. 8B. I-3 times as wide as long. B AB B A

17. Fleshy scutal setae: a. Present.
B. Absent. B B A AB

18. Basalare: a. Joining pleural wing
process to mesepisternum. B. Vestigial A A A B*

19. Median ridge of basisternum:
A. Complete. 3B. Reduced or absent. A AB B A

20. Fleshy postmesospiracular setae:
A. Present. 3B. Absent. B B A A

21. Antemetaspiracular setae:
A. Present. 3B. Absent. B B A A

22. Fleshy dorsospiracular setae:
A. Present. B. Absent. B B A A

23. Fleshy anterior metasternal setae:
A. More than 36. B. 0-24. B B B A*®

OF THE FAMILY COCCIDAE 141

TABLE 2 LIsT OF CHARACTERS WHICH SEPARATE MAJOR GROUPS OF GENERA

Characters Eule- Erio- Ingli- Coccus
canium peltis sia

THORAX—contd.
24. Posterior metasternal setae:
A. More than 4. B. Less than 4. B* A A A

25. Wings: A. Less than 2} times longer

than wide. B. 2}-2} times longer than

wide. c. More than 2} times longer than

wide. AB ce B A

26. Total number of setae on front tibia:
A. More than 60. B. Less than 60. AB B A A

27. Hind tibia: a. With more fleshy than
hair-like setae. B. With more hair-like
setae. AB B A A

ABDOMEN:
28. Pleural sclerotization: a. Present on
segm. VII. B. Present on segm. IV—VII.
c. Absent. c* A B* A

29. Caudal extensions of segm. VII:
A. Very prominent, tapering. B. Small,
broadly rounded or somewhat pointed. B B B I Nes

30. Cicatrix on caudal extension of
segm. VIII. a. Absent. 8B. Present. A A A B*

31. Fleshy pleural setae on segm. I-III:
A. Present. sB. Absent. B B B A*

32. Fleshy ventral setae: a. Absent
posteriorly beyond segm. III. B. Present
posteriorly beyond segm. IIT. As B B B

33. Fleshy setae on 8th sternite: a. More
than 6. 3B. Not more than 3. B B A* B

34. Ventral hair-like setae on segm. III,
usually: a. Two setae medially. B. No
setae medially. AB B A B

Total number of exclusive characters 4 3 4 5

142 MORPHOLOGY AND TAXONOMY OF ADULT MALES
TABLE 2 A

Number of Characters

Pairs of Groups (a) (b) (c)
shared of which exclusive differentiating
Eulecanium-Eriopeltis 14 6 6
Eulecanium-Inglisia 8 I 15
Eulecanium-Coccus 6 I 16
Eriopeltis-Inglisia 9 I 21
Eriopeltis-Coccus 9 2, 17
Inglisia-Coccus 16 6 16

1. The COCCUS group. To this group have been assigned five genera which
appear to be very closely related. They are Pulvinaria, Genus B, Coccus, Parthenole-
canium and Ceroplastes. Pairs of genera share from 75 to go characters and the
group as a whole has about 60 characters in common. The exclusive characters of
this group are (i) the basalare vestigial or absent, (ii) the caudal extension of
abdominal segment VII very prominent and tapering, (iii) the presence of a mem-
branous or weakly sclerotized cicatrix on the caudal extension of abdominal segment
VII, (iv) the presence of more than 36 fleshy anterior metasternal setae, and (v) the
presence of fleshy pleural setae on the first three abdominal segments. In all the
genera (except Ceroplastes) the shape of the head is rather peculiar in having a
pronounced anterodorsal bulge with the medioventral bulge sharply drawn back.
Except for this and some other small differences, the males of genus Ceroplastes are
very similar to those of the other members of this group. This is rather interesting
because the females of the genus Ceroplastes are considered to be quite distinct and
different from the other Coccidae.

The COCCUS group is probably the most specialized of the Coccidae, as a com-
paratively large number (8) of specialized characters are found in it. They are :
(i) the presence of only four simple eyes, (ii) the tubular scutellum, (iii) the reduced
metathorax, with the episternum small, the pleural ridge short and the halteres
lacking, (iv) a basalare which is vestigial and incorporated into the pleural wing
process, (v) considerable desclerotization of the abdomen, (vi) the presence of
prominent caudal extensions on abdominal segment VII (although reminiscent of
the “ fleshy tassels ’’ of the Monophlebidae), (vii) the presence on abdominal seg-
ment VIII of caudal extensions of various shapes which bear a membranous or
weakly sclerotized cicatrix, and (vii) the presence of pleural sclerotization on
abdominal segments VII-VIII. (This does not occur in the more primitive
Coccoidea, nor in what is regarded as the more primitive members of this family.)

From a study of limited material of Akermes andersoni Newst., the drawings and
descriptions of Chloropulvinaria aurantu (Ckll.) by Borchsenius (1957) and Neo-
pulvinania imeretina Hadz. by Hadzibejli (1955), and the drawing of the posterior
part of the abdomen of Sazssetia nigra (Nietner) by Green (1904-1909) it seems
certain that the genera Akermes, Chloropulvinaria, Neopulvinaria and Saissetia
should also be included in this group.

OF THE FAMILY COCCIDAE 143

2. The INGLISIA group comprises only the genus /nglista, which shows a
number of distinctive characters justifying its separation from all other groups. A
brief study of a species of Ceroplastes (C. chiton Green) showed that this species also
belongs to the JNGLISIA group. The exclusive characters of the group are the
following : (i) lateral eyes about as large as the dorsal and ventral ones, (ii) pro-
sternum spinose, (iii) pleural sclerotization extending from the IVth—VIIth ab-
dominal segments, (iv) more than 6 fleshy setae on abdominal sternite VIII.

The group features a number of primitive and specialized characters. The
specialized ones are : (i) the considerably reduced midcranial ridge, (ii) the small
spines present on the prosternum and on the front coxae, (ili) the reduced median
ridge of the basisternum, (iv) the reduced metathorax which lacks halteres, suspen-
sorial sclerites and a complete pleural ridge, and (v) the extensive pleural sclerotiza-
tion which, as discussed earlier, is probably of secondary development and therefore
a specialized feature. The primitive features include: (i) eight simple eyes,
which is probably a primitive condition as it more closely resembles the compound
eye of the margaroid ancestors, (ii) sternal plates on all the abdominal segments,
(iii) a scutellum which is not tubular, (iv) a basalare connecting the pleural wing
process with the episternum, and (v) the absence of prominent caudal extensions
on abdominal segment VII, or caudal extensions with a cicatrix on the VIIIth.

3. The ERIOPELTIS group: This group consists of the two closely related
genera Eriopeltis and Luzulaspis, which share about go characters between them.
Three characters are exclusive to this group: (i) the head comparatively flat
dorsoventrally, (ii) the postocular ridge not forking below ocellus, and finally,
(iii) the long and narrow wings which are more than 2? times longer than wide.
Another characteristic of this group is the presence of an interocular ridge, but this
is sometimes shared by Genus A of the EULECANIUM group.

The ERIOPELTIS group exhibits both primitive and specialized features. The
two genera share the following specialized characters : (1) an interocular ridge,
connecting the pre- and postocular ridges, (ii) a more or less reduced midcranial
ridge, (ili) four simple eyes, (iv) the absence of the preoral ridge, (v) a tubular
scutellum, (vi) a reduced metathorax with a small episternal plate, short pleural
ridge and no halteres or suspensorial sclerites, and (vil) the presence of pleural
sclerotization on abdominal segment VII. In each of the two genera an additional
specialized feature is found : in Luzulaspis the cluster of pores (and the pouch) on
abdominal segment VIII has secondarily been lost (the pores are present in the more
primitive margaroid Coccoidea (Morrison, 1928 ; Theron, 1958, 1962), the Pseudo-
coccidae (Giliomee, 1961) and all the other species studied here) ; in Eriopeltis the
median ridge of the basisternum is reduced.

The prevailing primitive features in this group are : (i) the presence of distinct
anterior tentorial pits, (ii) the presence of a basalare connecting the episternum and
the pleural wing process, (iii) the absence of prominent caudal extensions on
abdominal segment VII or caudal extensions with a cicatrix on the VIIIth. In
addition, in Eriopeltis the ventromedial part of the epicranium is well sclerotized,
and tergites and sternites are present on all the abdominal segments.

144 MORPHOLOGY AND TAXONOMY OF ADULT MALES

The ERIOPELTIS group can be regarded as being more specialized than the
next group, the EULECANIUM group. This conclusion is at variance with the
views of Borchsenius (1957), who held that they were the most primitive Coccidae.

4. The EULECANIUM group: In this group the rest of the genera studied
have been included, i.e. Eulecanium, Nemolecanium, Physokermes, Rhodococcus,
Parthenolecanium, Phyllostroma, Filippia, Ctenochiton, Evicerus, Genus A and
Sphaerolecanium. There is little doubt that this group is a temporary assemblage
of genera, related to each other to a greater degree than to the three groups already
discussed. The relationships between these genera are very complex and until
more representative material has been studied in detail it seems best to treat them
together, although further subdivision appears to be inevitable. The heterogeneity
of this group of genera is reflected in the comparatively small number of characters
(about 25) that are common to all of them. Four of these are exclusive to this
group, namely (i) pleural sclerotization on the abdomen absent, (ii) fleshy ventral
head setae absent, (i11) posterior metasternal setae less than 4, and (iv) fleshy setae
absent posteriorly beyond abdominal segment III.

All the genera included in the EULECANIUM group appear to be more primitive
than those of the other groups, especially the genera Eulecanium, Nemolecamum,
Physokermes, Rhodococcus, Palaeolecanium, Ericerus and Genus A. They have in
common the following characters, which can be regarded as being primitive : (i) a
scutellum which is not tubular, (11) a basalare connecting the pleural wing process
with the episternum, (ili) a comparatively less specialized metathorax with the
episternum and metapleural ridge well developed, the latter with a vestigial pleural
wing process which supports the haltere, which is in turn connected to a small
suspensorial sclerite, (iv) the absence of prominent caudal extensions on abdominal
segment VII, (v) the absence of prominent caudal extensions with a cicatrix on
abdominal segment VIII. Furthermore, three of these genera namely Eulecanium,
Nemolecanium and Physokermes share a sixth primitive character in possessing small
tergites and sternites on all abdominal segments (this is also shared by Phyllostroma).
The other four genera share some of the above-mentioned characters, though not all
of them. All the genera except Physokermes, Palaeolecanium and Sphaerolecanium
share an additional feature which is probably primitive, i.e. having more than 4
simple eyes.

Of the genera in this group, Sphaerolecanium appears to be the most specialized.
In this genus the scutellum is tubular (though the ventral foramen is large), the
halteres are absent and the metathorax reduced, the caudal extension of the VIIIth
abdominal segment forms a prominent cylindrical lobe, and only 4 eyes are present.
This genus might be regarded as linking the EULECANIUM group with the
ERIOPELTIS or COCCUS groups.

If relationships are estimated by calculating the total number of characters
shared by pairs of genera it appears that the genera Eulecanium, Nemolecamium,
Physokermes, Rhodococcus and Palaeolecanium are very closely related, with pairs of
genera sharing between 75 and 85 characters. Phyllostroma also comes close to
this group, especially to Palaeolecanium. The two genera Filippia and Ctenochiton,

OF THE FAMILY COCCIDAE 145

while not very close to these, share between themselves a large number of characters
(almost go). The three remaining genera, i.e. Evicerus, Genus A and Sphaerole-
canium are not very close to each other or to any of the other above-mentioned
subgroups (although Evicerws comes close to Rhodococcus). It is therefore clear
that the EULECANIUM group, in which 11 out of 19 genera studied have been
included, is very heterogeneous and as more material becomes available it will
undoubtedly be further subdivided.

B. From Tables 2 and 2A it can be seen that the ERIOPELTJS and EULE-
CANIUM groups agree in a large number of characters (14), of which 6 are common
to them alone (i.e. exclusive). Conversely in the COCCUS and JNGLISIA groups
the alternative condition of these 6 characters obtains and they share 15 characters
amongst themselves. Any other way of coupling pairs of groups results in a
comparatively small number of characters being shared or exclusive. It would
appear therefore that there are two major groups, ERIOPELTIS and EULE-
CANIUM on the one hand and JNGLISIA and COCCUS on the other, which can
be separated mainly by the absence or presence of setae in various regions of the
body, as follows :

INGLISIA and COCCUS groups ERIOPELTIS and EULECANIUM
groups

1. Fleshy dorsal head setae present. Fleshy dorsal head setae absent.

2. Setae between and behind ventral Setae between and behind ventral

eyes present. eyes absent.

3. Genal setae present. Genal setae absent.

4. Fleshy postmesospiracular setae Fleshy postmesospiracular setae

present. absent.

5. Antemetaspiracular setae present. Antemetaspiracular setae absent.

6. Fleshy dorsospiracular setae present. Fleshy dorsospiracular setae absent.

The fact that these two groups can only be separated by differences in the
chaetotaxy indicates that this family consists of a morphologically rather homo-
geneous group of species. This is particularly true in comparison with the
Diaspididae where Ghauri (1962) found sharp and very distinct morphological
differences between major groups.

At present no rank is suggested for these groups of genera indicated by male
characters, but they may be considered to be equivalent to each other and repre-
senting subfamilies, while the further subdivisions in the EULECANIUM group,
and possibly the COCCUS group could represent tribes.

The diagnostic characters of each group can be found in the detailed key which
follows later.

C. The classification proposed above unfortunately differs considerably from
the existing classification suggested by Borchsenius (1957) and a few examples are
discussed as an illustration of these differences.

146 MORPHOLOGY AND TAXONOMY OF ADULT MALES

It will be remembered that Borchsenius based his classification on a few characters
of the adult female, with considerable emphasis on the way in which the body and
the eggs are protected. In his subfamily Filippiuinae, Borchsenius included the
three genera Luzulaspis, Eniopeltis and Filippia. From the male characters it is
clear that whereas Luzulaspis and Eriopeltis are very closely related (sharing about
go characters) they differ from Filippia in a number of striking features (and share
only about 55 characters), e.g. the shape of the head, the number of eyes, the
presence or absence of the interocular and preoral ridges, the shape of the wing, and
in the presence or absence of fleshy ventral head setae and fleshy abdominal setae.
On the other hand, Filippia shares up to 80 characters with some of the other
genera of the EULECANIUM group, e.g. Ctenochiton and Palaeolecanium. It
is therefore clear that Filippia and Eniopeltis are not closely related, a conclusion
which supports the views of Bodenheimer (1953), who put them into different
subfamilies.

In his tribe Pulvinariini Borchsenius included, among others, the two genera
Pulvinaria and Phyllostroma. The males of these two genera, however, are widely
different and have only a comparatively small number of characters (about 50) in
common. Some of the more striking differences are found in the number of eyes,
the condition of the scutellum and basalare, the development of the metathorax,
the condition of the caudal extensions of abdominal segments VII-VIII, and in
the chaetotaxy of the head and abdomen. Phyllostroma shares about 80 characters
with some members of the EULECANIUM group (e.g. P. bituberculatum) and
Pulvinaria about go characters with the genera of the COCCUS group.

On the other hand males of some genera which were widely separated by Borch-
senius indicate close relationships. Thus Borchsenius considered the genera
Pulvinaria and Coccus to represent two tribes of one subfamily and Ceroplastes a
different subfamily altogether. As discussed earlier the males of these three genera
are very similar and belong to the same group. The close relationship between
Pulvinaria and Coccus has also been indicated by Steinweden (1959) and Boden-
heimer (1953), who grouped them together into the same taxon. Both workers
considered Ceroplastes to be distinctly different from Coccus and Pulvinaria.

Further detailed studies are needed on both males and females to show if these
differences in classification of the two sexes illustrate the dependance of the
grouping on the stage (or selection of characters) as found by Morrison (1928) in
his studies of various stages of the Margarodidae, or reflect the inadequate know-
ledge of the females of the family, and finally which, if any, of the two classifications
reflects the true relationships.

Genera.

In Table 3 the characters are tabulated which were found to be useful in separating
genera in at least one of the three groups for which more than one genus were
available. These characters are marked “‘ G’”’ in the columns where they operate ;
where they only operate at the specific level in any of the remaining groups they are
marked ‘“‘S’’, while the symbols “GS” are used where the characters can ap-

OF THE FAMILY COCCIDAE 147

parently be used to separate both genera and species within the same group ; a
dash (—) signifies that the character does not operate in that particular group.
The table contains about go characters. This very high number is due to the fact
that in most cases only one species per genus was available and many characters
may eventually be shown to be of specific value only.

TABLE 3
List OF CHARACTERS WHICH MAINLY SEPARATE GENERA
Characters Eulec- Erio- Coccus
anium peltis
Heap:
1. Shape of the head. G - G
2. Absence or presence of polygonal reticulation
on median crest. G -
3. Presence of interocular ridge. G -
4. Nature of postocular ridge, dorsally. G G
5. Nature of postocular ridge, posteromedially. G ~ s
6. Midcranial ridge dorsally represented or not. G - GS
7. Nature of sclerotization and reticulation
around ventral part of midcranial ridge. G G -
8. Nature of preocular ridge. G - -
9. Number of simple eyes. G - -
10. Distance between dorsal eyes in relation to
diameter of cornea. G G G
11. Distance between ventral eyes in relation to
diameter of cornea. G ~ G
12. Presence of preoral ridge. G - ~

13. Presence of ventral sclerite near mouth opening. G - -

14. Length of cranial apophysis. G G G
15. Nature of apex of cranial apophysis. G GS =
16. Presence of polygonal reticulation on genae. G — Ss

17. Presence of a pair of long, median, hair-like,
ventral head setae. G G -

18. Location of ventral head setae. G - -

148

MORPHOLOGY AND TAXONOMY OF ADULT MALES

TABLE 3. LisT OF CHARACTERS WHICH MAINLY SEPARATE GENERA

Characters Eulec-
anium
ANTENNA:
19. Length in relation to body length. G
20. Length in relation to length of posterior leg. G
21. Length in relation to length of penial sheath. G
22. Length of 3rd segment in relation to its width. G
23. Relative widths of rst and 2nd segments. G
24. Length of fleshy setae on 3rd segment. G
25. Presence of hair-like setae on 3rd segment. G
26. Number of capitate subapical setae on roth
segment. G
27. 1oth segment distally constricted or not. ~
28. Length of two smallest antennal bristles in
relation to length of fleshy setae. G
THORAX:
29. Number of circular pores behind pronotal
sclerite. G
30. Prosternum reticulated or not. -
31. Number of medial pronotal setae. G
32. Presence of post-tergital setae. -
33. Presence of fleshy prosternal setae. G
34. Nature of emargination of mesoprephragma. G
35. Nature of reticulation on prescutum. G
36. Length of membranous area of scutum in
relation to its width. G
37. Scutellum tubular or not. G
38. Width of scutellum in relation to its length. G
39. Length of-basisternum in relation to length
of membranous area of scutum. G

Erio-
peltis

Coccus

GS

G

GS

OF THE FAMILY COCCIDAE

TABLE 3. List oF CHARACTERS WHICH MAINLY SEPARATE GENERA

Characters

THORAX—contd.

40.
41.
42.
43.
44-
45-
46.
47:
48.
49.
50.

51.

54.

55-

Presence of fleshy scutal setre.

Number of hair-like scutal setae.

Presence of postalary setae.

Number of fleshy postmesospiracular setae.
Presence and type of basisternal setae.
Presence of suspensorial sclerite.

Nature of metapleural ridge.

Number of fleshy anterior metasternal setae.
Length of wing in relation to its width.
Presence of alar setae.

Presence of halteres.

Number of haltere setae.

Length of hind leg in relation to body length.
Presence on coxal bristles on front coxa.

Total number of setae on front tibia.

Length of apical seta on front trochanter in

relation to width of trochanter.

56.

57-

Shape of hind femur (length/width).

Relative number of fleshy and hair-like setae

on hind tibia.

58.

59.

Relative length of fleshy setae on tibiae.

Relative numbers of fleshy and hair-like

setae on hind tarsus.

60.

61.

Length of hind tarsus in relation to its width.

Relative length of hind claw.

Eulec-
anium

G

G

Erio-
peltis

Coccus

G

GS

149

MORPHOLOGY AND TAXONOMY OF ADULT MALES

TABLE 3. LIST OF CHARACTERS WHICH MAINLY SEPARATE GENERA

Characters Eulec- Erio- Coccus
anium peltis
ABDOMEN:
62. Number of tergites. G G -
63. Nature of tergite between segm. I and II. G - G
64. Number of sternites. G - G

65. Absence or presence of glandular pouch on

segm. VIII. - G -
66. Shape of caudal extension of segm. VIII. G _ G
67. Location of cicatrix on segm. VIII. - - G
68. Length of external in relation to “‘internal’”’

part of setae of glandular pouch. G G G
69. Number of fleshy dorsal setae. - - G
70. Number of hair-like dorsal setae on segm. I. G - G
71. Number of hair-like dorsal setae on segm. IIT. G - G
72. Presence of fleshy ante-anal setae. G G G
73. Presence of 2 long hair-like ante-anal setae. G - G
74. Number of fleshy setae around glandular pouch. — - G
75. Presence of circular pores in ante-anal region. G G G
76. Presence of fleshy pleural setae. G - =-
77. Presence of fleshy ventral setae. G - -

78. Number and arrangement of hair-like setae
ventrally on segm. II. G - GS

79. Number of hair-like ventral setae on segs. VI
and VII. G - G

80. Lateral sclerotization of penial sheath fused
anterior to anus or not. G - G

81. Presence and size of membranous extension
of penial sheath. - - G

82. Length of penial sheath in relation to body
length. G G G

OF THE FAMILY COCCIDAE 151
TABLE 3. List OF CHARACTERS WHICH MAINLY SEPARATE GENERA

Characters Eulec- Erio- Coccus
anium peltis

ABDOMAN—cond. :
83. Length of basal rod in relation to length of
aedeagus. G G GS

84. Basal rod reaching basal membranous area
or not. G ~ ~

85. Length of aedeagus in relation to length of
penial sheath. G - -

86. Length of aedeagus in relation to length of
basisternum. G - Ss

COCCUS group: All the genera of this group are very closely related but each
has a characteristically shaped caudal extension on abdominal segment VIII.
Ceroplastes also differs from the others in the shape of the head.

INGLISIA group: Only one genus was studied and generic differences can
therefore not be discussed.

ERIOPELTIS group: The two genera Eriopeltis and Luzulaspis are very
closely related but Luzulaspis is distinct in lacking the glandular pouch and pores.

EULECANIUM group. The genera Eulecanium, Nemolecanium, Physokermes
and Rhodococcus are very closely related, but can easily be separated by the number
of eyes. Palaeolecanium differs from these genera in possessing a comparatively
short penial sheath. Phyllostroma is also rather closely related to these genera but
is characterized in lacking halteres and in having the metapleuron reduced. The
two genera Ctenochiton and Filippia form a distinct subgroup that can be separated
from the aforementioned in having a tubular scutellum and from each other by the
number of eyes and the development of the basal rod of the aedeagus. The three
remaining genera of the group are all very distinct. Evicerus is characterized by
the long, narrow head and the extremely long fleshy setae on the appendages,
Genus A by the exceptionally short and thick fleshy setae on the appendages and
the possession of a ventral sclerite, whereas Sphaerolecanium is the only genus in
this group with a prominent caudal extension on abdominal segment VIII, a basal
rod longer than the aedeagus and fleshy prosternal, anterior metasternal and
abdominal setae.

Species.

In Table 4 the characters are listed which separate species in the four genera in
which more than one species were available ; a “ x ”’ sign indicates the genus in
which the character operates. From the table it can be seen that differences
between species are not only indicated by differences in the chaetotaxy and

152 MORPHOLOGY AND TAXONOMY OF ADULT MALES

reticulation in various regions of the body, but also by differences in the develop-
ment of certain ridges such as the midcranial ridge, the postoccipital ridge and the
median ridge of the mesobasisternum.

TABLE 4
List OF CHARACTERS WHICH SEPARATE SPECIES
Characters Erio- Pulvin- Partheno- Cero-
peltis aria lecanium _ plastes

1. Degree of sclerotization of postero-
median part of postocular ridge. - -

x
|

2. Presence of lateral arms of mid-

cranial ridge. x Zz = zs
3. Shape of apex of cranial apophysis. x = = =
4. Presence of reticulation on gena. - x = 2
5. Presence of reticulation on pedicel. ~ K a =

6. No. of fleshy setae between and
behind ventral eyes. - = = x

7. No. of fleshy setae on 4th antennal
segment. - — - x

8. Size of two shortest antennal bristles. — x — -

g. Presence of reticulation on the
post-tergital region. x = = a

to. Nature of emargination of
mesoprephragma. x = ~ _

It. Presence of reticulation on
mesoprescutum. = x = 1

12. Presence of reticulation on anterior
arms of mesoscutum. x = a s

13. Presence of reticulation on
membranous area of scutum. Xx = = =

14. Size of ventral foramen of scutellum. — = x =

15. Development of median ridge of
mesosternum.

x
|
|
|

16. Length of hind leg in relation to
body length. = x = =

OF THE FAMILY COCCIDAE 153

TABLE 4. List OF CHARACTERS WHICH SEPARATE SPECIES

Characters Erio- Pulvin- Partheno- Cero-
peltis aria lecanium __ plastes

17. Length of hind tarsus in relation to
its width. se x = -

18. Presence of median pronotal setae. - — -
19. Presence of coxal bristles. = ~

20. No. of fleshy pleural setae on 7th
abdominal segment. -

21. No. of fleshy ventral setae on
7th abdominal segment. - ~

22. Size of aedeagus (e.g. in relation to
basal rod or aedeagus). =

Relationships of the Coccidae with other Subdivisions of the Coccoidae

The division of the Coccoidea into the margaroid, lecanoid and diaspidoid types
by Balachowsky (1937, 1942) has been generally supported by the results of more
detailed investigations of the adult males (Theron, 1958), as well as by cytological
studies (Hughes-Schrader, 1942 ; Brown, 1959). Of the families of which the
males have been studied intensively, the margaroid type includes the Margarodidae
and Phenacoleachiidae (Theron, 1958, 1962), the lecanoid type the Pseudococcidae
and Coccidae, and the diaspidoid type the Diaspididae. Theron (1958) showed
that the Pseudococcidae and Coccidae were very closely related and that their
closest relatives were found in the margarodid Steingelia on the one hand and the
Diaspididae on the other. He regarded the diaspidoids as being more closely related
to the lecanoids than the latter are to the margaroids and this view was supported
by Ghauri (1962).

Theron’s conclusions regarding the affinities of the Coccidae were based mainly
on the one species that he studied in detail, Parthenolecanium pomeranicum. The
results of the present investigation, in which a fairly representative sample of the
Coccidae was studied, generally confirm and supplement Theron’s observations.
A few structures of morphological importance, which were either overlooked or
absent in the species studied by Theron, can be recorded here :

1. Vestigial dorsal part of the nudcranial ridge. This short median ridge is present
in some members of the EFULECANIUM and COCCUS groups, though not in the
species studied by Theron. It is more fully developed in the Pseudococcidae
(Theron, 1958 ; Giliomee, 1961), but absent in the Diaspididae (Ghauri, 1962).
The greater reduction or absence of the dorsal section of the midcranial ridge (con-
sidered to be primary) is a specialization in the Coccidae.

2. Interocular ridge. This ridge, which connects the pre- and postocular ridges
below the ocellus, is present and always well developed in the ERIOPELTIS group.

154 MORPHOLOGY AND TAXONOMY OF ADULT MALES

It also occurs, although in a more or less reduced condition, in most specimens of
Genus A. One is tempted to suggest that this condition was the forerunner of the
situation in the Pseudococcidae where the pre- and postocular ridges are fused, but
the fact that both the ERIOPELTIS group and Genus A are comparatively special-
ized, though with little affinity between them, makes this seem unlikely. Pre-
sumably the ridge evolved independently to support the preocular ridge near the
point where it articulates with the antenna, which is exceptionally long in Eviopeltis
and fairly long in Genus A.

3. Anterior tentorial pits. These structures can be seen in the ERIOPELTIS
group and in some members of the EULECANIUM group (e.g. Ericerus and
Phyllostroma). They are absent in the Pseudococcidae,where the anterior tentorial
arms are fused with each other and with the cranial apophysis. In the Diaspididae
the tentorium is absent altogether, but vestigial anterior and posterior tentorial
pits remain in some species (Ghauri, 1962).

4. Lateral pronotal sclerite. A small lateral pronotal sclerite is present in all
Coccidae. Homologous structures are found in the Pseudococcidae (Giliomee,
1961) and Diaspididae (Ghauri, 1962).

5. Post-tergite. The absence of this sclerite in the Coccidae is listed as a special-
ized character by Theron (1958). It was observed, however, in all the species
studied here, including P. pomeranicum. It is present in all the other Coccoidea,
except Steingelia (Theron 1958).

6. Metapleural apophyses. Vestigial metapleural apophyses, of the same nature
as found in Psewdococcus (Giliomee, 1961), are present.

7. Basalave. This structure is vestigial or absent in P. pomeranicum (and the
other members of the COCCUS group) and this was accordingly regarded by Theron
as a specialized feature of the Coccidae. It was found, however, in all the members
of the EULECANIUM, ERIOPELTIS and INGLISIA groups, where the con-
dition is very similar to that in the Pseudococcidae (Theron, 1958) and the
Diaspididae (Ghauri, 1962).

8. Metasternal sclerite. A distinct metasternal plate was found in most of the
species studied, including P. pomeranicum. This is a primitive character, present
in all the Coccoidea except the Pseudococcidae. The Pseudococcidae, on the other
hand, have metasternal apophyses (Giliomee, 1961), which are absent in the
Coccidae.

9. Sclerotization of the abdomen. In P. pomeranicum and the Pseudococcidae
studied by Theron and Giliomee, the abdomen is considerably desclerotized and this
is regarded as a specialized condition which the lecanoids share with the diaspidoids
(Ghauri, 1962). However, it was found that in some of the genera of the Coccidae,
like Luzulaspis, Nemolecamum, Eulecanium, Physokermes and Phyllostroma both
tergal and sternal plates (although reduced) were present. This is undoubtedly
primitive.

This study has shown therefore that, as far as the relationship with the margaroid
type is concerned, only minor alterations are necessary in the list of characters
given by Theron (1958) that separate the lecanoid type from the basic margaroid

OF THE FAMILY COCCIDAE 155

type. That is in so far as metasternal and abdominal sclerites are not absent in
the Coccidae.

The relationships with the basic margaroid male and the diaspidoid male, as
expressed by the characters shared with the lecanoid male, is shown in Table 5.

TABLE 5

SHOWING RELATIONSHIPS OF LECANOID MALE WITH THE MARGAROID
AND DIASPIDOID TYPES

Margaroid Lecanoid Diaspidoid

A. PRIMITIVE CHARACTERS:

1. Head well sclerotized. x x -

2. Tentorium present. x- x -

3. Distinct propleural apophyses present. x x -
B. SPECIALIZED CHARACTERS:

1. Compound eyes absent. - x x

2. Lateral branches of midcranial ridge

present. = x x

3. Cranial apophysis long. — x x

4. Pronotal ridges present. - x x

5. Post-tergites small. - x x

6. Prescutal ridges shifted medially. - x x

7. Prealare differentiated into a

triangular sclerite. = x x

8. Subepisternal ridge detached from

marginal ridge. - x x

9. Abdominal spiracles absent. - x x

From Table 5 it can be seen that the lecanoids and diaspidoids have a large
number (9g) of specialized characters in common, whereas 3 primitive ones are
shared by the lecanoids and margaroids. It can therefore be stated that the
lecanoids are more closely related to the diaspidoids than to the margaroids and
also that they are more specialized than the margaroids but less than the diaspidoids
This conclusion confirms the views of Theron (1958) and Ghauri (1962).

156 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Two genera of which the males were studied by Theron (1958, 1962), i.e. Steingelia
and Phenacoleachia, were regarded by him as occupying an intermediate position
between the more primitive margaroids (Margarodes and Pseudaspidoproctus) on
the one hand and the lecanoids (Coccidae and Pseudococcidae) on the other. He
showed that Steingelia and Phenacoleachia were closely related, but that they
differ in a number of striking features. When one investigates whether these
differences throw any light on the relationships of the two genera with the two
lecanoid families, rather interesting results are found, as shown in Table 6. In this
table the characters known to differentiate male Stengelia and Phenacoleachia are
listed and compared with conditions in the Coccidae and Pseudocociddae.

TABLE 6
CHARACTERS DIFFERENTIATING MALE Steingelia AND Phenacoleachia COMPARED WITH
CONDITIONS IN COCCIDAE AND PSEUDOCOCCIDAE

Cocc- Stein- Phenaco- Pseudo-
idae gelia leachia _—coccidae

1. Postoccipital ridge completely absent. x — -

2. Postocular ridge present. x = x x

3. Cranial apophysis extended. x » = Ee

4. Membranous area of scutum present. x < - -

5. Scutellum short, transverse. x x S a

6. Median ridge of basisternum present. x % - =

. Disc pores on body (other than
glandular pouch) absent. x x = a

From the table it can be seen that, in 6 out of 7 cases, similar conditions obtain
in Steingelia and the Coccidae whereas, of the opposite conditions, all 7 characters
are shared between Phenacoleachia and the Pseudococcidae. These results indicate
that the Coccidae are more similar to Steingelia than to Phenacoleachia, and the
Pseudococcidae more to Phenacoleachia than to Steingelia. At this stage of research
it is difficult to determine, however, whether the similarities have any phylogenetic
significance or whether they are merely due to convergence.

In a number of characters, male Coccidae differ from male Pseudococcidae, the
only other lecanoid family of which the male has so far been studied in detail
(Theron, 1958 ; Giliomee, 1961). These characters are listed in Table 7 and com-
pared with conditions obtaining in the basic margaroid (i.e. excluding Steingelia
and Phenacoleacha) and diaspidoid types.

OF THE PAMILY ‘COCCIDAE 15

“I

TABLE 7

CHARACTERS DIFFERENTIATING MALE COCCIDAE AND PSEUDOCOCCIDAE COMPARED
WITH CONDITIONS IN MARGAROID AND DIASPIDOID TYPES

Margar- _Pseudo- Coce- Diasp-
oid coccidae idae idoid
A. PRIMITIVE CHARACTERS:
rt. Anterior tentorial pits/arms separate. = x x

2. Membranous area of scutum present. - x -

3. Median ridge of basisternum present. - x x
4. Metasternal sclerite present. x —
5. Ostiole absent. x - x x

B. SPECIALIZED CHARACTERS:

1. Postoccipital ridge absent. - - x -
2. Scutellum short, transverse. - ‘% x
3. Metasternal apophyses absent. - - x x
4. Penial sheath elongated. - - x x

5. Disc pores on body (other than
glandular pouch) absent. - - x x

A few remarks should be made concerning the interpretation of the phylogenetic
significance of the membranous area of the scutum (character A2) as indicated in
Table 7. In the primitive margaroid type it is absent in Margarodes (Theron, 1958)
and also in Phenacoleachia (Theron, 1962) but developed to some extent in Pseud-
aspidoproctus, Steingelia (Theron, 1958) and IJcerya (Balachowsky, 1937); in the
specialized diaspidoid type it is absent (Theron, 1958 ; Ghauri, 1962). Similar
considerations make the interpretation of the median ridge of the basisternum
(character A3) difficult. The occurrence of both these structures over a wide
range within the superfamily, including decidedly more primitive forms, does
suggest that, as far as the Coccidae and Pseudococcidae are concerned, their presence
signifies a primitive condition. For the opposite to be true it would mean that
these structures were absent in their common ancestor and have been evolved
independently in the Coccidae, which seems unlikely. As mentioned earlier, two

158 MORPHOLOGY AND TAXONOMY OF ADULT MALES

of the specializations that Theron (1958) recorded, i.e. the absence of a post-tergite
and basalare are not valid as they were found to be present in this family.

From Table 7 it can be seen that the two families share an equal number of
primitive and specialized features and on the basis of these facts alone it is difficult
to decide which of the two families is more specialized. It is clear from the table
that the Coccidae show an overall affinity with the more specialized Diaspididae,
sharing 8 out of 10 characters with that family, whereas the Pseudococcidae have
only 2 out of 10 characters with the Diaspididae in common. It can therefore be
said that the Coccidae are possibly more specialized than the Pseudococcidae. It
will also be remembered that the membranous area of the scutum and the median
ridge of the basisternum have somewhat speculatively been interpreted as being
primitive features. Should the opposite be true, it would prove decisively (by 7
specialized characters to 3 primitive) that the Coccidae were more specialized than
the Pseudococcidae. The conclusion that the Pseudococcidae are more primitive
than the Coccidae supports the opinions based on the evidence of female characters
(Morrison, 1928 ; Balachowsky, 1942 ; Borchsenius, 1958), cytological studies
(Hughes-Schrader, 1948) and earlier, less comprehensive studies of the male (Theron,
1958 ; Giliomee, 1961).

From the existing literature on the males of other families of the lecanoid type,
scanty as it is, a few tentative conclusions can be reached regarding their relation-
ships to the Coccidae. Thus the illustrations and descriptions by Green (1904-
1909), Russel (1941) and Borchsenius (1960) indicate that the males of the Aster-
olecaniidae have a large membranous area in the scutum, a transverse scutellum
and an elongated penial sheath. In these respects they resemble the Coccidae and
at the same time differ from the Pseudococcidae. It can therefore be suggested that
the Asterolecaniidae and Coccidae are closely related, as has been advocated by
Balachowsky (1948) and Ferris (1955) from studies of the females.

From the description of the male of Kermococcus quercus (L.) by Borchsenius
(1960), it appears that the Kermococcidae are more closely related to the Coccidae
than to any of the other Coccoidea of which the males have been studied in detail.
Thus K. quercus possesses 5 pairs of simple eyes ; separate pre- and postocular
ridges ; a large membranous area in the scutum ; a comparatively short, transverse
and possibly tubular scutellum ; a deeply invaginated glandular pouch ; an elong-
ated penial sheath, long aedeagus and a ridge-like basal rod. In all these respects it
differs from the Pseudococcidae. The only character which at present appears to be
exclusive to the Kermococcidae and Pseudococcidae is the two-segmented tarsus.
The males of the Kermococcidae therefore appear to show different relationships
from those assigned to these forms by various workers on female Coccoidea.
Balachowsky (1942, 1948) grouped them together with the pseudococcids and
eriococcids into one family ; Ferris (1957b) regarded them as a subfamily of the
Eriococcidae and grouped them with the Pseudococcidae, amongst others, into the
ramus Eriococci, whereas the Coccidae and Asterolecaniidae were assigned to the
ramus Cocci. Obviously, more detailed studies are necessary before any firm
conclusions can be reached regarding the true relationships of these groups.

OF THE FAMILY COCCIDAE 159
KEYS

As is the case with the preceding descriptions of the species studied, the following
keys contain many more characters than would eventually be necessary for the
purposes of identification. However, it is considered wise to follow Ghauri (1962)
and compile detailed keys because (i) they constitute provisional definitions of the
spuraspecific categories and (ii) because there is little doubt that some of the
characters will eventually prove to be insignificant at the taxonomic levels at which
they are employed here. It is clear from the following keys that both higher and
lower taxa can easily be separated.

KEY TO THE GROUPS OF GENERA

1 With fleshy dorsal head setae, setae between and behind the ventral simple eyes,
genal setae, fleshy postmesospiracular setae, antemetaspiracular setae and ng 2
dorsospiracular setae all present

— With fleshy dorsal head setae, setae between and behind the ventral simple eyes,
genal setae, fleshy postmesospiracular setae, antemetaspiracular setae and fleshy
dorsospiracular setae all absent. F J < 3

2 Antenna shorter than posterior leg ; midcranial ridge ventrally complete ; with 4
simple eyes ; prosternum not spinose ; scutellum tubular, usually more than 3
times (range 2-8—4-1) as wide as long ; basalare not joining pleural wing process
to mesepisternum ; median ridge of basisternum complete ; anterior metasternal
setae more than 36 ; wings usually less than 2} times (range 1-9—2-3) longer than
wide ; with pleural sclerotization only on 7th abdominal segment ; caudal
extension of abdominal segment VII very prominent, tapering ; with a cicatrix
on caudal extension of abdominal segment VIII ; fleshy pleural setae on ab-
dominal segments I-III present ; fleshy setae on sternite VIII not more than 3,

ii Present |. : : . COCCUS group

— Antenna 1-07—1°15 (average ie x0) times longer thin posterior leg ; midcranial ridge
ventrally reduced ; with 8 simple eyes ; prosternum spinose ; scutellum not
tubular, 2:11-2:86 (average 2-36) times as wide as long ; basalare joining pleural
wing process to mesepisternum ; median ridge of basisternum reduced or absent ;
anterior metasternal setae 13-24 (average 18) ; wings 2:33-2-45 (average 2-38)
times longer than wide ; with a band of pleural sclerotization stretching from the
IVth—-VIIth abdominal segments ; caudal extension of abdominal segment VII
small, broadly rounded ; without a cicatrix on caudal extension of abdominal
segment VIII ; fleshy pleural setae on abdominal segments I-III absent ; fleshy
setae on sternite VIII more than 5 ; ‘ INGLISIA group

3 Head (in lateral view) flat ; fleshy ventral head setae present ; broad interocular
ridge present ; postocular ridge not forking below ocellus ; posterior metasternal
setae present, minimum 7 ; wings more than 2} times longer than wide ; ab-
dominal segment VII with pleural sclerotization ; with fleshy ventral setae
occurring posteriorly beyond abdominal segment IV : . ERIOPELTIS group

— Head (in lateral view) dorsoventrally elongated or rounded ; fleshy ventral head
setae absent ; interocular ridge absent or (in Genus A) very narrow ; postocular
ridge forking below ocellus ; posterior metasternal setae usually absent, maxi-
mum 3; wings less than 2? times longer than wide ; abdomen without pleural
sclerotization ; fleshy ventral setae not occurring posteriorly beyond abdominal
segment IV ; ; - ; : : - : . EULECANIUM group

tN

160

nN

MORPHOLOGY AND TAXONOMY OF ADULT MALES

KEYS TO GENERA AND SPECIES
KEY TO THE EULECANIUM Group OF GENERA

Head with anterodorsal bulge somewhat pronounced ; fleshy prosternal setae
present ; fleshy anterior metasternal setae 11-18 (average 14) ; abdominal seg-
ment VIII with a large, cylindrical caudal extension ; fleshy pleural setae present
on posterior abdominal segments and fleshy ventral setae present on abdominal
segments II and sometimes III ; abdominal segments VI and VII usually with
4 and 2 setae respectively ; basal rod 13-2 times as long as aedeagus.

SPHAEROLECANIUM (S. prunastri (Fonsc.))

Head with anterodorsal bulge not pronounced ; fleshy prosternal setae absent ;
fleshy anterior metasternal setae absent ; abdominal segment VIII with a small
convex caudal extension ; fleshy abdominal setae absent ; abdominal segments
VI and VII with either 4 or 2 setae on both segments ; basal rod shorter or as
long as aedeagus . : : : ; : 5 : : : :

Head (in lateral view) rounded ; ventral sclerite near mouth opening present ;
fleshy setae very short and thick, being less than 0-60 times as long as width of 3rd
antennal segment and shorter than width of hind tibia ; roth antennal segment

with 4-6 (average 5) capitate subapical setae : ; ‘ : : Genus A

Head (in lateral view) dorsoventrally elongated ; ventral sclerite absent ; fleshy
setae medium-sized or long, being 0-67—5-00 times as long as width of 3rd antennal
segment, and as long, to 5 times as long as width of tibia ; roth antennal segment
with 2 or 3 capitate subapical setae

Third antennal segment more than 4 times (range 4° pars -3, av erage 5° 3) longer we
wide ; fleshy setae very long, 4—5 times as long as width of 3rd antennal segment
and tibia ; sternites on abdominal segment II absent ; abdominal segment II
with one hair-like seta on each side :

Third antennal segment less than 4 times (range 1-8—3-6) longer than wide ; fleshy
setae medium-sized, less than twice as long as width of 3rd antennal segment and
tibia ; sternite(s) on abdominal segment II present ; abdominal segment II with
2 medially situated hair-like setae : F : : ; : :

Median crest not polygonally reticulated ; scutellum tubular ; with more than 6
hair-like scutal setae E : : : : : : ;

Median crest polygonally rehiouibed scutellum not tubular ; with less than 5
hair-like scutal setae : , : : : ; ‘ :

With 8 simple eyes; with 1-4 Gree 2.2) dorsal ocular setae ; with small
circular pores behind pronotal sclerites, around metatergal setae, on abdominal
segment I and in the ante-anal region ; suspensorial sclerite spot-like ; fore
wings 2:36-2:56 (average 2-46) times longer than wide, with 1-2 (average 1-1)
alar setae on each side ; hind tarsus 5-6—7-o0 (average 6-3) times longer than wide;
without a separate median tergite between abdominal segments I and II ; setae
of glandular pouch with external part 3—4 times as long as the section within the
pouch ; lateral sclerotizations of penial sheath narrowly joined anterior to anus ;
length of basal rod 3 to equal that of aedeagus, basal rod not reaching apex of
basal membranous area ; aedeagus short, the basisternum being 1:-74—1:92

ERICERUS (E. pela (Chav.))

On

(average 1-85) times longer . 3 . CTENOCHITON (C. sp.)

With ro simple eyes ; with no dorsal aed ee ; circular pores absent ; suspen-
sorial sclerite elongate ; fore wings 2:04-2:25 (average 2-09) times longer than
wide, with no alar setae ; hind tarsus 5-1-5-3 (average 5-2) times longer than
wide ; with a separate median tergite between abdominal segments I and II ;
setae of glandular pouch with external part about twice as long as the section
within the pouch ; lateral sclerotizations of penial sheath not joined anterior to
anus ; basal rod less than half as long as aedeagus, reaching apex of basal mem-
branous area ; aedeagus long, the basisternum being only 1:23-1:39 (average

1°34) times longer : : ; : : . FILIPPIA (F. viburni (Sign.))

9

10

OF THE FAMILY COCCIDAE 161

Halteres and suspensorial sclerites absent, metapleural ridge reduced ; fleshy
postmetaspiracular setae present ; abdominal segment III with no hair-like
setae medially ; pleural sclerotizations of penial sheath narrowly joined anterior
to anus ‘ é P PHYLLOSTROMA (P. myrtilli (Kalt.))
Halteres and Socpensecal edlemtes present, metapleural ridge complete ; fleshy
postmetaspiracular setae absent ; abdominal segment III usually with 2 setae
medially, rarely one ; pleural sclerotizations not joined anterior to anus. : 7
With 1-4 (average 2-7) scutal setae ; with only one haltere seta on each side ; with
2 hair-like setae dorsally on abdominal segment I ; penial sheath medium-sized,
the ratio of its length in relation to body length being 1 : 5-09—5-63 (average 5-39).
PALAEOLECANIUM (P. bituberculatum (Targ.))
With no scutal setae ; with more than one haltere seta on at least one side (usually
2, range 1-4) ; usually with no hair-like setae dorsally on abdominal segment I,
occasionally with one ; penial sheath very long, the ratio of its length in relation
to body length being 1 : 3-15—4-15 ‘ ‘ f ‘ : - 2 , 8
Dorsal part of midcranial ridge absent ; with 8 simple eyes ; apex of cranial
apophysis truncate ; antenna longer than half body length, the ratio of its length
in relation to body length being 1 : 1-52~-1-61 (average 1°57) ; antenna also
2°4-2°6 (average 2:5) times longer than penial sheath ; 3rd antennal segment
without hair-like setae ; scutellum 1-5-1-7 (average 1-6) times wider than long ;
legs moderately long, the ratio length of hind leg to body length being 1 : 1-72-
1-83 (average 1-78) ; claws long, hind claw about 14 times as long as width of
tarsus ; abdominal tergites and sternites absent on segments IV and V ; abdom-
inal segment II usually with none, but occasionally with one median hair-like seta
ventrally . , 2 . RHODOCOCCUS (R. spiraeae (Borchs.))
Vestigial dorsal part of tnidcranial ridge present ; with 4, 6 or to simple eyes ;
apex of cranial apophysis bifurcate ; antenna shorter than half body length, the
ratio of its length in relation to body length being 1 : 2-02—2-92 ; antenna only
1-3—2'1 times longer than penial sheath ; 3rd antennal segment with one or
more hair-like setae ; scutellum 1-8—2-3 times wider than long ; legs short, the
ratio length of hind leg to body length being 1 : 2:07-2:47 ; claws medium-sized,
as long or a little longer than width of tarsus ; tergites and sternites present on
all abdominal segments ; abdominal segment II with 2 median hair-like setae
ventrally . : : ; ; . ; ; : 2 . ? . 9
With 4 simple eyes ; mesoprephragma without an emargination ; prescutum with
distinct polygonal reticulation ; basisternum 1-7—2-4 (average 2-1) times longer
than membranous area of scutum ; alar setae absent ; coxal bristles absent ;
apical seta on trochanter 2-1-2 -6 (average 2-4) times as long as width of trochanter;
abdominal segments VI and VII usually with 2 setae on both segments; basal rod
short, length 4—4 that of aedeagus : . PHYSOKERMES (P. piceae (Schr.))
With 6 or 10 simple eyes ; mesoprephragma with aaa emargination ; prescutum
with weak or no reticulation ; basisternum 2-5—3-6 times longer than mem-
branous area of scutum ; alar setae present on oa least one side (range 0-3) ;
coxal bristles present ; apical seta on trochanter 3-4 times as long as width of
trochanter ; abdominal segments VI and VII usually with 4 setae on both
segments ; basal ridge medium-sized, length 2—% that of aedeagus . : 10
Degree of sclerotization of postocular ridge not reduced posteromedially ; with 6
simple eyes ; genae not reticulated ; antenna longer than posterior leg, ratio
I : I'O4—-1'14 (average 1:07) ; 3rd antennal segment 2-5—3-0 (average 2-8) times
longer than wide ; basisternum usually with 1 or 2 hair-like setae ; setae of
glandular pouch with external part about twice as long as the section within the
pouch ; basal rod extending ;%ths of the length from aedeagus to apex of basal
membranous area : j : 6 NEMOLECANIUM (N. abietis Borchs.)

162 MORPHOLOGY AND TAXONOMY OF ADULT MALES

— Degree of sclerotization of postocular ridge reduced posteromedially ; with 10
simple eyes ; genae weakly polygonally reticulated ; antennae shorter than
posterior leg, ratio 1 : 0-73-0°87 (average o-80) ; 3rd antennal segment 1-8—2-2
(average 2) times longer than wide; basisternum without setae ; setae of
glandular pouch with external part about 14 times as long as the section within
the pouch ; basal rod extending : or 3 of the length from aedeagus to apex of
basal membranous area F 3 : . EULECANIUM (E. tiliae (L.))

KEY TO THE GENERA AND SPECIES OF THE ERIOPELTIS Group

1 With a pair of median, hair-like ventral head setae distinctly longer than the rest ;
3rd antennal segment 3-0-3:7 (average 3-3) times longer than wide ; prescutum
distinctly polygonally reticulated ; median ridge of basisternum complete ;
tergites present on all the abdominal segments ; glandular pouch absent, repre-
sented by a slight depression with one long seta ; ante-anal region with no fleshy
setae, but 2-6 circular pores ; penial sheath short, the ratio of its length in
relation to body length being 1 : 6-06—7:28 (average 6-66).
LUZULASPIS (L. luzulae (Dufour))
— Without a pair of long, median, hair-like ventral head setae ; 3rd antennal segment
4-6 times longer than wide ; prescutum not distinctly reticulated ; median
ridge of basisternum to a greater or lesser degree reduced ; tergites absent on
abdominal segments III-VI ; glandular pouch present, containing 2 long setae ;
ante-anal region with 1-9 fleshy setae but no pores ; penial sheath medium-sized,
the ratio of its length in relation to body length being 1 : 4:46-4:92. ERIOPELTIS 2
2 Lateral arms of midcranial ridge absent ; apex of cranial apophysis truncate or
rounded ; median ridge of basisternum interrupted, combined length of separate
parts more than half the length of the basisternum ; post-tergital region, and
anterior arms and membranous area of scutum not reticulated.
Eriopeltis ?festucae (Fonsc.)
— Lateral arms of midcranial ridge present ; apex of cranial apophysis trifurcate ;
median ridge of basisternum reduced to a few vestiges of which the combined
length is less than half the length of the basisternum ; post-tergital region, and
anterior arms and membranous area of scutum weakly reticulated . Eriopeltis sp.

KEY TO THE GENERA AND SPECIES OF THE COCCUS GROUP

1 Head with anterodorsal bulge not pronounced ; with 14-32 fleshy postmeso-
spiracular setae ; with 1-6 postalary setae ; segment VIII laterally with a large
semicircular caudal extension, which carries a large cicatrix posterodorsally ;
without 2 long, hair-like ante-anal setae ; with 2-7 fleshy setae lateral to gland-
ular pouch ; abdominal segment VI ventrally with 2 or less hair-like setae, usually
one on each side ; penial sheath with a large, see membranous extension
at the apex ; . CEROPLASTES 2

— Head with anterodorsal bullee Setiouces! - atl more ‘fem 43 fleshy postmeso-
spiracular setae ; rele without postalary setae ; abdominal segment VIII
laterally with a geniculate, mammillate or papilla-shaped caudal extension which
carries a small lateral or large posterior cicatrix ; with 2 long, hair-like ante-anal
setae ; without fleshy setae lateral to glandular pouch ; abdominal segment VI
ventrally with 3 or more hair-like setae, usually 2 laterally and 2 medially ; penial
sheath with a very small or no membranous extension at the apex . : ; 3

un

OF THE FAMILY COCCIDAE 103

Vestigial dorsal part of midcranial ridge present on anterior margin of head ; with
11-21 (average 16) fleshy setae between and behind ventral simple eyes ; 4th
antennal segment with 23-29 (average 26) fleshy setae ; with 2-4 (average 3-2)
coxal bristles ; abdominal segment VII ventrally with 6-11 (average 9-2) fleshy
setae ; aedeagus short, basal rod about 2 times longer and penial sheath
4°I-4'9 (average 4-5) times longer : : : : . Ceroplastes sp.
Dorsal part of midcranial ridge absent ; with 5-9 9 (awerage 7:1) fleshy setae between
and behind ventral simple eyes ; 4th aietectial segment with 11-19 (average 15)
fleshy setae ; coxal bristles absent ; abdominal segment VII with 1~5 (average 3)
fleshy setae ; aedeagus medium-sized, about as oo as basal rod, and penial
sheath 3-0-3-4 (average 3:2) times longer. : . C, berliniae Hall
Post-tergital setae absent ; hind tarsus usually with more hae: like than fleshy setae;
tergites between abdominal segments I and II consisting of a small sclerite on each
side ; abdominal segment VIII laterally with a papilla-shaped caudal extension,
which carries a small cicatrix laterally . : ‘ PARTHENOLECANIUM 4
Post-tergital setae present on at least one side ; hind tarsus with more fleshy than
hair-like setae ; tergites between abdominal segments I and II consisting of a
small sclerite on each side and a separate median sclerite ; abdominal segment
VIII with a geniculate or mammillate caudal extension which carries a large
posterior and small lateral cicatrix respectively F : - = :
Usually with a pair of hair-like medial pronotal setae ; ventral foramen of scutellum
large, length usually more than half that of scutellum ; caudal extension of ab-
dominal segment VII with 4-8 (average 5) fleshy setae.
Parthenolecanium pomeranicum (Kaw.)
Usually without hair-like medial pronotal setae ; ventral foramen of scutellum
small, length usually less than half that of scutellum ; caudal extension of
abdominal segment VII with 11-25 fleshy setae. Parthenolecanium corni (Bouché)
Vestigial dorsal part of midcranial ridge present ; distal part of terminal antennal
segment not constricted ; mesoprephragma with a shallow emargination ; fleshy
scutal setae absent ; femora of medium length, hind femur 3-86—4-75 times
longer than wide ; lateral margin of abdominal segment VIII with a mammillate
caudal extension which carries a very small cicatrix laterally ; a few fleshy ante-
anal setae usually present (range o-7) ; length of basal rod about equal to 1}
times that of aedeagus , : . PULVINARIA 6
Dorsal part of midcranial ridge isan: distal part a terminal antennal segment
somewhat constricted ; ee with deep emargination ; with 7-32
fleshy scutal setae ; femora long and narrow, hind femur 5:98-6:71 times longer
than wide ; lateral margin of abdominal segment VIII with a geniculate or
cylindrical caudal extension which carries a large cicatrix posteriorly ; fleshy
ante-anal setae absent ; length of basal rod more than 1} times that of aedeagus . 7
The two shortest antennal bristles on the terminal antennal segment about half as
long as the fleshy setae ; pedicel, gena and mesoprescutum distinctly polygonally
reticulated ; abdominal segment II usually with one or more (range o-6) hair-
like setae ventrally : : : ; Pulvinaria ?betulae (L.)
The two shortest antennal bristles « on the tenninal antennal segment longer and
thicker than the fleshy setae ; reticulation on pedicel, gena and mesoprescutum
absent or very weak ; abdominal segment II ventrally without hair-like setae.
Pulvinaria acericola (Walsh & Riley)
With 10-22 (average 16) fleshy dorsal head setae ; 4th antennal segment 9-11
(average 10) times longer than wide ; length of 2 shortest antennal bristles about
equal that of the fleshy setae ; prosternum not reticulated ; with 2-9 (average
5-4) fleshy basisternal setae ; abdominal segment VIII with a geniculate caudal
extension ; setae of glandular pouch with external part about 2-24 times as long

UW

164 MORPHOLOGY AND TAXONOMY OF ADULT MALES

as the section within the pouch ; abdominal segments II and III usually with 2

hair-like setae dorsally ; abdominal segment II usually with 2 (range 1-2, average

1-7) hair-like setae ventrally, one on each side ; ante-anal region with 1-5 (average

2°5) small, circular pores : : . Genus B sp. (nr. PULVINARIA)
— With 26-42 (average 35) fleshy dorsal head setae ; 4th antennal segment 5-7

(average 6) times longer than wide ; length of two shortest antennal bristles

about half that of the fleshy setae ; prosternum reticulated ; without basisternal

setae ; abdominal segment VIII with a cylindrical caudal extension ; setae of

glandular pouch with the external part about 4?-6 times as long as the section

within the pouch ; abdominal segments II and III usually without hair-like setae

dorsally ; abdominal segment II usually without hair-like setae ventrally ; ante-

anal region without pores. F é : : COCCUS (C. hesperidum L.)

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SCHRANK, F. von P. 1801. Fauna Boica 2, pt. 1, Niirnberg. 374 pp.

OF THE FAMILY COCCIDAE 167

SIGNORET, V. 1871. Essai sur les Cochenilles ou Gallinsectes (Homopteres — Coccides), 8-e

partie. Annis Soc. ent. Fr. (Sér. 4) 1 : 421-434.

1873. Essai sur les Cochenilles ou Gallinsectes (Homoptéres—Coccides), 1oe et 11e parties.

Annls Soc. ent. Fr. (Sér. 5) 3 : 27-48, 395-448.

1874. Essai sur les Cochenilles ou Gallinsectes (Homopteres—Coccides), 12e et 13e parties.

Annls Soc. ent. Fr. (Sér. 5) 4 : 87-106, 545-558.

SILVEsTRI, F. 1919a. Contribuzioni alla conoscenza degli insetti dannosi e dei loro simbionti.
IV. La coccinigha del prugno (Sphaerolecanium prunastri Fonsc.) Boll. R. Lab. Ent. agyr.
R. Scuola sup. Agric. Portici, 13 : 70-126.

——1919b. Contribuzioni alla conoscenza degli insetti dannosi e dei loro simbionti. V. La

cocciniglia del nocciuolo (Eulecaniwm coryli L.) Ibid. 13 : 127-192.

1920. LDescrizione e notizie del Ceroplastes sinensis 1). Guerc. (Hemiptera, Coccidae).

Ibid. 14 : 3-17.

SNEATH, P. H. A. & SoxaL, R. R. 1962. Numerical Taxonomy. Nature, 193 : 855-860.

SnoperRass, R. E. 1935. Principles of insect morphology. McGraw-Hill Book Co., Inc.,
New York and London. 667 pp.

Soka, R. R. & SNEatTH, P.H. A. 1963. Principles of numerical taxonomy. WV. H. Freeman
and Company, San Francisco and London. 359 pp.

STEINWEDEN, J. B. 1929. Bases for the generic classification of the coccoid family Coccidae.
Ann. ent. Soc. Am. 22 : 197-245.

STICKNEY, F. S. 1934a. The external anatomy of the red date scale Phoenicococcus marlatti

Cockerell and its allies. Tech. Bull. U.S. Dep. Agric. 404 : 162 pp.

1934b. The external anatomy of the Parlatoria date scale, Parlatoria blanchardi Targioni

Tozzetti, with studies of the head skeleton and associated parts. Tech. Bull. U.S. Dep.

Agric. 421, 67 pp.

Sutc, K. 1908. Towards the better knowledge of the genus Lecanium. Entomologist’s mon.

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1931. O skladbé voskovych Stét6 a jejich prislusnych Zl4z u samcu puklic (Lecanium,

Coccidae). Sb. priv. Spol. Mor. Ostravé 1930-1931 : 85-96.

1932. Ceskoslovenské druhy rodu puklice (gn. Lecanium, Coccidae, Homoptera). Acta

Soc. Sci. nat. moravo-Siles. 7 : 1-134.

1941. Mohelnia festuceti n. gn., n. sp. Lecaniidae, Eriopeltini, Coccoidea, Hemiptera).

(In Czech.) Acta Soc. Sci. nat. moravo-Siles. 13 : 1-17.

1942. Diagnosis differentialis inter gn. Pulvinaria et Phyllostroma n. gn. Ent. Listy

5: 5-8.

1943. Zevni morfologie, metamorfosa a béh zivota cervse Phenacoccus aceris Sign. Acta.
Soc. Sct. nat. moravo-Siles. 15 : 1-52.

TARGIONI Tozett!I, A. 1867. Studii sulle Cocciniglie. \WJemorie Soc. ital. Sci. nat. 3 : 1-87.

1868. Introduzione alla seconda memoria per gli studi sulle Cocciniglie, e catalogo dei

generi e delle specie della famiglia dei Coccidi. Atti Soc. ital. Sci. nat. 11 : 694-738.

THERON, J.G. 1958. Comparative studies on the morphology of male scale insects (Hemiptera:
Coccoidea). Annale Univ. Stellenbosch, 34 (A), 71 pp.

1962. Structure and relationships of the male of Phenacoleachia zealandica (Maskell)

(Hemiptera : Coccoidea). Proc. R. ent. Soc. Lond. (A) 37 : 145-153.

Watsu, B. D. & Ritey,C. V. 1868. A new bark louse on the Osage orange. Am. Ent. 1 : 14.

Weser, H. 1928. Skelett, Muskulatur und Darm der schwarzen Blattlaus Aphis fabae Scop.

Zoologica, Stuttg. 28 Heft 76, 120 pp.

1935. Der Bau der Imago der Aleurodinen. Zoologica, Stuttg. 33 Heft 89, 71 pp.

WILKey, R. F. & McKenziz, H. L. 1961. Systematic status of the Pseudococcus maritimus-
malacearum complex of mealybugs. (Homoptera : Coccoidea : Pseudococcidae). Bull.
Calif. Dep. Agric. 50 : 245-249.

168 INDEX

abietis, Nemolecanium, 41 myrtilli, Phyllostroma, 56
acericola, Pulvinaria, 104

Nemolecanium, 41
berliniae, Ceroplastes, 117

?betulae, Pulvinaria, 100 Palaeolecanium, 53
bituberculatum, Palaeolecanium, 53 Parthenolecanium, 108
pela, Ericerus, 67
Ceroplastes, 117 Phyllostroma, 56
Ceroplastodes, 92 Physokermes, 45
aia Ceroplastodes, 92 piceae, Physokermes, 45
peetay 2 : pomeranicum, Parthenolecanium, 113
corni, Parthenolecanium, 108 prunastri, Sphaerolecanium, 74
Ctenochiton, 63 Pulvinaria, 100

Ericerus, 67
Eriopeltis, 77
Eulecanium, 37

Rhodococcus, 49

Sphaerolecanium, 74

sp., Ceroplastes, 121

sp., Ctenochiton, 63

sp., Eriopeltis, 77

sp., Genus A, 70

sp., Genus B, 96

spiraeae, Rhodococcus, 49

?festucae, Eriopeltis, 81
Filippia, 59

Genus A, 70
Genus B, 96

hesperidum, Coccus, 92 ;
theobromae, Inglisia, 88

Te see 88 tiliae, Eulecanium, 37
luzulae, Luzulaspis, 85 viburni, Filippia, 59

Luzulaspis, 85

A (LIST : OF! SUPPLEMENTS
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. Masner, L. The types of Proctotrupoidea (Hymenoptera) in the British
Museum (Natural History) and in the Hope Department of Entomology, Oxford.
Pp. 143. February, 1965. £5.

. Nixon, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera :
Braconidae). Pp. 284; 348 Text-figures. August, 1965. 6.

. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177:
18 plates, 270 Text-figs. August, 1965. {£4 4s.

. SanDs, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera,
Termitidae) from the Ethiopian Region. Pp. 172 : 500 Text-figures. September,
1965. £3 5S.

. AuMAD,I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156:
475 Text-figures. November, 1965. {2 15s.

. Oxapa, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso-
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G04. E.

A REVISION OF
THE ETHIOPIAN SPECIES
AND A CHECK LIST OF THE
WORLD SPECIES OF CLEORA
(LEPIDOPTERA: GEOMETRIDAE)

DS FLETCHER

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Supplement 8
LONDON : 1967

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® REVISION OF
THE ETHIOPIAN. SPECIES
Peon. CeCe List. Or THE
WORLD SPECIES “OF ‘CLEORA

(LEPIDOPTERA: GEOMETRIDAE)

BY
D. S. FLETCHER

British Museum (Natural History)

146 Text-figs. ; 14 Plates ; 9 Maps.

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Supplement 8
LONDON : 1967

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), <tmnstituted im 1949, 1s
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veady. Volumes will contain about three or four
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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. 8 of the Entomo-
logical series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

World List abbreviation: Bull. Br. Mus. nat. Hist.
Entomology.

© Trustees of the British Museum (Natural History) 1967

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 14th February, 1967 Price $3 10s

A REVISION OF
Pre THIOPIAN “SPECIES
AND: A CHBCK LIST .OF -THE
WORLD SPECIES OF CLEORA

(LEPIDOPTERA : GEOMETRIDAE)
By D. S. FLETCHER

CONTENTS

Page

SYNOPSIS 3
MATERIAL EXAMINED 5
ACKNOWLEDGEMENTS . 4
TREATMENT 4
Cleova CURTIS : 4
Taxonomic History 6
Affinities 7
Distribution : : ; : - : : 8
Biology : : : : : ; é - : 9
Species-Groups. : : : : : : : 9
Key to Species. : 4 - : - . : 10
Description of Species : 2 2 : : : 16
CuHeEcK List OF SPECIES : - : : : . : EI2
SPECIES TO BE MOVED FROM Cleova WHEN GENERA ARE AVAILABLE : 114
SPECIES MOVED FROM Cleova TO OTHER GENERA : . 4 115
REFERENCES ; : : $ . : : : 116
INDEX 2 : s : - ; ; : c 118

SYNOPSES

The species of the genus Cleova occurring in the Mascarene islands and continental Africa
south of the Sahara are described and illustrated. A key to the species based on the male
genitalia is included. The 24 species previously known are re-characterized and 22 species and
Io subspecies are described for the first time. A brief history of the genus is given and the
affinities of Cleora with other Old World genera are discussed. <A check list of the world species
of Cleova is included. Species described originally in the genus, but now known to belong
elsewhere, have also been listed ; wherever it has been possible to place these species with
precision, the new binomen is given.

MATERIAL EXAMINED

THE holotypes or lectotypes of all but three of the Ethiopian species have been
studied ; the type of Cleora acaciaria (Boisduval) has not been found, but the
original illustration is adequate for identification ; C. quadrimaculata (Janse) and
C. nigrisparsalis (Janse) have also been determined from original illustrations.

In addition to the material in the collection of the British Museum (Natural
History), which now includes the large and very rich collections of Lord Rothschild,
J. J. Joicey, Charles Oberthtir and L. B. Prout, material has been loaned by the
following :

a D. S. FLETCHER

Muséum national d’Histoire naturelle, Paris ; Monsieur Cl. Herbulot, Paris ;
Musée Royal de l'Afrique Centrale, Tervuren, Belgium ; Zoologisches Museum der
Humboldt-Universitat, Berlin ; Staatliches Museum fiir Naturkunde, Stuttgart ;
National Museum, Nairobi (Coryndon Museum) ; National Museum of Rhodesia,
Bulawayo ; Transvaal Museum, Pretoria.

The material has been listed geographically and, unless otherwise stated, specimens
are in the British Museum (Natural History).

ACKNOWLEDGEMENTS

I wish to thank the many people who have helped by lending material, who have
allowed me to study type specimens, who have answered questions and who have
sent genitalia drawings : Dr. C. W. Benson, Dr. L. A. Berger, Mr. R. H. Carcasson,
Mr. I. F. B. Common, Dr. A. Diakonoff, Dr. H. J. Hannemann, Monsieur Cl.
Herbulot, Dr. H. B. D. Kettlewell, Prof. O. Lindner, Dr. E. C. G. Pimhey andor:
L. Vari. Mr. Carcasson, with his wide knowledge of the topography and vegetation
of continental Africa, has given much valuable help, especially in tracing localities
and describing their vegetation.

TREATMENT

Previously known species have been re-characterized in the same style as those
species described for the first time. The colour names used in the descriptions are
taken from Ridgway (1912, Color Standards and Color Nomenclature).

The wing measurements are those of the smallest and the largest specimens of
each sex, the measurement being double that from the apex of the fore wing to the
centre of the mesothorax. The genitalia of both sexes, where known, of each
species have been described and illustrated. Drawings of the genitalia of a male
and a female have been fabricated to illustrate the range of characters referred to in
the descriptions (Text-figs. 1-3). The scale placed by each drawing represents
one millimetre. The half-tone illustrations have been prepared from photographs
supplied by Mr. M. G. Sawyers of the Photographic Section of this Museum.

Bibliographic references appearing in the text and in the synonymy are given in
abbreviated form. A full bibliography appears at the end of the paper.

In examining the species previously included in Cleova, a number of Old World
species have been found to belong to other genera. These are listed separately after
the check list of included species to be found at the end of this paper. Wherever
it has been possible to place these with precision they have been transferred to
appropriate genera and new combinations are indicated, but a number of species
will probably need new genera erected to accommodate them.

CLEORA Curtis

Cleora Curtis, 1825 : pl. 88. Type-species : Geometra cinctaria Denis & Schiffermiiller, 1775, by
original designation.

Cerotricha Guenée, 1857 : 284. Type-species : Cevotvicha licovnavia Guenée, by monotypy.
Syn.n.

REVISION OF AFRICAN SPECIES OF CLEORA 5

Barsine Meyrick, 1883 : 530 nec Walker, 1854. Type-species : Scotosia panagrata Walker, by
subsequent designation ; citation by Meyrick, 1917 : 266. Syn.n.

Meyrickia Butler, 1884 : 133, n.n. pro Barsine Meyrick, 1883 nec Walker, 1854.

Chogada Moore, 1887: 415. Type-species : Boarmia alienaria Walker, 1860, by original
designation. [First synonymy by Prout, 1928 : 155.]

Carecomotis Warren, 1896 : 402. Type-species : Carecomotis perfumaria Warren, by mono-

typy. Syn.n.
Neocleova Janse, 1932 : 266. Type-species : Boarmia tulbaghata Felder, by original designation.

Syn.n.

Proboscis well developed. Frons and head moderately rough-scaled, with a slight tuft
between antennae and another above proboscis, the latter tuft formed by meeting of long
scaling from each side. Palpus : first segment long-scaled beneath, scaling equal in length to
that of segment ; scaling on second segment one-half as long as that on first ; third segment
smooth-scaled, usually depressed. Male antenna bipectinate to about two-thirds of shaft, the
longest pectinations about one-eighth as long as shaft, decreasing in length apicad and basad ;
the pectinations, arising from the apical margin of each segment, are densely ciliate ventrally
and fully scaled dorsally and occasionally tipped with a bristle ; in some species-groups (cinc-
tavia Denis & Schiffermiiller, sublunaria Guenée, vepetita Butler and alienaria Walker) a second,
unscaled pair of pectinations, two and one-half times as long as the diameter of the shaft, arises
from the basal margin of each segment. In the species rvepetita Butler and perlepidaria Warren
the second pair of pectinations occurs irregularly. Female antenna shortly bipectinate and
ciliate in species groups previously referred to Carecomotis, the pectinations shortening in length
apicad and basad ; the longest pectinations equal in length to twice the diameter of the shaft,
each pair ciliate ventrally and scaled dorsally and arising from the apical margin of each segment ;
in remainder of genus, antenna minutely ciliate with a pair of bristles to each segment. Meta-
thorax with a slight crest ; pectus with long hair-scales. In male, first abdominal tergite
crested ; third abdominal sternite with a cluster of spines on anterior margin in species in which
a hair-pencil is present on the hind tibia. Fore tibia with process extending to tarsal joint ;
male hind tibia usually dilate with a hair pencil and two pairs of spurs. Fore wing : termen
oblique and very slightly crenulate ; a weak fovea in male ; veins #2; and Re free from radius,
rarely from a point or shortly stalked (munditibia Prout, dodonaeae Prout) ; R3-5 on long stalk ;
Mp2 a little closer to M, than to M3 ; Cu, from just before lower angle. Hind wing : termen
slightly crenulate ; Sc + A, approximated to basal half of upper margin of cell ; 5 from before
upper angle. Discal spots on both wings consist of raised scales ringed with fuscous or black.
Wing pattern in genus characteristic, varying mainly in degree of emphasis of certain fasciae and
suffusion of colour. Fore wing : antemedial fascia from one-third costa, shallowly lunulate,
bowed distally in discal area, then inclined to one-fourth inner margin ; medial fascia lunulate
from discal spot to one-half inner margin ; postmedial fascia shallowly lunulate from two-
thirds costa, bowed distally in discal area, then inclined to five-eighths inner margin ; area
distad of postmedial fascia traversed by several ill-defined, lunulate fasciae ; some irroration
proximad of antemedial fascia, but medial area often little marked ; a fuscous streak between
veins M, and Me distad of discal spot. Hind wing : lunulate medial fascia from discal spot to
one-half anal margin ; postmedial fascia lunulate and parallel to termen from two-thirds costa
to two-thirds anal margin ; area distad of postmedial fascia similar to that of fore wing. Ter-
minal interneural spots fuscous on both wings.

Male genitalia. Uncus usually simple and tapered, tip minutely produced (apex spatulate in
some subspecies of onycha Fletcher and in sabulata Fletcher and evenly rounded in albobrunneata
Fletcher) ; gnathus well developed, usually forming a spiculate plate medially ; dorsal margin
valve sclerotized, apex dilate into a membranous pad usually clothed with hair-scales or weak
spines, rarely spiculate (sabulata, albobyunneata) ; ventral margin usually sclerotized and bearing
one or more processes ; a tuft of long hair-scales arises from base of valve near base of juxta ;
aedeagus variable in shape, but usually tapered apicad ; vesica usually with one or more cornuti.

6 DoS. HEE TeGRE IR

Female genitalia. Ovipositor long and retractile, greater in length than the distance from the
ostium to the anterior tip of the bursa copulatrix. Sterigma sclerotized, specifically diagnostic
in form. Bursa copulatrix sometimes wholly membranous, sometimes sclerotized and ribbed
anteriorly ; stellate or disc-like signum present in all species with two pairs of pectinations on
each segment of male antenna, absent from all African species, present or absent in remaining
species.

Taxonomic history.

The genus Cleora is richly represented in the tropical regions of the Old World
and its range extends into the temperate region of both the Old and New Worlds,
being represented in North America by the two species C. sublunaria (Guenée) and
C. projectaria (Walker).

Hitherto, on the grounds of convenience, Cleora has been treated taxonomically
in a variety of ways. Interpreted strictly, the genus has included only those species
in which the male antenna bears two pairs of pectinations on each pectinate segment,
one pair very long and densely ciliate, the other shorter and simple, and in which the
female antenna is simple.

Carecomotis, occurring in the tropical parts of the Oriental region and in the
Australian and Pacific regions, included 29 known species and was separated from
Cleora on the basis of the antennae; the pectinate segments of the male antenna
each bearing one pair of long, densely ciliate pectinations arising from the anterior
edge of the segment, the female antenna bearing one pair of similar, but shorter
pectinations. In both Cleora and Carecomotis the male genitalia conform to a
similar basic pattern, a well developed but simple and tapered uncus, a well
developed gnathus with a scobinate medial plate, a weak juxta, the valve with a well
defined but weakly sclerotized dorsal area, a lobe-shaped cucullus without processes
and a well developed and sclerotized sacculus displaying specifically diagnostic
characters; the aedeagus is usually at least twice as long as broad, usually with one
or more cornuti adorning the vesica. The female genitalia bear a well developed
and specifically diagnostic sterigma and usually a clearly defined colliculum ; the
posterior part of the bursa copulatrix is usually ribbed and sclerotized and the
remainder is membranous, with or without a signum ; when present the signum is
disc-like or stellate.

The Ethiopian representatives of Cleora were separated by Janse (1932 : 266)
on the basis of the antennae, which in the male bear only one pair of long, densely
ciliate pectinations on each pectinate segment and in the female are simple, and
were placed in the genus Neocleora which he erected for them. Subsequent examina-
tion of the female genitalia of these Ethiopian species has shewn them to lack a
signum on the bursa copulatrix. The neuration, male genitalia and habitus of
both sexes are, however, closely similar in general pattern to those species that were
included separately in Cleora and Carecomotis.

Whilst the groups of species that were included separately in Cleova, Carecomotis
and Neocleora each display certain well defined characters in one sex only and each
group has evidently speciated extensively, the more satisfactory treatment groups
all these species in one genus under the oldest available name, Cleora.

The North American species of Cleora were re-described and illustrated by

REVISION OF AFRICAN SPECIES OF CLEORA 7

McDunnough in 1920. The greater part of the Indo-Australian species of Cleora
were treated by Prout in two revisions, I92ga and 1937. The species previously
included in Carecomotis were revised by Fletcher in 1953. The Ethiopian species
of Cleora then known were treated by Janse in 1932 under the name Neocleora.

Affinities. Cleora is closely related to Ascotis Hiibner (1825 : 313), which is
represented in southern Europe by A. selenaria selenaria (Denis & Schiffermiiller),
by selenaria reciprocaria (Walker) in continental Africa and by further subspecies in
India, Ceylon, China and Japan. The neuration is almost identical to that of Cleora,
but Rz sometimes anastomoses briefly with Rs. In Ascotis the male antenna is
ciliate and in the male genitalia the uncus is short and shallowly bifurcate ; in the
female genitalia the sterigma is weakly developed and the presence of a signum
separates it from the Ethiopian species of Cleora ; the characteristic shape of the
signum (with the anterior edge toothed, often strongly) at each corner, separates it
from the remaining species of Cleora.

Also closely related to Cleova is the genus Alcis Curtis (1826 : 113) occurring in
the Palaearctic and Indo-Australian regions. Though the neuration is identical
with that of Cleora, there are a number of other structural differences which clearly
separate it. In the males of species examined (except A. gomphica Wehrli and
A. flavolinearia Leech) the pectinations of the male arise from the apical edge of
each pectinate segment and are unscaled. In the male genitalia the juxta is two-
pronged, the valve has the ventral margin membranous and unadorned, but with a
process at mid-valve arising from mid-dorsal margin, and a characteristically
halberd-shaped cornutus on the vesica. In those examples of the female genitalia
of Alcis so far examined, there are two weakly spiculate signa on the bursa copulatrix.

Hypopalpis Guenée (1862 : 29) was erected for H. terebraria Guenée and H.
perforaria Guenée, since shown by Vinson (1938 : 38) to be synonymous ; terebraria
was placed by Vinson in Cleora. Hypopalpis terebraria and H. antemelaria Mabille
(1893), island endemics on Réunion and Mauritius respectively, have male antennae
of the cinctaria type, with two pairs of scaled pectinations from each pectinate
segment, but in the structure of the genitalia of both sexes these species are closely
related to Ascotis selenaria and they have been transferred to the genus Ascotis.
Hypopalpis terebraria Guenée is here selected for the first time as type-species of
Hypopalpis, which becomes a junior synonym of Ascotis. Syn. n.

Prout (1938 : 154) erected the genus Colocleora for 32 species from the Ethiopian
region, which probably belong to two or more genera. The species agreeing in
structure with the type-species of Colocleora, Alcis ansorget Warren, show some
similarity in habitus to that of Cleora, but structurally are closely related to Ascotis.
In the male the neuration is identical with that of Cleora, but the pectinations of the
antennae arise from the base of each segment and are unscaled ; in the females of
those species of Colocleora available for study, veins Ri and R2 in the fore wing are
long-stalked. In the male genitalia the uncus is usually very short, squat and
triangular and the form of the valve in the male and that of the signum in the
female of those species available for study are similar to those of Ascotis.

The genus Scotorythra Butler (1883 : 177), in which Zimmerman (1958 : 1-2) has

8 D. S. FLETCHER

listed 37 species, is endemic in the Hawaiian Islands. Structurally the species are
related to Cleora, possibly deriving from an ancestor of C. nausori B.-Baker from
Fiji or of C. cheesmanae Prout from the New Hebrides ; they differ markedly,
however, in habitus, in the narrow and proportionately longer wings and in the
unpatterned hind wings.

In the Indo-Australian region the genera Paradromulia, Catoria, Ophthalmodes,
Pseudalcis and Cusiala occur with and approach closely in habitus many species of
Cleora, but are structurally distinct.

Paradromulia Warren (1896 : 300) has veins Ry; and Re of the fore wing long-
stalked in both sexes, Rg anastomosing briefly with Rs. The genitalia differ in
both sexes ; in the male, the uncus is bifurcate and in the female the ovipositor is
appreciably shorter than that found in Cleora.

Catoria Moore (1887 : 414) has veins Rj and Re of the fore wing long-stalked, the
stalk anastomosing briefly with Sc. The male antenna has two pairs of pectinations
to each pectinate segment, as in part of Cleora, but the pectinations are equal in
length and unscaled ; each pair of pectinations is in part fused basally. The genitalia
of both sexes differ in structural pattern from those of Cleora ; in the male the valve
is produced and the uncus is very short and broad with two dorsal digitate processes
recalling those found in Semiothisa Hiibner ; in the female genitalia the ovipositor
is appreciably shorter than that found in Cleora. The genus Catoria was revised
by Prout, 1929c.

Ophthalmodes Guenée (1857 : 283) has neuration similar to that of Catoria ; in
both sexes the antennae are bipectinate, the pectination in the male being longer
than those in the female ; the pectinations in both sexes are unscaled. In the male
genitalia the uncus is reduced and bears lateral processes and the eighth sternum is
specialized. In the female genitalia the ovipositor is short.

In Pseudalcis Warren (1897 : 96) veins Ri and Re in the fore wing are long-
stalked, Ry anastomoses briefly with Sc and R2 with Rs to form a long, slender areole.
The male antennae are bipectinate for the greater part of their length ; the long
pectinations are unscaled and arise from the base of the segment. The genitalia of
both sexes differ appreciably from the pattern of Cleora.

In Cusiala Moore (1887 : 407) veins Ri and Rez of the fore wing are long-stalked,
the male antennae are lamellate and ciliate and the genitalia of both sexes differ
in pattern from those of Cleora.

Distribution.

The species of Cleova in the Ethiopian region, to judge from the limited information
on data labels, appear to be forest insects ; of the 38 species known from continental
Africa, 25 are recorded from the equatorial forest of West Africa, Congo and Western
Uganda ; 12 other species are from areas of montane, lowland, riverine and tem-
perate forests ; C. oligodranes differs in having apparently adapted to the drier
conditions of open, grassy plains.

Eight species appear to be island endemics, five in the Indian Ocean, acaciaria on
Réunion, legrast and macracantha on Madagascar, transversaria and angustivalvis

REVISION OF AFRICAN SPECIES OF CLEORA 9

on the Comoro Is., and three species in the Gulf of Guinea, famsz on Principe I., and
vietter and prosema on Sao Thomé I. Subcincta subcincta is an endemic subspecies
on Sao Thomé I. and the most widely distributed species, rothkirchi, is represented
by an endemic subspecies on the island of Socotra and by another on Madagascar.

The other species occurring on Madagascar, qguadrimaculata, is distributed also on
the coasts of East Cape Province and Natal ; this limited distribution, without
any apparent subspeciation, suggests that its arrival in continental Africa is of
recent origin.

Biology.

Little is known of the biology of the Ethiopian species of Cleora ; herbuloti is
known to have defoliated introduced timber trees, Pinus patula and species of
Eucalyptus, showing an adaptability comparable with the African forest dwelling
species of Buzura, B. abruptaria Walker and B. edwardsi Prout, which extensively
damaged introduced softwoods in Western Uganda in 1961 (W. K. Brown, 1962,
Uganda Forest Department, Technical Note No. 99 (62) ; 1-8, figs.).

Species-Groups.

Among the Ethiopian species of Cleora a number of species-groups are recognizable,
but there remain nevertheless several species which appear to be structurally remote
and which it has not been possible to place with any certainty.

The acaciaria group, which includes acaciaria, transversaria, betularia and flavi-
venata, possesses in common in the male genitalia a smoothly tapered process on the
sacculus, partially enclosed in a membranous fold from the ventral margin of the
valve. In the female genitalia there is a simple lamella postvaginalis and a well-
developed colliculum, tapered anteriorly ; papillifer and cancer are included on the
basis of the male genitalia ; melanochorda is included on the basis of habitus and on
the form of the process on the sacculus, but differs in not having the membranous
fold on the valve developed. The distribution of the acaciaria group is shown on
Map I.

Allied to the preceding is the oculata group, which includes oculata, anacantha,
prosema and epiclithra. In the male genitalia the process on the sacculus extends
parallel with the ventral margin of the valve and is partially enclosed in a mem-
branous fold ; the apex of the process is incurved and bears a dense cluster of spines.
The female of epiclithra is unknown, but the genitalia of the other three species are
diverse in form. Carcassoni is tentatively associated with the oculata group ; the
ventral fold is present and the aedeagus is similar to that of anacantha. The distri-
bution of the group is shown on Map 2.

Species of the vaphis group, which includes vaphis, aculeata and panarista, are
recognizable in the male by the presence of a coarsely scobinate area just basad of
the cucullus and a stout tapered process on the sacculus ; in the female the series
of concentric, medial ridges on the lamella postvaginalis and the form of the ostium
bursae are distinctive. Echinodes and boetschi are placed in the vaphis group on
the basis of the male genitalia. Quadrimaculata and lacrymata are associated

10 D.S. BEECHER

tentatively with the vaphis group, principally on the female genitalia ; though the
males of these two species lack areas of coarse scobination on the valves, quadn-
maculata possesses a well developed process on the valve comparable with that of
echinodes and lacrymata has a well sclerotized apex to the aedeagus and the form of
the cornuti and the presence of a well-defined, sclerotized band on the vesica are
closely similar to those found in vapiis. The distribution of the group is shown on
Map 3.

The tulbaghata group is characterized in the male by the form of the larger of two
cornuti which is either of even width or dilate apically ; the apex is serrate, the
serration extending basad along one side. In the female genitalia the posterior
half of the bursa copulatrix is heavily sclerotized and ribbed, with a shoulder-like
projection at the left side posteriorly. In addition to tulbaghata, the group includes
darget, dactylata, thyris, nigrisparsalis, plax and munda. The distribution of the
tulbaghata group is shown on Map 4.

The rostella group is characterized in the male by the presence on the vesica of
two strongly developed, tapered cornuti which are fused basally, by the weakly
developed apical process on the sacculus and by the presence near the base of the
sacculus of one or more short, setose, digitate processes. The female genitalia are
diverse in form ; there is usually a well-defined lamella postvaginalis with a sclerot-
ized lobe medio-dorsally, often mitre-shaped ; the posterior part of the bursa
copulatrix is usually well sclerotized and sharply contrasted with the membranous,
often globular, anterior part ; the female genitalia of oligodranes and rothkirch are
weakly sclerotized ; those of macracantha are asymmetrical and aberrant. In
addition to rvostella, the species group includes legrasi, angustivalvis, serena, oligo-
dranes, macracantha, derogaria, pavlitzkiae, lima, scobina, radula and rothkircht.

The species toulgoetae and lamotte: are tentatively placed in the vostella group on
the form of the cornuti on the vesica.

The distribution of the vostella group is shown on Maps 5-8.

The species tamst and vietter, endemics on the islands of Principe and Sao Thomé
respectively, though closely related to each other, have been tentatively placed in
the genus ; herbuloti and acerata, another closely related pair, have also been placed
only tentatively ; swbcincta and bicornis appear to be isolated species and each has
been placed in Cleora arbitrarily. Biological data, when available, may help clarify
the affinities of these species. Their distribution is shown on Map 9.

KrEyY TO SPECIES BASED ON MALE GENITALIA

I Valve with area of dense, coarse scobination ventrally just basad of cucullus,
but not extending to dorsal margin : : ‘ : : ; 2
- Valve not so developed 5 6

2 (1) Scobinate area in form of a slender, anne and eon pela euch seashire
ventral margin at one-third ; arm of sacculus curved, spatulate, of even satin
and scobinate at apex, extending in length to mid-cucullus ; short, setose
digitate process medially at one-third valve (Text-fig. 69) : boetschi iss 54)
- Scobinate area and sacculus not so formed . : : ‘ 3

8 (7)
9 (8)
10 (9)
Ir (10)
12 (9)
13 (12)
14 (13)

REVISION OF AFRICAN SPECIES OF CLEORA

Scobinate area extending to cucullus ; process from sacculus almost rhomboid,
posterior dorsal angle produced in short, digitate form (Text-fig. 54)

echinodes (p.

Scobinate area separated from cucullus ; process from sacculus aculeate (Text-
figs. 57, 61, 64) 2
Vesica with three cornuti, but without Scobinate Band (Text: fies 65)

panarista (p.

Vesica with two cornuti and scobinate band (Text-figs. 58, 60) . -
Two smooth, tapered cornuti equal in length ; scobinate band equal in length to
width of aedeagus ; aculeate arm of sacculus two-thirds as long as dorsal

margin of valve (Text-figs. 57, 58) : ’ raphis (p. 4

Two cornuti of unequal length, the longer tipped with four spines ; scobinate
band equal in length to shorter cornutus ; aculeate arm of sacculus one-half

as long as dorsal margin of valve (Text-figs. 60;161)i) ~~. : aculeata (p.

Vesica with single cornutus, shallowly bifurcate at apex ; aculeate arm of
sacculus parallel with and extending to cucullus (Text-figs. 16, 17)

melanchorda (p.

Vesica and sacculus not so formed ,

Arm of sacculus three-fourths as long as dorsal margin oe valve, apex incensed
and densely spined but not covered by a membraneous fold from ventral
margin of valve ; aedeagus smooth ; no cornutus (Text-figs. 37, 38)

anacantha (p.

Sacculus and aedeagus not so formed

Arm of sacculus partially or entirely covered by a setose, nemibranaws fold from
ventral margin of valve ; aedeagus with one or more cornuti

Sacculus not so formed :

Arm of sacculus parallel with ventral eicieren of sealer e, apex incurved and bear-
ing a cluster of spines

Sacculus not so formed

Aedeagus smooth ; vesica with one - tapered cornutus sub- equal in length ia
mid-width of aedeagus ; arm of sacculus three-fifths as long as dorsal margin

of valve (Text-figs. 39, 40) . epiclithra (p.

Apical third of aedeagus scobinate along left half of dorsal surface ; vesica with
two or more cornuti

Arm of sacculus three-fifths as long as dorsal margin of ie, — a short,
semicircular process at base ; vesica with three a sclerotized cornuti

(Text-figs. 34, 35) : : prosema (p.

Arm of sacculus nine-tenths as = as ei margin of valve, without basal
process ; vesica with two cornuti, broad based and strongly sclerotized

(Text-figs. 31, 32) . ; : : oculata (p.

Arm of sacculus with Meise, ne like apex ; ae broadened apicad ;
vesica with two tapered cornuti and a longitudinal scobinate band (Text-figs.

22 2B" ; : : : : cancer (p.

Arm of sacculus with simple, i ae apex ; aedeagus not broadened apicad ;
vesica with two or three tapered cornuti

Arm of sacculus not extended beyond two-thirds bate margin of valve,
tapered and angled aie go° at middle aia -fig. a: ; endemic on island

of Réunion . . acaciaria (p.

Sacculus not so fend species not occurring on ‘alba of Réunion

Arm of sacculus parallel with and extending to four-fifths ventral margin a
valve (Text-fig. 19) ; : : ‘ é papillifer (p

Arm of sacculus arcuate or sinuous, tip extending to dorsal margin of valve

It

31)

28)

13

16)
14

. 25)

15

20

26

(18)

D. 1S) PLEDMCHER

Arm of sacculus bluntly tapered and sinuous (Text-fig. 7) ; wing-span of moth

usually 30-35 mm. ; endemic on Comoro Is. _ . : transversaria (p.

Arm of sacculus finely tapered ; wing-span of moth usually 40-45 mm. ; not
found on Comoro Is.
Aedeagus six times as long as bead one ad one- ale oe as long as 3 deral

margin of valve (Text-fig. 14) : . flavivenata (p.

Aedeagus almost eight times as a ae as bead Pee as long pedoral margin of

valve (Text-fig. 11) . betularia (p.

Arm of sacculus of almost even width, slightly spiral and extending parallel
with and almost to apex of folded ventral margin of valve ; vesica without

cornuti (Text-figs. 41, 42) . : 2 ; F . carcassoni (p.

Sacculus not so formed :

Arm of sacculus slender and spatulate, doce ead panel faced arte dorsi
margin of valve, ventral margin serrate and parallel with ventral margin of
valve ; long, very slender digitate process at base of sacculus (Text-figs. 47,
48) : :

Sacculus not so feed ,

Serrate-edged process on sacculus one ae one- half tees as tone as as:
width of valve (Text-fig. 47) ; endemic on Sao Thomé I.

subcincta subcincta (p.

Serrate-edged process on sacculus twice as long as greatest width of valve
(Text-fig. 48) ; continental Africa. ,

Arm of sacculus extending parallel with apical half of ventral margin of valve,
rasp-like, being densely and coarsely spined for full length of dorsal surface ;
a short, blunt digitate process at base of sacculus i -fig. ”

Sacculus not so formed :

Vesica with two cornuti (Text-fig. tes ‘ : F . herbuloti (p.
Vesica without cornuti c : . : : : acerata (p.

Sacculus with one or more well sclerotized, tapered processes, each as long as,
or longer than uncus, the apices curving through 90°

Sacculus not so formed ‘ : ‘ ‘ :

Sacculus with two tapered processes, each concen to tip ; vesica with one

cornutus equal in length to width of aedeagus (Text-figs. 72, 73) bicornis (p.

Sacculus with one tapered process, the apex serrate-edged ; vesica with two
stout, tapered cornuti fused at base, each two-thirds as long as aedeagus

(Text-figs. 66,67) . . quadrimaculata (p.

Cucullus almost circular ae ee Te ventral margin of valve ;
sacculus with a curved, tapered process arising from near mid-valve, with or
without additional processes

Cucullus and sacculus not so formed ‘

Sacculus with a short truncate process at one- ahird “ete maeete oe nace ;
vesica with one long and one short cornutus (Text-figs. 25, 26) ; endemic

on Principe I. : : : : F : : tamsi (p.

Sacculus smooth along ventral margin and without additional process ; vesica
with one tapered cornutus (Text-figs. 28, 29) ; endemic on Sao Thomé I.

viettei (p.

Scobinate medial plate of gnathus broader than greatest width of cucullus and
with two scobinate arms projecting dorsally, one at each side ; process
from sacculus extends along mid-valve and is tipped with stout spines,
varying in number from I-13 ; vesica with two cornuti, one scobinate, one

tapered and smooth (Text-figs. 51, 52) . : 3 . lacrymata (p.

Genitalia not so formed

subcincta longifibulata (p.

30)

44)
27

27

28

29

30

31

35

(26)

(27)

(29)

REVISION OF AFRICAN SPECIES OF CLEORA 13

Vesica with two cornuti fused at base, the longer finely or coarsely scobinate for
varying distances to and including apex, which may be tapered, truncate or

dilate : 28
Vesica with two cornuti fused at base, ‘the longer asaally smooth, but if scobinate

only lightly so and always smooth in apical eighth : : 35
Longer cornutus coarsely scobinate, scobination extending basad fecal a

but not into basal half ; arm of sacculus not scobinate : 29

Longer cornutus of even width, slightly arcuate and finely scobinate in weppieal
two-thirds ; arm of sacculus ema and densely but finely scobinate

(Text-figs. 141, 142) : radula (p. 104)
Apex of larger cornutus dilate to one and one- chal times width at middle,

apical perimeter with 8—1o spines : - : ; : 30
Apex of larger cornutus of even width or tapered 31

Scobinate medial plate of gnathus broad and shallow, equal i in w idth to @reniliec:
apex of sacculus truncate with a short digitate process inclined towards mid-
valve ; larger cornutus slightly sinuous (Text-figs. 90, 91) : . plax (p. 71)
Scobinate medial plate of juxta produced and narrowed, tip equal in width to
one-third width of cucullus ; apex of sacculus with short, triangular, mem-
branous projection at two-thirds ventral of valve ; larger cornutus
straight (Text-figs. 87, 88) : ; . tulbaghata (p. 67)
Apex of aedeagus produced and narrowed by one-half, irregular in shape ;
are half of shorter cornutus bulbous, apical half slender and curved through

° (Text-fig. 79) . F dactylata (p. 60)

tees of aedeagus narrowed slightly and evenly rounded. - 32
Apex of shorter cornutus curved through 45° and usually bearing 2— 5 spines

varying in size (Text-fig. 76) ‘ 4 : ; : dargei (p. 58)

Shorter cornutus straight and tapered : . : 33

Sacculus with process developed at two-thirds eae wars of valve. : 34

Sacculus without process (Text-fig. 93) : : munda (p. 71)

Scobination of gnathus extending along basal part of subscaphium ; sacculus
with curved, spatulate process at two-thirds ventral margin of valve, the
surface minutely pustulate and setose ; shorter cornutus one-half as long
as longer one (Text-figs. 81, 82) . : 3 thyris (p. 62)
Scobination of gnathus not extending along subscaphium ; a short, smooth
spatulate process from apex of sacculus at two-thirds ventral margin of
valve ; shorter cornutus one-third as long as longer one (Text-figs. 84, 85)
nigrisparsalis (p. 64)
Mid-dorsal margin of valve with a heavily spined, semi-circular dilatation
extending towards mid-valve, spining continuous with that of cucullus ;
apex of sacculus in form of broad, blade-like process (Text-fig. 97)
lamottei (p. 73)

Valve not so developed ; : . : : . : : 36
One or both cornuti spined . : : : : - : : 42
Both cornuti quite smooth . é : : : : 37

Tip of uncus produced, tapered aad dearer apical process of sacculus
slight, about as long as diameter of mid-uncus ; two groups of setose, digitate
processes, one at mid-valve, one nearer venteal margin ; apex of juxta

broadly V-shaped and rugose ; gnathus slender (Text-fig. 103) rostella (p. 77)
Uncus, valve and gnathus not so formed . : 38
Apical process on sacculus sclerotized and ereater in ieee Wes deere of

uncus . 39

Apical process on sagariire = SnISenens aad ious fen ees: of uncus, or
wanting : - : 2 : ; : : 3 e 40

14 D. SURLETCHER

39 (38) Apical process on sacculus strongly sclerotized and extending to seven-eighths
ventral margin of valve (Text-fig. 118) ; endemic on Madagascar
macracantha (p. 87)
= Apical process on sacculus sclerotized as above, but extending only to two-thirds
ventral margin of valve (Text-fig. 121) ; continental Africa only
derogaria (p. 89)
40 (38) Ventral margin of valve quite smooth ; cucullus and most of dorsal margin of
valve densely clothed with very long setae (Text-fig. 105) ; endemic on
Madagascar z ‘ ‘ legrasi (p. 80)
- Valve not so developed . : 41

UNCUS

VENTRAL FOLD «

x WN
NN

HN AN
TINT EASA”

SUBSCAPHIUM |

DORSAL MARGIN

GNATHUS

ARM OF SACCULUS
MEDIAL PROCESS
OIGITATE PROCESS

VENTRAL MARGIN

CRISTAE HAIRS JUXTA SACCULUS

AEDEAGUS

CORNUTI

2

Fics. 1-3. Cleova fabricated genitalia. 1, jg; 2, aedeagus ; 3, 2 (opposite page).

41 (40)

42 (36)

REVISION OF AFRICAN SPECIES OF CLEORA 15

Minute membranous apical process on sacculus at two-thirds ventral margin of
valve ; one or more setose digitate processes at mid-valve (Text-fig. 112)
serena (p. 84)
Valve slightly narrowed at two-thirds ventral margin, but no apical process
developed on sacculus ; two or more minute digitate processes at mid-valve ;
cucullus with 4-8 tubercle-like projections from dorsal margin towards mid-

valve (Text-fig. r09) ; endemic on Comoro Is. . ; angustivalvis (p. 82)
Longer of two cornuti smooth and without spines ; shorter cornutus tipped
with a cluster of spines or edged with a few spines - 43

Longer of two cornuti scobinate on one surface from one-half a8 seven- eighths,
apical eighth eons an” shorter cornutus smooth or partially
scobinate ; : : f , : - ; 44

LAMELLA POSTVAGINALIS
STERIGMA

LAMELLA ANTEVAGINALIS

COLLICULUM

RIBBING

BURSA COPULATRIX

16

Ais)

44

(42)

(49)

D. S. FEBTCHER

Sacculus sclerotized and dilate towards mid-valve in apical third ; apex of
sacculus minutely produced at two-thirds ventral margin of valve ; shorter
cornutus tipped with a cluster of spines or spined at one side in apical eighth
(Text-figs. 144, 145) : : : . rothkirchi (p. 108)
Apex of sacculus setose, sometimes with a dinute, membranous projection at
two-thirds ventral margin of valve ; shorter cornutus finely tapered at apex,
apical half of one surface with 6-7 adpressed spines (Text-figs. 115, 116)
oligodranes (p. 86)
Process from apex of sacculus cylindrical, cygnate and scobinate, one and one-
fourth times as long as width of cucullus ; shorter cornutus smooth (Text-
figs. 136, 137) E ; : : : : lima (p. 100)
Sacculus not so formed F 45
Apical third of sacculus dilate and scobinate, more heavily on inner surkies,
scobination extending to cover over-curved, spatulate apex ; shorter cornu-
tus usually smooth, but occasionally one or two adpressed spines mid-way
along one surface (Text-figs. 99, 100) , : : - mite (p. 76)
Sacculus not so formed . 46
Process from apex of sacculus eae ae acd omen equals in length
to uncus and incurved slightly apicad ; base of gnathus one and one-fourth
times as broad as cucullus, scobination extending posteriorly along sub-
scaphium for distance equal to one-half of its width ; shorter cornutus with
adpressed spines along one surface (Text-figs. I 139) ; scobina (p. 101)
Sacculus not so formed 5 47
Apical third of spacculus pegs spatulate, er produced aod eteen ie
rounded, minutely and densely scobinate ; gnathus broader at base than
cucullus, scobination extending posteriorly along subscaphium (Text-fig. 134)
spake <a saltuensis (p. 98)

Sacculus not so formed : 48
Apical projection from sacculus spatulate and eos aces not jaca
(Text-fig. 127) : : é pavlitzkiae lamella {P. 94)
Sacculus not so formed, apex sensed : 49
Apical projection from sacculus blunt, setose but not scobinate and Gals one-
eighth as broad as cucullus (Text-fig. 124) . pavlitzkiae pavlitzkiae (p. 92)
Apical projection from sacculus spatulate, scobinate and broader than one-
eighth width of cucullus’ . 50
Apical RUSE ne from sacculus one- oer as need as Sisal and incuegee
through 90° (Text-fig. 129) : pavlitzkiae etesiae (p. 94)
Apical projection from sacculus one- Seedy as broad as cucullus and obtusely
incurved (Text-fig. 131) : ; ‘ . pavlitzkiae oriadelpha (p. 98)

Cleora acaciaria (Boisduval)
(Text-figs. 4-6 ; Pl. 1, figs. 147-151 ; Map 1)

Boarmia acaciaria Boisduval, 1833 : 264.

Boarmia acaciaria Boisduval, 1833a : 116, pl. 16: 4.
Boarmia acaciaria Boisduval ; Guenée in Maillard, 1862 : 29.
Cleova acaciaria (Boisduval) Vinson, 1938 : 38.

Neocleova acaciaria (Boisduval) Viette, 1954 : 500.

Neocleova acaciaria (Boisduval) ; Herbulot, 1957 : 234.

3. Crest on first abdominal segment white. Vestiture and wings tilleul buff varyingly
suffused with cinnamon brown and tawny ; tilleul buff ground colour clearly discernible on
wings only in medial area of two examples ; postmedial fascia black, slender and sharply

REVISION OF AFRICAN SPECIES OF CLEORA 17

defined ; in two examples discal spot wholly black (Pl. 1, fig. 147). Under surface of wings
tilleul buff densely suffused with fuscous (PI. 1, fig. 150).

é. Genitalia (Text-figs. 4, 5). Tip of uncus evenly tapered and rounded, and sclerotized as
in figure ; arm of sacculus incurved through 90°, finely tapered and covered, except for tip,
by asetose, membranous fold from ventral margin of valve ; short setose, digitate process at base
ofarm ; apex of aedeagus rounded ; vesica with two cornuti, each one-third as long as aedeagus.

®. Ground colour of wings white, lightly irrorate with drab and fuscous ; postmedial fascia
on each wing black, slender and sharply defined. Under surface of wings white, irrorate and
patterned with fuscous. In two examples, area proximad of antemedial fascia on fore wing and
a broad band distad of postmedial fascia on each wing cinnamon brown (PI. 1, figs. 149, 151).

9. Genitalia (Text-fig. 6). Lamella postvaginalis as in figure ; colliculum narrowed to
two-thirds in anterior half, equal in length to posterior width ; bursa copulatrix pyriform ;
anterior third membranous, remainder lightly sclerotized.

Measurements. 4 38-42mm.; @ 38-48 mm.

Figs. 4-6. C. acaciaria genitalia. 4,3; 5, aedeagus ; 6, 9.

18 Diss PEE UGHER

Ascotis terebraria (Guenée), the only other species known from the island of
Réunion with which acaciaria might be confused, differs externally in the antennae
of both sexes, there being two pairs of pectinations instead of one pair to each
pectinate segment in the male ; in the female there are two pairs of short bristles
instead of one pair to each segment. In both sexes the genitalia are diagnostic.

Acaciaria has been recorded widely in the literature from other parts of the
Mascarene region, from continental Africa and from the Indo-Australian region ;
these records are based on misidentifications, for acaciavia is endemic on the island
of Réunion.

The type, which should have passed to Oberthtir with the rest of the Boisduval
collection, cannot be found either in the British Museum (Natural History) or in the
National Museum in Paris. Boisduval’s illustration is, however, adequate for
identification.

Distribution (Map 1). Island of Réunion.
Material examined. REUNION: (Dr. Roussel), 4 5, 3 2; ibid. (Maillard), rt 9 ;
ibid., 21-30.v.1922 (G. F. Leigh), 3 3, 4 9.

Cleora transversaria (Pagenstecher) comb. n.
(Text-figs. 7-9 ; Pl. 1, figs. 152-165 ; Map 1)

Boayrmia tvansversaria Pagenstecher, 1907 : 97, pl. 6 : 4.

6. Genitalia (Text-figs. 7, 8). Arm of sacculus curved through 90° towards dorsal margin
of valve, then ventrad and again towards dorsal margin, the greater part covered by a setose,
membranous fold from ventral margin of valve ; apex of arm bluntly tapered ; a small, setose
process at base of arm ; apex of aedeagus narrowed and rounded ; vesica with two tapered
cornuti, one with a curved apex and two-fifths as long, the other straight and one-third as long
as aedeagus.

®. Genitalia (Text-fig. 9). Lamella postvaginalis weakly sclerotized and shaped as in
figure ; bursa copulatrix pyriform, posterior two-thirds ribbed and lightly sclerotized, anterior
third membranous.

Measurements. 4 29:5-35mm.; Y 33-37 mm.

Closely related to acaciaria and to the following two species, betularia and
flavivenata, and sharing with acaciaria and betularia a remarkably wide and similar
polymorphism, examples of which are illustrated on Pl. 1. Figs. 152 and 160
illustrate a male and a female of the form most commonly represented in collections.
In the male the tilleul buff ground colour of the wings is irrorate with drab and
fuscous, more densely proximad of the medial fascia on the fore wing and distad
of the postmedial fascia on each wing ; distad of the postmedial fascia on each wing
there is an irregular band of cinnamon buff to cinnamon brown. The female differs
in having the ground colour of the wings white. Figs. 154 and 158 represent a form
similar to the preceding, but with wholly black discal spots. Figs. 157, 161 and
162 represent a form in which the fuscous black medial fascia is strongly developed
on each wing and from which a lateral streak extends distally in the discal fold of
the fore wing. Figs. 153 and 159 represent a form in which the medial area of each

REVISION OF AFRICAN SPECIES OF CLEORA 19

wing and the basal area of the hind wing are almost immaculate, except for the
discal spots. In the male (fig. 153) the remainder of the wings are densely irrorate
with drab ; in the female (fig. 159) the corresponding dark areas are cinnamon
brown. Fig. 155 represents a form in which the wings are densely suffused with
fuscous, except along the costa of the fore wing and in the posterior half of the
hind wing. Fig. 156 represents a form with a dark tilleul buff ground colour to
the wings, intensified terminad ; the medial area in this form, posterior of the
subcostal vein, is intensely fuscous black. Fig. 163 represents a female in which
there is a broad fascia of warm buff instead of the postmedial fascia on each wing
and a similar fascia on the fore wing proximad of the antemedial fascia.

The species evidently represents acaciaria in the Comoro Islands, as betularia and
flavivenata do in continental Africa. So far no representative of this species-group
has been found in Madagascar.

Fics. 7-9. C. trvansversaria genitalia. 7,3; 8, aedeagus ; 9, 9.

20 D. Sv FLETCHER

With so wide a range of colour and pattern, the best diagnostic characters are
to be found in the genitalia ; in the male the shape of the arm of the sacculus and
in the female the form of the sterigma and the proportions of the bursa copulatrix are
distinctive.

Distribution (Map 1). Comoro Islands ; endemic.

Material examined. Holotype g: Gr.-Comoro (Voeltzkow S.), in Zoological
Museum, Berlin.

GRANDE CoMORO : 26.1x.r011 (G. J. Leigh), 1 g; ibid.} 1.1921, 2 9 joa
X.1921, 2 6} sbid., xi-rto2zr, © oO ; abid.,; bred;“x.vin-ro2y, 2g Grande: Gemane
(L. Humblot), 70 3, 70 9. MAYOTTE: 1I-5.v.1g1I (G. F. Leigh), 1 f ; ibid., 27-31.
v.IQII, I 9.

Cleora betularia (Warren) comb. n.
(Text-figs. 10-12 ; Pl. 2, figs. 166-175 ; Map 1)

Chogada betularia Warren, 1897 : 93.
Chogada funesta Warren, 1905 : 398.
Neocleova betularia (Warren) Janse, 1932 : 271, pl. 8 : 7, text-fig. roo.

6. Genitalia (Text-figs. 10, 11). Arm of sacculus arcuate and tapered and, except for tip,
covered by a setose, membranous fold from ventral margin of valve ; aedeagus long and slender,
eight times as long as mean width and twice as long as dorsal margin of valve, with a narrowly
rounded apex ; vesica with two tapered cornuti, each slightly longer than one-third of length of
aedeagus.

9. Genitalia (Text-fig. 12). Sterigma sclerotized as in figure; anterior half of bursa
copulatrix membranous, remainder ribbed and sclerotized, strongly at left side.

Measurements. 4 45-50 mm. (/funesta holotype 35mm.) ; 950mm.

An exceedingly variable species, displaying a range of polymorphism in the male
(Pl. 2, figs. 166-175) similar to that found in tvansversaria ; the few females for study
vary little. In the male lectotype the light buff ground colour of the upper surface
of the wings is evenly irrorate with fuscous, comparable with the typical form of the
palaearctic Biston betularia (Linnaeus) ; several male examples from Marieps Mtn.
are entirely suffused with smoke grey, others have on the fore wing a strongly
developed black medial fascia fusing with a black lateral streak in the discal fold
(Pl. 2, figs. 170, 172). In some examples the medial area of the fore wing and the
proximal half of the hind wing are suffused with black (Pl. 2, fig. 174) ; in other
examples these areas are of the ground colour only very sparsely irrorate with fuscous
(Pl. 2, fig. 168) ; in each of these latter two forms the remainder of each wing is
cinnamon buff. The underside of each wing is white to tilleul buff suffused and
marked with drab, the pattern similar to that of the upperside (PI. 2, figs. 167, 169,
17 Ly 173275)

Closely related to tvansversaria, differing externally in the appreciably greater
size and the rather more produced apex of the fore wing ; differing structurally in
the shape of the arm of the sacculus in the male genitalia and in the shape and
sclerotization of the sterigma and in the degree and extent of the sclerotization of
the bursa copulatrix in the female genitalia.

REVISION OF AFRICAN SPECIES OF CLEORA

Fics. 10-12. C. betularia genitalia. 10,3; 11, aedeagus ; 12, 9.

21

22 D. S. FLETCHER

The holotype of fwnesta collected by G. F. Leigh at Durban and another male
with identical data each has a wing-span of 35 mm. ; these may represent a dry
season form, or if reared, as much of Leigh’s material is, they may be starvlings.

Biology. A pupa was found on Pinus patula in Malawi.

Distribution (Map 1). Malawi ; Rhodesia ; Transvaal ; Natal ; Cape Province.

Material examined. Types. Of the two male syntypes of Chogada betularia
labelled ‘“S. Africa’’, I select as LECTOTYPE the specimen from which the
genitalia slide Geometridae No. 2143 has been made; the paralectotype was
labelled by Warren “ Chogada betularia Warren, Type 3’, but is without abdomen.

Holotype g of Chogada funesta Warren: Natal, Durban, (G. F. Leigh)
Geometridae genitalia slide No. 1575.

MALAWI: Zomba Plateau, pupa ex Pinus patula, emerged I9.xii.1962 (R. C. H.
Sweeny), I § ; Little Ruo Plateau, Mt. Mlanje, 6400 ft., 3.viii.1956 (A. W. R.
McCrae), I §. TRANSVAAL: Marieps Mtn., I-9.i.1926 (G. van Son), 10 3, 4 9, in
British Museum (Natural History), 5 g in Transvaal Museum. NATAL: I g;
Karkloof, 17.1.1917 (E. E. Plait), 1 3 ; ibid., 13.1.1918, 1 § ; Durban (G. F. Leigh),
I$; Balgowan, 5.ii.1951 (K. M. Penniton), 1 g in Transvaal Museum ; ibid.,
12.11.1951, Ig; ibid., g.xii.1951, 1 g ; Mont-aux-Sources, Natal National Park,
I g in Transvaal Museum. CAPE PROVINCE: Katberg, 4000 ft., x.1933 (R. E.
Turner), 1 8 ; Wilderness, Knysna Forest, 20.x-8.xi.1952, at M.V. light (H. B. D.
Kettlewell), 1 9 ; Engcobo, 3.1.1954 (D. A. Swanepoel), 1 § in Transvaal Museum.

Cleora flavivenata sp. n.
(Text-figs. 13-15 ; Pl. 3, figs. 176-179 ; Map 1)

6. Crest on first abdominal segment white ; remainder of vestiture white, irrorate with
drab grey and fuscous black. Upper surface of wings white, irrorate with drab grey and fuscous
black ; transverse fasciae and lateral streak in discal fold of fore wing fuscous black, slender and
sharply defined ; broad cinnamon brown to russet bands proximad of antemedial fascia on
fore wing and distad of postmedial fascia on each wing ; veins, except subcostal vein of fore
wing, streaked with ochraceous buff (Pl. 3, fig. 176). Under surface of wings tilleul buff,
irregularly suffused and patterned with fuscous (PI. 3, fig. 177).

¢. Genitalia (Text-figs. 13, 14). Arm of sacculus arcuate and finely tapered and, except for
tip, covered by a setose, membranous fold from the ventral margin of the valve ; aedeagus six
and one-half times as long as mean width and one and one-half times as long as dorsal margin of
valve, with broadly rounded apex ; vesica with two tapered cornuti, each slightly less than one-
third as long as aedeagus.

©. Similar to male, but drab grey and fuscous black irroration sparse, except in distal third
of each wing ; cinnamon brown to russet bands narrower ; in some examples veins less strongly
streaked with ochraceous buff (Pl. 3, fig. 178). Under surface of wings usually less suffused with
fuscous proximally (Pl. 3, fig. 179).

@. Genitalia (Text-fig. 15). Sterigma sclerotized as in figure ; bursa copulatrix pyriform
with slight projection medio-laterally ; anterior two-thirds membranous, remainder weakly
ribbed and weakly sclerotized.

Measurements: 4.40-45mm.; 9 46-50 mm.

The series from Kirstenbosch and Cape Town displays very little variation in
colour and pattern, except in the degree of dark irroration of the upper surface of

REVISION OF AFRICAN SPECIES OF CLEORA 23

the wings ; specimens from the Knysna localities, Wilderness, Garden of Eden,
Sourflats and the Groot River Pass however are paler, the males markedly so,
resembling the females in colour. The single example from Camps Bay, Cape Town
is also pale ; the examples from Port St. Johns are intermediate.

Closely related to betularia, differing externally, especially in the male, in the
clear ochraceous buff streaking on the veins on the upper surface of the wings.
Structurally the size of the aedeagus in the male genitalia and the pattern of
sclerotization of the sterigma and the shape and weak sclerotization of only the
posterior third of the bursa copulatrix in the female genitalia are diagnostic.
Externally similar in colour and pattern to forms of twlbaghata and munda ; from
these species flavivenata may be clearly separated by the genitalia of both sexes.

Distribution (Map 1). South Africa, Cape Province.

15

Fics. 13-15. C. flavivenata genitalia. 13, g; 14, aedeagus ; 15, 9.

24 D. S. FLETCHER

Holotype 3. Cape Town, Kirstenbosch, 5~-29.xii.1954 (A. J. T. Janse), in
Transvaal Museum.

Paratypes : 13 g, 5 2 with same data as holotype ; Cape Province, Camps Bay,
17.x11.1955 (A. J. Duke), 1 J, all in Transvaal Museum ; Cape Town, iii.1g12 (Lord
Gladstone), 1 $ ; ibid., v.1912, 1g ; Cape Province, Harkerville, 9.iii.1g21 (Dr. H. G.
Breyer), I 3; Knysna, Wilderness, 20.x-8.xi.1952, at M.V. light (H. B. D.
Kettlewell), 6 3, all in British Museum (Natural History) ; Knysna, Sourflats,
22-24.x1.1954 (L. Vari), 1 § ; Knysna, Garden of Eden, 16—20.1.1955 (A. J. T.
Janse), 5 9; Groot River Pass, 21-23.1.1955 (A. J. T. Janse), 12 ; Port St. Johns,
10-22.11.1955 (A. J. T. Janse), 3 3, all in Transvaal Museum.

Cleora melanochorda (Fletcher) comb. n.
(Text-figs. 16-18 ; Pl. 3, figs. 180-182, 184-186 ; Map 1)
Neocleova melanochorda Fletcher, 1958 : 137, figs. 42, 76, 216.

6. Fore wing : medial area white except along costa and distad of broad, black, medial
fascia posterior of vein M3 ; remainder of wing cinnamon drab, lightly irrorate with fuscous and,
distad of postmedial fascia, white ; ante- and postmedial fasciae black, sharply defined ; sub-
terminal fascia fuscous, represented only by spots between costa and vein Scgs and between vein
My,and M3. Hind wing, proximad of medial fascia, white, lightly irrorate with fuscous at base,
otherwise similar to fore wing (PI. 3, fig. 180). Underside of wings tilleul buff ; costa of fore wing
and distal third of each wing varyingly suffused with cinnamon drab and with fuscous at apex of
fore wing ; discal and terminal interneural spots fuscous (Pl. 3, fig. 184).

6. Genitalia (Text-figs. 16,17). Arm of sacculus aculeate, parallel with ventral margin of
valve and extending to cucullus ; a small setose, digitate process at base of arm ; vesica with a
stout cornutus shallowly bifurcate at apex, rather longer than one-half length of aedeagus.

2. Not known from type locality.

@. Genitalia (Text-fig. 18), based on a specimen from Kenya. Sterigma wrinkled laterally ;
two parallel, longitudinal ribs along lamella antevaginalis ; posterior two-thirds of bursa
copulatrix ribbed, posterior third lightly sclerotized ; anterior third membranous.

Measurements. Holotype ¢50mm.; Kenya g$46mm.; 9248 mm.

The colour and pattern of the holotype is striking, but probably represents only
one form of a polymorphic species having a range of variation comparable with that
of betularia. The two specimens from Kenya (Pl. 3, figs. 181, 182, 185, 186) have
the ground colour of the wings light drab ; the transverse fasciae are black and the
postmedial fascia on each wing is broadly edged distally with bister ; the genitalia
of the male are identical with those of the holotype from Uganda ; those of the
female are described above.

Related to betularia and flavivenata and representing the most northerly
penetration so far known of the species-group. The shape of the arm of the sacculus
and of the cornutus in the male genitalia and the configuration of the sterigma in
the female genitalia are diagnostic.

Distribution (Map 1). Uganda (upper limits of montane rain forest on Ruwenzori) ;
Kenya.

REVISION OF AFRICAN SPECIES OF CLEORA 25

Material examined. Holotype g. UGAnpba : Ruwenzori, Nyinabitaba, 8650 ft.,
7-13.Vii.1952 (D. S. Fletcher),Geometridae genitalia slide 1751.

Kenya : Kitale, vii.1958 (C. Howard), 1 9 ; Mt. Elgon, x.1961 (T. H. E. Jackson),
I g in the Coryndon Museum.

Cleora papillifer Prout
(Text-figs. 19-21 ; Pl. 3, figs. 183, 187 ; Map 1)
Cleova papillifey Prout, 1934 : 89.

6. Vestiture white, very lightly irrorate with ochraceous tawny and cinnamon brown.
Fore wing white ; transverse fasciae slender and broken, cinnamon brown ; some cinnamon

18

Fics. 16-18. C.melanochorda genitalia. 16,3; 17, aedeagus ; 18, 9.

26 D.S. PLETCHER

brown irroration in preterminal area of discal fold and near base of subcostal vein ; remainder of
wing irrorate with ochraceous tawny, very lightly proximad of postmedial fascia ; discal spot
weakly outlined in cinnamon brown. Hind wing white, lightly irrorate with ochraceous tawny
distad of postmedial fascia ; medial and postmedial fasciae as on fore wing ; tornus cinnamon
brown (PI. 3, fig. 183). Underside of both wings white, patterned as illustrated with cinnamon
brown (PI. 3, fig. 187).

d. Genitalia (Text-figs. 19, 20). Arm of sacculus tapered, parallel with and extending to
four-fifths ventral margin of valve ; apical part covered by a small, setose, membranous fold
extending from ventral margin of valve ; minute, setose ridge at base of arm ; vesica with two
tapered cornuti, each one one-third as long as aedeagus.

®. Similar to male externally.

@. Genitalia (Text-fig. 21). Lamella postvaginalis with angular projection medio-poste-
riorly ; colliculum weakly sclerotized, shorter than broad ; bursa copulatrix dilate anteriorly,
anterior fifth membranous with a short bulbous projection at right side, remainder strongly
sclerotized and ribbed.

Measurements. ¢24mm.; 9 34 mm.

Fics. 19-21. C. papillifer genitalia. 19,3; 20, aedeagus ; 21, 9.

REVISION OF AFRICAN SPECIES OF CLEORA 27

The wings are only sparsely scaled with the white ground colour, giving them a
pearly, hyaline appearance, recalling, as the author noted in his original description,
species of the neotropical genus Ividopsis ; this quality and the presence of cinnamon
brown irroration in the preterminal area of the discal fold on the fore wing give the
species a distinctive appearance. The structure of the valve indicates a close
affinity with the acaciaria species-group ; the detail of the sacculus in the male and
the form of the sterigma and bursa copulatrix in the female genitalia are diagnostic.

Distribution (Map 1). Central and Western Congo (Leopoldville).

Material examined. Holotype 2: [Conco (Leopoldville)}: Kisantu, 1929
(R. P. J. Van Wing), in Musée Royal de |’Afrique Centrale.

Paratype g¢ : [Conco (Leopoldville)] : Lusambo, Kassai, 12.ix.19g19, in British
Museum (Natural History).

Fics. 22-24. C. cancer genitalia. 22,4; 23, aedeagus ; 24, 9.

28 D. S. FLETCHER

Cleora cancer sp. n.
(Text-figs. 22-24 ; Pl. 3, figs. 188-191 ; Map 1)

6. Vestiture white, densely irrorate with vinaceous buff and fuscous. Fore wing white,
irrorate with vinaceous buff and fuscous, densely along costa ; apex suffused with light drab ;
some ochraceous buff irroration in preterminal area of discal fold and in sub-basal area ; ante-
and postmedial fasciae slender and fuscous black, the latter right-angled on vein My. Hind
wing white, sparsely irrorate with vinaceous buff and fuscous ; some ochraceous buff irroration
in preterminal area of discal fold ; medial fascia, from discal spot to anal margin, and slender
postmedial fascia, the latter right-angled in discal fold, fuscous black. Underside of wings
white patterned with fuscous black.

In two male and three female examples the medial fascia on each wing is strongly developed
and from it a fuscous black horizontal streak extends along discal fold to termen (PI. 3, figs. 188,
189).

o. Genitalia (Text-figs. 22, 23). Arm of sacculus extending from one-half to five-sixths
ventral margin of valve, apex bifurcate and claw-like, partially covered with a setose, membra-
nous fold from the ventral margin of the valve ; process near base of claw-like structure broad
and dentate on inner margin ; aedeagus broadened apicad ; vesica with two tapered cornuti,
each three-sevenths as long as and a longitudinal, scobinate band one-third as long as aedeagus.

©. Similar to male externally (Pl. 3, figs. 190, 191).

2. Genitalia (Text-fig. 24). Lamella postvaginalis strongly sclerotized and shaped as
illustrated ; colliculum with a short lip projecting ventrad ; anterior fifth of bursa copulatrix
membranous, remainder ribbed and sclerotized.

Measurements. 4 44-47mm.; 9 45-49 mm.

The structure of the male genitalia shows a close affinity with betularia ; a
tendency to a comparable variation in wing pattern is also apparent. Externally
the marked right-angling of the postmedial fascia and structurally the shape of the
sacculus and ornamentation of the vesica in the male and the lip on the colliculum
in the female are diagnostic.

Distribution (Map 1). Ethiopia ; Kenya ; Burundi ; Malawi ; Transvaal.

Holotype §. Krnya: Ngong, Nairobi, vii.1954 (Fowler & Coulson).

Paratypes : [ETHIoPIA] ABYSSINIA: Harar, 22.xii.1938 (R. E. Ellison), 1 9;
B.E.A. > Kibwezi, 18.v.i917 (W. Feather), 1 9); 1bid., u-x9z9, 1 9. BuRoNDEE
Usumbura, 900 m., 8.i.1962 (Dv. M. Fontaine), 1 3, in Musée Royal de 1’Afrique
Centrale. [Matawi] Nyasa: Zomba, vi.1923 (H. Barlow), 1 9. RHODESIA :
Salisbury, 25.vi.1918 (O’Neil), 1 J. TRANSVAAL : Louws Creek, near Barberton,
26° S., 31 20” E., x1.19022 (7. G. Walkams), 1S.

Cleora tamsi sp. n.
(Text-figs. 25-27 ; Pl. 4, figs. 1g2-196 ; Map 9)

g. Head cinnamon brown ; patagia fuscous with a few tilleul buff scales ; abdominal crest
tilleul buff ; remainder of vestiture tilleul buff, irrorate with fuscous. Fore wing tilleul buff,
irrorate with ochraceous buff, drab and fuscous ; sub-basal fascia broad and ill-defined ; ante-
and postmedial fasciae fuscous black, sharply defined; horizontal fuscous black streak distad of
postmedial fascia between veins Wz and M3 ; broad fascia of ochraceous buff and drab distad of
and parallel with postmedial fascia ; discal spot white, outlined with fuscous black. Hind
wing similar, but antemedial fascia wanting and medial fascia present (Pl. 4, fig. 192). Under-
side of both wings white, patterned as illustrated with bister ; discal spots fuscous black
(Pl. 4, fig. 193).

REVISION OF AFRICAN SPECIES OF CLEORA 29

g. Genitalia (Text-figs. 25, 26). Membranous cucullus dilate and extending beyond ventral
margin of valve ; sacculus with one truncate and one tapered, spine-tipped process, from which
a second small spine sometimes arises, and also a minute setose digitate process ; vesica with
two cornuti, one one-half as long as and one one-sixth as long as aedeagus.

Q. Similar to male externally, but less densely irrorate proximad of postmedial fascia (Pl. 4,
figs. 194-195).

9. Genitalia (Text-fig. 27). Sclerotization of lamella postvaginalis mitre-shaped in pattern;
posterior edge of lamella antevaginalis semi-circular ; colliculum tapered anteriorly, posterior
edge produced slightly medially ; bursa copulatrix membranous and dilate anteriorly, very
weakly sclerotized and ribbed posteriorly.

Measurements. ¢ 35-40mm.; 9 40-44 mm.

Fics. 25-27. C. tamsi genitalia. 25,3; 26, aedeagus ; 27, 9.

30 D. S$. FLETCHER

A variable species developing forms comparable with those of acaciaria from
Réunion and transversaria from the Comoro Is. (Pl. 4, fig. 196). The male genitalia
of this and the following species, vzetter from Sao Thomé I., each has a markedly
dilate, membranous cucullus, a similarly stout uncus and a similarly shaped
gnathus ; whilst their affinity with each other is clear, their relationship with other
species in the genus is obscure. Because of the similarity of range of variation in
wing-pattern, the dilate cucullus and the well developed sacculus, tamsi and viettei
are placed after those species of the acaciaria group.

It is with pleasure that I name this species after Mr. W. H. T. Tams of this
department, in token acknowledgement of his frequent, kindly advice.

Distribution (Map g). Principe I. ; probably endemic.

Holotype 3. W. Arrica : Principe I., 19.xii.1932 (W. H. T. Tams).

Paratypes : ST. PRINCIPE : 1500-2000 ft., iv—-v.1923 (T. A. Barns), 15 3,32;
PRINCIPE I., 19-22.x11.1932 (W. H. T. Tams), 9 3, 3 9.

Cleora viettei (Herbulot) comb. n.
(Text-figs. 28-30 ; Pl. 4, figs. 197-202 ; Map 9)

Neocleora viettet Herbulot, 1958 : 103, fig. 1.

6 9. Externally similar in both sexes to the preceding species and displaying a similar
range of colour and wing pattern.

6. Genitalia (Text-figs. 28, 29). Cucullus almost circular and extending beyond ventral
margin of valve ; ventral margin of valve smooth and without process from sacculus ; broad-
based, curved and tapered process at mid-valve ; vesica with one tapered cornutus, one-half as
long as aedeagus.

®. Genitalia (Text-fig. 30). Lamella postvaginalis produced and rounded medio-posteri-
orly ; lamella antevaginalis crenulate at each side ; colliculum slenderly produced at each side
posteriorly, tapered anteriorly ; anterior third of bursa copulatrix membranous, remainder
ribbed and very weakly sclerotized.

Measurements. 4 37-43mm.; 2 39-43 mm.

Closely related to the preceding species, tamsz, from which it may be distinguished
structurally ; in the male genitalia by the smooth ventral margin of the valve and
the absence of a process from the sacculus, by the broader-based and more slender
process at mid-valve and by the loss of the second, shorter cornutus on the vesica ;
in the female by the crenulate pattern of the lamella antevaginalis and the long
posteriorly produced and anteriorly tapered colliculum.

Distribution (Map 9). Sado Thomé I. ; probably endemic.

Material examined. Holotype g. SAo THomE: Bombaim, Traz-os-Montes
(450 m.), 6-8.vi.1956, in Muséum national d’Histoire naturelle, Paris.

St. THOME, x—xi.1899 (Mocquerys), 8 g, 10 29; Sao Thomé, 10—24.1.1926, edge of
virgin forest ([. A. Barns), 20 3, 11 9.

REVISION OF AFRICAN SPECIES OF CLEORA 31

Fics. 28-30. C. viettei genitalia. 28, dg; 29, aedeagus ; 30, 9.

Cleora oculata sp. n.
(Text-figs. 31-33 ; Pl. 4, figs. 203-206 ; Map 2)

3. Vestiture white, very lightly irrorate with cinnamon brown, patagia edged with cinnamon
brown. Wings white ; postmedial fascia on each wing slender and bister, edged distally with a
fascia of cinnamon brown, and marked on veins with ochraceous buff ; area distad of postmedial
fascia irregularly irrorate with cinnamon brown ; proximad of medial area, fore wing cinnamon
brown with a broad, sub-basal fascia heavily marked and irrorate with ochraceous buff ; medial

32 D. S. FLETCHER

area of fore wing and area proximad of postmedial fascia on hind wing very sparsely irrorate with
cinnamon brown ; in a few examples medial fascia weakly marked posterior of discal spots,
which are smoke grey broadly ringed with cinnamon brown (PI. 4, fig. 203). Underside.
Costa of fore wing light buff, striate with bister ; remainder of wings white, patterned as illus-
trated with bister (Pl. 4, fig. 204).

g. Genitalia (Text-figs. 31, 32). Valve rhomboid ; arm of sacculus nine-tenths as long as
dorsal margin of valve and straight to incurved and densely spined apex ; apical half of arm
partly covered by a setose, membranous fold ; apical third of aedeagus scobinate along left half
of dorsal surface ; vesica with two short cornuti, one-fourth as long as aedeagus, one rather
longer.

2. Similar to male externally, but white area of upperside of both wings more densely
irrorate with cinnamon buff (Pl. 4, figs. 205, 206).

9. Genitalia (Text-fig. 33). Lamella antevaginalis with deep V-shaped incision medially ;
ductus bursae lightly sclerotized, twice as long as broad ; colliculum not developed ; bursa
copulatrix ovate, anterior half membranous, posterior half sclerotized and ribbed.

Measurements. 4 38-44mm.; 240mm.

Fics. 31-33. C. oculata genitalia. 31, ¢; 32, aedeagus (dorsal view) ; 33, 9.

REVISION OF AFRICAN SPECIES OF CLEORA 33

The structure of the sacculus and the covering of the apical half in a membranous
fold in the male genitalia indicate an affinity between oculata and the following
species, prosema and anacantha. Externally the broad sub-basal fascia on the fore
wing and the large, diffuse discal spots on both wings and structurally the form of
the sacculus and vesica in the male and of the lamella antevaginalis in the female
are diagnostic.

Distribution (Map 2). Nigeria ; Cameroun ; Angola ; Congo (Leopoldville) ;
Uganda.

Holotype g. [CAMEROUN]: Afriq. Occid., Station Kamerun, Johann-Albrechts
Hohe, 1896 (L. Conradt).

Paratypes: NIGERIA: Lokoja, x.1904, rainy season (D. Cator), 1 @.
[CAMEROUN]: Afriq. Occid., Station Kamerun, Johann-Albrechts Hohe, 1898
(L. Conradt), 1 9; Lolodorf, 1894-1895 (L. Conradt), 1 g ; Epulan [Efulen],
I.v.1926 (G. Schwab), 1 3. ANGOLA: Quicolungo, 120 km. N. of Lucala, iv.1936,
800 m. (R. Braun), i 3. [CoNnco (Leopoldville)] : Leopoldville, 27.vii.1954, I gd ;
Uele, Paulis, 8.iv.1956, 2g ; ibid., 5.vi.1956, 1g ; Sankuru, Djeka, 17—18.xii.1952,

Fics. 34-36. C. prosema genitalia. 34, $ ; 35, aedeagus (dorsal view) ; 36, 9.

34 D. S. FLETCHER

26; Dimbelenge, 24.x.1950, 1 § ; Lusambo, 29.viil.1949, 1 g ; ibid., 23.vii—
28.1x.1950, 3d, 3 2 ; Katako-Kombe, 9.xi.1951, 1 J, all collected by Dr. M. Fontaine
and deposited in Musée Royal de l'Afrique Centrale ; W. Kivu, Upper Lowa
Valley, Nr. Masisi, 5000-6000 ft., forest and long grass, ii.1924, wet season (T. A.
Barns), 3 3. UGANDA: Entebbe, 25.vii.1961, light trap (K. W. Brown), 4 d.

Cleora prosema Prout
(Text-figs. 34-36 ; Pl. 4, figs. 207, 208 ; Map 2)

Cleova prosema Prout, 1927 : 194, pl. 20: IT.
Neocleova prosema (Prout) Herbulot, 1958 : 103.

g. Vestiture and wings white, very sparsely irrorate with bister. Fore wing : transverse
fasciae bister, heavily marked at costa, slender and broken elsewhere ; antemedial fascia double
and marked heavily also at inner margin. Hind wing : medial and double postmedial fasciae
slender and bister. Discal spots on both wings very slenderly outlined in bister (Pl. 4, fig. 207).
Underside of wings white, patterned as illustrated with fuscous (Pl. 4, fig. 208).

g. Genitalia (Text-figs. 34, 35). Valve rhomboid. Arm of sacculus three-fifths as long as
dorsal margin of valve, stout and straight and covered by a membranous fold from ventral
margin ; apex of arm incurved and densely spined, base with small, rounded, setose process ;
apical third of aedeagus scobinate along left half of dorsal surface ; vesica with three tapered
cornuti, two closely depressed, the third distinctly separate, each one-seventh as long as
aedeagus.

2. Similar to male externally.

2. Genitalia (Text-fig. 36). Lamella antevaginalis with two slender, tapered ridges
medially ; ductus bursae sclerotized, more heavily anteriorly ; anterior half of bursa copulatrix
membranous, remainder ribbed and lightly sclerotized.

Measurements. 4 35-40mm.; 9 39 mm.

Differs externally from the closely related oculata in the white, weakly patterned
wings ; differs structurally in the male in the shorter arm of the sacculus and the
relatively shorter cornuti on the vesica and in the female in the shape of the lamella
antevaginalis.

Distribution (Map 2). Sao Thomé I. ; endemic.

Material examined. Holotype g. Sado Thomé, 10-24.1.1926, edge of virgin
forest (T. A. Barns), Geometridae genitalia slide No. 2125.

SAo THOME : data as type, 2g, 1 9 ; Sao Thomé, 24.1-25.ii1.1926 (T. A. Barns),
26; ibid., ix-x.1926, 1g; Sao Thomé I., 30.x—24.x1.1932 (W. H. T. Tams), 10 dg.

Cleora anacantha sp. n.
(Text-figs. 37-38; Pl. 4, figs. 209-210; Map 2)

6. Vestiture tilleul buff, patagia tipped with and abdomen lightly irrorate with bister.
Wings white ; area distad of slender, bister postmedial fascia of each wing irrorate with cin-
namon brown to bister, very sparsely on fore wing, densely on hind wing ; cinnamon brown
fascia, marked with ochraceous buff on veins, distad of and parallel with postmedial fascia on
each wing ; proximad of antemedial fascia, fore wing densely irrorate with bister and marked
on veins with ochraceous buff ; basal area of hind wing irrorate with bister ; antemedial fascia

REVISION OF AFRICAN SPECIES OF CLEORA 35

on fore wing and medial fascia on each wing bister (Pl. 4, fig. 209). Underside of wings tilleul
buff, weakly suffused with drab (PI. 4, fig. 210).

6. Genitalia (Text-figs. 37, 38). Ventral margin of valve evenly curved to rounded apex ;
arm of sacculus short and stout, three-fourths as long as dorsal margin of valve, apex incurved

Fics. 37-40. Cleova 3 genitalia. 37-38, anacantha. 37, 3; 38, aedeagus (ventral view) ;
39-40 epiclithrva. 39,3; 40, aedeagus.

36 DS. FLETCHER.

and densely spined ; membranous fold slight ; apical third of aedeagus scobinate along left half
of dorsal surface ; no cornutus.

2. Not known.

Measurement. 4 39 mm.

The structure of the sacculus and the scobinate aedeagus relate anacantha closely
to the preceding two species oculata and prosema. The shape of the valve and the
sacculus, the reduction of the membranous fold and absence of cornuti are diagnostic.

Distribution (Map 2). Congo (Leopoldville).

Holotype g. [Conco (Leopoldville)] : Sankuru, Lusambo, 9.vi.1950.

Paratypes : [ConGo (Leopoldville)] : Lusambo, 15.vi.1949, 1 g ; Uele, Paulis,
I.v.1956, Ig ; all collected by Dr. M. Fontaine, all in Musée Royal de |’ Afrique
Centrale.

Cleora epiclithra sp. n.
(Text-figs. 39, 40; Pl. 5, figs. 211, 212 ; Map 2)

¢. Vertex and first abdominal segment tilleul buff, thorax tilleul buff, irrorate with drab ;
remainder of abdomen snuff brown, irrorate with bister (?discoloured). Upper surface of wings
tilleul buff, suffused with pinkish buff, strongly proximad of sub-basal fascia and along costa on
fore wing and distad of postmedial fascia on each wing and irrorate with black ; sub-basal
fascia broad, straight-margined and black ; remaining pattern black, as illustrated (Pl. 5, fig.
211). Under surface of wings tilleul buff, very weakly patterned with fuscous, as illustrated
(Pies uss 2r2)

3d. Genitalia (Text-figs. 39, 40). Sacculus three-fifths as long as dorsal margin of valve ;
a setose, digitate process at base ; apical half slightly sinuous, the tip bearing a cluster of
spines ; apex covered by a setose membranous fold from ventral margin of valve ; aedeagus
smooth, apex narrowly rounded ; vesica with a short tapered cornutus, subequal in length to
width of aedeagus.

2. Not known.

Measurement. 4 40 mm.

Externally somewhat similar to oculata and to some forms of herbuloti but
differing in the uniformly pinkish buff basal area of the fore wing, the straight-
margined sub-basal fascia and the black colour of the irroration and pattern.
Structurally closely akin to anacantha, but differing in the much shorter sacculus,
the smooth aedeagus and the presence of a short cornutus.

Distribution (Map 2). N.E. Congo (Leopoldville).

Holotype 3. [Conco (Leopoldville)] : Kibali-Ituri, Nioka, 31.viii.1954 (J. Hecq),
in Musée Royal de l'Afrique Centrale.

Cleora carcassoni sp. n.
(Text-figs. 41, 42 ; Pl. 5, figs. 213, 214 ; Map 2)

6g. Vestiture white, irrorate with drab and bister ; patagia edged with bister. Wings white,
irregularly and densely suffused with bister, as illustrated ; posterior distal fourth of fore wing
suffused with warm buff in paratype ; cilia on anal margin of hind wing tilleul buff in type,
warm buff shading to drab tornad in paratype (PI. 5, fig. 213). Underside of both wings white,
irregularly blotched with fuscous ; margins, except inner margin of fore wing, light buff (Pl. 5,

fig. 214).

REVISION OF AFRICAN SPECIES OF CLEORA 37

6. Genitalia (Text-figs. 41, 42). Arm of sacculus of almost even width, slightly spiral, one
surface scobinate in apical half, apex spatulate ; process extends parallel with folded ventral
margin of valve, almost to its apex ; apex of aedeagus narrowly rounded, apical third scobinate
at one side ventrally ; no cornutus.

2. Not known.

Measurement. <4 40-41 mm.

The partially scobinate process arising from the sacculus, the scobinate apical
part of the aedeagus and the absence of cornuti appear to relate carcassoni closely
to anacantha. Strikingly distinct in colour and pattern ; structurally the shape of
the process arising from the sacculus is diagnostic.

It is with pleasure that I name this species after Mr. R. H. Carcasson of the

Coryndon Museum.
Distribution (Map 2). W. Kenya ; S.W. Uganda.

Holotype g. Kenya: Kakamega, ix.1g61 (N. Mitton), genitalia slide Geo-
metridae No. 5292.

Paratype : UGAnpA: Ankole, Kalinzu Forest, xi.1g61 (R. H. Carcasson), t $ in
Coryndon Museum.

42

Fics. 41-43. Cleova genitalia. 41-42, carcassomi. 41,3; 42,aedeagus. 43, acevata, 9.

38 D.S. PFLE@DCHER

Cleora herbuloti (Fletcher) comb. n.
(Text-figs. 44-46 ; Pl. 5, figs. 215-222 ; Map 9)

Neocleova herbuloti Fletcher, 1958 : 136, figs. 44, 85, 215.
Neocleova herbuloti Fletcher ; Anon., 1963 : 4.

3. External characters of colour and pattern vary geographically and are discussed under
respective subspecies.

6. Genitalia (Text-figs. 44, 45). Arm of sacculus in form of a slender rod, densely spined
and rasp-like on its dorsal surface, spiralling ventrad apically, separated by a semi-ovate excava-
tion from a small digitate, setose basal process. Vesica with two cornuti, each subequal in
length to width of aedeagus.

2. Genitalia (Text-fig. 46). Sclerotized part of lamella postvaginalis in form of a horizontal
band with a slight projection medio-posteriorly and a strongly sclerotized, raised lobe medio-
anteriorly ; colliculum one and one-half times as long as broad. Bursa copulatrix lightly
sclerotized in two small areas, one near colliculum, the other anteriorly ; remainder mem-
branous and weakly ribbed.

Two subspecies are described, distinguished by colour and pattern.
Examination of the male genitalia of the limited material available suggests some
degree of geographical variation in the shape of the rasp-like arm of the sacculus.

Fics. 44-46. C. herbuloti genitalia. 44,3; 45, aedeagus ; 46, 2.

REVISION OF AFRICAN SPECIES OF CLEORA 39

In the nominate subspecies, from W. Kivu and W. Uganda, the process is slender
and only the apical fifth of the rasp-like surface is turned ventrad. In specimens
from Kenya, the arm of the sacculus is shorter and broader and the apical third of
the rasp-like surface is turned ventrad ; the genitalia of the single male from Angola
are similar. In specimens from Tanzania, Malawi and Cape Province the arm of
the sacculus is slender and the rasp-like surface is turned ventrad for almost the
whole length. In the single male from Mt. Cameroon the arm of the sacculus is
intermediate in form between that of the nominate subspecies and that of the
Angolan male and the vesica bears three cornuti. There is also some degree of
variation in the stoutness of the aedeagus and of the cornuti ; in examples from
Kenya, Angola, Malawi and Cape Province the aedeagus is scobinate apicad on the
dorsal surface. Until adequate series are available all material, with the exception
of the series of subsp. phaea, has been listed under the nominate subspecies.

The well developed arm of the sacculus has a setose digitate basal process similar
to that found in acaciaria, prosema and epiclithra. Externally the colour and
pattern and structurally the partially scobinate apical area of the aedeagus in some
examples and the short cornuti on the vesica are closely similar to oculata ; the
female genitalia, however, lack the strongly developed ductus bursae of that species.

The excurvation of the arm of the sacculus, between the digitate basal process
and the rasp-like apical part in the male genitalia and the form of the sclerotization
of the lamella postvaginalis in the female genitalia are specifically diagnostic.

Distribution (Map g). Fernando Po ; Angola ; Cameroun ; Nigeria ; Congo
(Leopoldville) ; Uganda ; Kenya; Tanzania ; Malawi; Rhodesia ; Transvaal ;
Cape Province.

Cleora herbuloti herbuloti (Fletcher)
(Pl. 5, figs. 215-218 ; Map 9)

6. Vestiture white, irrorate with cinnamon brown ; abdomen irrorate with fuscous.
Wings white, irrorate with cinnamon brown, densely proximad of antemedial fascia and in
apical area of fore wing and around discal spot and proximad of postmedial fascia on hind wing ;
elsewhere very sparsely ; antemedial fascia on fore wing and postmedial fascia on each wing
slender and cinnamon brown (PI. 5, figs. 216, 217). Underside of wings white, patterned with
fuscous (Pl. 5, fig. 218).

9. Differs from male in the even and denser, darker irroration of the wings with bister and
fuscous.

Measurements. (4 36-45mm.; Q 38-45 mm.

In three male examples, one from Ibadan, Nigeria, one from Rwankwi in Kivu
Province, Congo (Leopoldville) and one from between Nanyuki and Meru, Kenya,
the upperside of the hind wing, proximad of the postmedial fascia, is suffused with
cinnamon brown (PI. 5, fig. 215). In the male from Mt. Mlanje, Malawi and a
female from Mt. Selinda, Rhodesia both fore and hind wings are similarly suffused
distad of the postmedial fasciae ; the basal and apical areas of the fore wings are
fuscous. The second female from Mt. Selinda has the remainder of both wings
fuscous black.

40 D. S. FLETCHER

The white ground colour of the wings and the contrasting concentration of dark
irroration proximad of the antemedial fascia and in the apical area of the fore wing
and around the discal spots of both wings in typical males are subspecifically
diagnostic.

Distribution (Map 9). Fernando Po ; Angola ; Cameroun ; Nigeria ; Congo
(Leopoldville) ; Uganda; Tanzania; Malawi; Rhodesia; Transvaal; Cape
Province.

Material examined. Holotype g. UGaAnpbaA: Kigezi, 5500 ft., (G. D. H.
Carpenter), genitalia slide Geometridae No. 735.

FERNANDO Po: Moka, 28.i-3.11.1933 (W. H. T. Tams), 1 9. NIGERIA: Near
Ibadan, 1g. Mr. CAMEROON : Musake, 6350 ft., 8.1.1932 (M. Steele), 1g. ANGOLA:
Quicolungo, 120 km. N. of Lucala, 800 m., iv.1936 (R. Braun), 1 g. CONGO
(Leopoldville) : W. Kivu, Upper Lowa River, nr. Masisi, 5000-6000 ft., 11.1924
(I. A. Barns), I g, allin British Museum (Natural History) ; Kivu, Rwankwi,
r5.vili.1947 (J. V. Leroy), 1 G5 ibid., 13-x1 19047, 1d) 5) ibid.; iv.1048, go
Mulungu (Tshibinda), 31.1.1956 (J. Hecqg), I g ; Mongbwalu (Kilo), 1937 (Me.
Harford-Jordens), i § ; Sankuru, Djeka, 17.xii.1952 (Dr. Fontaine), 1 § ; Katako-
Kombe, 2.1.1952 (Dr. Fontaine), 1 § ; Kibali-Ituri, Kilo (Mines), 1955 (R. Andry),
I dg, all in Musée Royal de l'Afrique Centrale. BurRuNpb1: Astrida, c. 400 m.,
6.vii.1g61 (Dr. Fontaine), 1 3 in Musée Royal de |’Afrique Centrale. UGANDA :
Ankole, Kalinzu Forest, xi.1961 (R. H. Carcasson), 1 g ; Mabira Forest, Jinja,
x.1962 (R. H. Carcasson), 2 9, all in Coryndon Museum ; Bundibugyo, 3440 ft.,
22.Vili-3.1x.1952 (D. S. Fletcher), 1 3 ; Entebbe, 25.viii.1g61 (W. K. Brown), 3 2.
Kenya : Mt. Kenya, vii.1930 (EF. Barns), 1 § ; Nanyuki-Meru, 6.vii.1930 (E. Barns),
2 6. TANZANIA: Amani, ili-iv.1936 (Cooper), 2 3. MALawi: Mt. Mlanje,
25.1v.1913 (Neave), I g. RHODESIA: Mt. Selinda, 8-9.11i.1954 (H. Cookson), 1 9
and ibid., g-17.iv.1956 (Van Son & Vari), i 9, both in Transvaal Museum.
TRANSVAAL : Kowyn’s Pass, Pilgrim’s Rest Distr., 22.1.1962 (Vari & Leleup),
I g in Transvaal Museum. CAPE PROVINCE: Transkei, Katberg, 1949 (H. B. D.
Kettlewell), i 3.

Cleora herbuloti phaea ssp. n.
(Pl. 5, figs. 219-222 ; Map 9)

6. Ground colour of wings vinaceous buff, white only at termen between veins M3 and Cu,
on each wing, irrorate and patterned with fuscous black as illustrated ; veins ochraceous buff
(Pl. 5, fig. 219). Underside pattern similar to that of nominate subspecies, but marked in iron
grey, postmedial fascia sharply defined on anterior half of each wing ; dark markings of upper-
side show through, giving wings a smoky appearance (PI. 5, fig. 220).

9. Similar to male, but pattern largely obscured by dense fuscous black irroration (Pl. 5,
figs. 221, 222).

Measurements. 4 38-41 mm.; 2 38-46 mm.

Biology. Larvae have been reported (Anon, 1963 : 4) from the Louis Trichardt
area of the Northern Transvaal completely defoliating or severely damaging foliage

REVISION OF AFRICAN SPECIES OF CLEORA 41

of Pinus patula, Eucalyptus cloesiana and E. grandis (saligna) in March and
November.
Distribution (Map 9). Transvaal, Louis Trichardt.

Holotype 3. Soutu Arrica : Entabeni, nr. Louis Trichardt, ex pupa 3.viii.1962,
Div. of Entomology, S. Africa ; genitalia slide Geometridae No. 5242.
Paratypes : Entabeni, nr. Louis Trichardt, 30.ili.1962, 1 g ; data as holotype

Ea,3 -

Cleora acerata sp. n.
(Text-fig. 43 ; Pl. 5, figs. 223, 224 ; Map 9)

g. Vestiture vinaceous buff, densely irrorate with fuscous black ; crest of first abdominal
tergite white. Fore wing vinaceous buff, densely irrorate and patterned with fuscous black ;
sub-basal and antemedial fasciae fused in some examples to form band ; veins ochraceous buff,
colour sometimes suffusing posterior half of wing. Hind wing white anteriorly, vinaceous buff
posteriorly, densely irrorate and patterned with fuscous black ; medial and postmedial fasciae
sometimes fused posterior of discal spot (Pl.-5, fig. 223). Underside white ; fuscous black
pattern of upper side marked in iron grey, but ill-defined (Pl. 5, fig. 224).

6. Genitalia. Arm of sacculus similar to that of preceding species, rasp-like, straight and
parallel with margin of valve, but separated from short, basal, digitate process by semicircular
excavation ; aedeagus not scobinate at apex ; no cornutus.

%. Similar to male externally.

2. Genitalia (Text-fig. 43). Sclerotized lamella postvaginalis in form of an irregular
transverse band of almost even width ; colliculum weakly sclerotized, length equal to width ;
bursa copulatrix membranous, weakly ribbed posteriorly.

Measurements. 4 39-44mm.; 9 42-44 mm.

Closely related to the preceding species and externally similar to the darker forms
of herbuloti phaea ; distinguished structurally in the male by the shape of the
excavation in the sacculus and the absence of cornuti ; distinguished in the female
by the sclerotized pattern of the lamella postvaginalis and the shorter, weaker
colliculum.

Distribution (Map 9). Mt. Cameroon.

Holotype g. Mr. CAMEROON: Mann’s Quelle, 7400 ft., 29.1.1932 (M. Steele),
genitalia slide Geometridae No. 5239.

Paratypes : Mr. CAMEROON: Musake, 6350 ft., 8.i.1932 (M. Steele), 2g, 1 9:
toid., 13.1-1932, 1 2; data as holotype, 2g, 1 9 ; ibid., 2.ii.1932, x 9.

Cleora subcincta (Warren)
(Text-figs. 47-50 ; Pl. 6, figs. 231-241, 245 ; Map 9)

Chogada subcincta Warren, IgoI : 16.
Cleova subcincta (Warren) Prout, 1927 : 194.
Neocleova subcincta (Warren) Herbulot, 1958 : 103.

62. An obscurely marked species varying geographically in size and in colour of underside
of wings ; these characters are discussed under respective subspecies.

42 D. S. FLETCHER

g. Genitalia (Text-figs. 47-49). Uncus long, slender and tapered ; valve broadened
apicad ; arm of sacculus slender and spatulate, partly fused with dorsal margin of valve, ventral
margin serrate and parallel with ventral margin of valve ; long, very slender, setose, digitate
process at base of sacculus ; vesica with two slender, tapered cornuti, one one-third and one
one-fourth as long as aedeagus.

48 49

Vics. 47-50. Cleova genitalia. 47, subcincta subcincta, 3. 48-50, subcincta longifibulata.
48, 3, left valve ; 49, aedeagus ; 50, 9.

REVISION OF AFRICAN SPECIES OF CLEORA 43

9. Genitalia (Text-fig. 50). Lamella postvaginalis weakly sclerotized with shallow, tapered
process medio-posteriorly ; lamella antevaginalis weakly sclerotized at posterior margin ;
colliculum weak and short ; anterior fourth of bursa copulatrix membranous, remainder
weakly sclerotized and ribbed.

Two subspecies are described, characterized by the colour of the underside of the
wings and by the modification of the male genitalia.

The precise relationship of swbcincta is obscure, but the serrate-edged process on
the valve and the digitate process on the sacculus suggest an affinity with
herbuloti. The form of these structures, together with the shape of the sterigma in
the female genitalia, afford diagnostic specific characters.

Distribution (Map 9). Sao Thomé I. ; Angola ; Cameroun ; Congo (Leopold-
ville) ; Uganda ; Kenya ; Tanzania.

Cleora subcincta subcincta (Warren)
(Text-fig. 47 ; Pl. 6, figs. 231-233, 236-238 ; Map 9)

dg. Underside of wings white, suffused with fuscous, except at apex of fore wing ; discal
spots and discal third of each wing more densely suffused than remainder (PI. 6, figs. 236-238).

3g. Genitalia (Text-fig. 47). Serrate-edged process on sacculus one and one-half times as
long as greatest width of valve.

9. Proximal two-thirds of underside of each wing less densely suffused than in male ;
white ground colour discernible at apex and mid-termen of each wing (PI. 6, fig. 237).

Measurements. ¢ 92 33-34 mm.

Distribution (Map 9). Sao Thomé ; endemic.

Material examined. LECTOTYPE 9. Sr. THome, x-—xi.1899 (Mocquerys),
labelled by the author : Chogada subcincta Warr., type 2, by present designation.

Paralectotypes : St. THOME, x—xi.1899 (Mocquerys), 4 9.

St. THOME, xii.1899-i.1901 (Mocquerys), 2 3, 6 9 ; SAo THOME, 10.i-24.11.1926,
edge of virgin forest (T. A. Barns), 5 3, 5 8.

Cleora subcincta longifibulata (Fletcher) comb. n.
(Text-figs. 48-50 ; Pl. 6, figs. 234, 235, 239-241, 245 ; Map 9)
Neocleora subcincta longifibulata Fletcher, 1958 : 137, fig. 84.

6°. Underside of wings in both sexes suffused as in the nominate subspecies, but ground
colour light to warm buff (Pl. 6, figs. 239, 240, 245).

6. Genitalia (Text-figs. 48, 49). Serrate-edged process on valve longer than in nominate
subspecies, being almost twice as long as greatest width of valve.

Measurements. 4 32-38mm.; 9 34 mm.

Distribution (Map 9). Angola ; Cameroun ; Congo (Leopoldville) ; Uganda ;
Kenya ; Tanzania.

Material examined. Holotype g. UGanpa: Bundibugyo, 3440 ft., 22.vili—
3.1x.1952 (D. S. Fletcher), genitalia slide Geometridae No. 1755.

44 DLS.) PVE ACHE ik

CAMEROUN : Johann-Albrechts Hohe, 1896 (L. Conradt), 2 3 ; ibid., 1898, 1 ¢.
Conco (Leopoldville) : Upper Uelle distr., v, 1g ; E. Upper Ituri Valley, 30 miles
S. of Irumu, 3000-3500 ft., vil.1924, dense forest (T. A. Barns), 1 g, all in British
Museum (Natural History) ; Boma a Banana, 1933 (Dr. Van Hoof), 1 g ; Uele,
Paulis, 14.11.1956 (Dr. M. Fontaine), 1 3 ; ibid., 6.viii.1956, 1g ; Mongbwalu (Kilo),
1937 (Me. Harford-Jordens), 1 3,1 2; Kivu, Rwankwi, iii,iv,ix (J. V. Leroy), 43;
Kivu, Nyamunyunye (Mulungu), 20.xii.1955 (J. Hecqg), 1 9 ; all in Musée Royal de
l'Afrique Centrale. UGANDA: Bwamba, v.1958 (R. Carcasson), 1 9 and Bwamba
Terr., ix.1g61 (N. Mitton), 1 J, both in Coryndon Museum ; data as holotype, I 2
and Kigezi, Mafuga forest, 25 miles N.W. Kabale, Rutenga, 7000 ft., 15.vil.1951
(J. A. Burgess), 1 g, both in British Museum (Natural History) ; Mabira forest,
Jinja, x.1962 (R. H. Carcasson), 2 g in Coryndon Museum. KENYA: Kakamega,
v.1957 (Mrs. Board), 1 g in Coryndon Museum. Tanzania: E. Usambara Mts.,
Amani, x.1953 (E. Pinhey), 3 3.

Cleora lacrymata sp. n.
(Text-figs. 51-53 ; Pl. 5, figs. 225, 226 ; Map 3)

g-. Vestiture and wings white, lightly irrorate with bister, apical area of fore wing usually
rather more densely irrorate. Antemedial fascia on fore wing and postmedial fasciae on both
wings slender and bister ; medial fascia present on hind wing posterior of discal spot, bister,
broad and ill-defined ; discal spots on both wings heavily outlined with bister (Pl. 5, fig. 225).
Underside of wings white, suffused and patterned with bister as illustrated (Pl. 5, fig. 226).

g. Genitalia (Text-figs. 51, 52). Medial plate of gnathus scobinate, rather broader than
greatest width of cucullus and with two scobinate arms projecting dorsad, one from each side ;
ventral margin of valve and of cucullus sparsely spined ; process arising from sacculus tipped
with heavily sclerotized spines varying in number, two on left valve and three on right valve in
holotype ; in a topotypical paratype the numbers are five and three respectively and the basal,
spinose process is enlarged ; vesica with two cornuti, one slender, tapered and tipped with two
spines, one-half as long as aedeagus, the other densely scobinate and one-third as long as aedea-
gus.

2. Similar to male externally.

@. Genitalia (Text-fig. 53). Sterigma two-thirds as broad as long and well sclerotized ;
lamella postvaginalis with a series of concentric, semicircular ridges medially ; colliculum rather
longer than broad ; bursa copulatrix cylindrical, anterior third membranous, remainder
ribbed and very weakly sclerotized.

Measurements. 4 40-43mm.; 92 43 mm.

In addition to the variation in the sacculus already mentioned a male from the
Central African Republic has thirteen spines on the left side and nine on the right ;
a male from Cameroun has a single spine on each side ; a male from Ghana is the
converse of the holotype, having two spines on the left side and three on the right.
More material might show some correlation between the ornamentation of the
sacculus and distribution.

When seen in a Series, the clean white ground colour of the wings, the slender,
bister pattern and the ill-defined medial fascia extending posteriorly from the
heavily marked discal spots, the species appears distinctive. In the male genitalia

REVISION OF AFRICAN SPECIES OF CLEORA 45

the shape of the sacculus, the scobinate arms of the gnathus and the cornuti on the
vesica and in the female genitalia the form of the sterigma are diagnostic.

Distribution (Map 3). Ivory Coast ; Ghana; Nigeria ; Cameroun ; Central
African Republic ; Congo (Leopoldville) ; Kenya.

Holotype g. Ivory Coast: Bingerville, I-5.vili.1g15 (G. Melou), genitalia
slide Geometridae No. 5255.

Paratypes : Ivory Coast: Bingerville, 25.v—3.vi.1g15 (G. Melou), 1 g ; ibid.,
II.vi, 1g ; ibid., 1-14.vii, 2g; ibid., 1-5.vili, 1 9; ibid., 1-3.ix, 1g. [GHANA]
GoLp Coast: Ashanti, Goaso (G. N. Gibbs), 1 g. NIGERIA : Mamfe, 11.1958, I ¢
in National Museum of Rhodesia. [CAMEROUN]: Afriq. Occid., Station Kamerun,
Johann-Albrechts Hohe, 1898 (L. Conradt), 1 §. ANGOLA: Quicolungo, 120 km.
N. of Lucala, 800 m., iv.1936 (R. Braun), 1 g ; Fazenda Congulu, Amboim district,

Fics. 51-53. C. lacrymata genitalia. 51,3; 52, aedeagus; 53, 9.

46 D. Ss] PLET CHER

7-800 m., 12-16.iv.1934 (Dr. K. Jordan), 1 g. [CENTRAL AFRICAN REPUBLIC]
OUBANGUI : Bangassou, vi-vii.1958 (P. Labour), 1 g in coll. C. Herbulot, Paris.
[ConGo (Leopoldville)| : Uele, Paulis, 12.1v.1956 (Dr. M. Fontaine), 1 3 in Musée
Royal de l’Afrique Centrale. KErNyA: Mt. Marsabit, 4500 ft., 1.1946 (T. H. E.
Jackson), I 9.
Cleora echinodes sp. n.
(Text-figs. 54-56 ; Pl. 6, figs. 242, 246 ; Map 3)

g. Vestiture white to pinkish buff, irrorate with bister. Fore wing white, varyingly suffused
with pinkish buff and irrorate with bister ; transverse fasciae bister, sharply defined. Hind
wing similar, but pinkish buff suffusion and bister irroration usually greatly reduced proximad of
postmedial fascia ; antemedial fascia wanting, discal spots heavily outlined in bister, but
definition diffuse (Pl. 6, fig. 242). Underside of each wing tilleul buff to light buff, varyingly
suffused with pinkish buff to bister except at apex and terminally between veins M3 and Cu,
(Pl. 6, fig. 246).

6. Genitalia (Text-figs. 54, 55). Arms of gnathus slender, bearing a scobinate medial plate,
incurved medio-ventrally, four-fifths as broad as greatest width of valve ; valve with a coarsely
scobinate area, triangular in shape, extending from mid-valve apicad to cucullus and ventrad
almost to ventral margin ; process arising from mid-valve almost rhomboid, posterior dorsal
corner produced in short, digitate form, slightly asymmetrical, vesica scobinate in apical half
and bearing two tapered cornuti fused at base, one one-fifth and one one-fourth as long as
aedeagus.

©. Similar to male externally.

2. Genitalia (Text-fig. 56). Lamella postvaginalis mitre-shaped ; colliculum twice as long
as broad with tapered lateral projections into bursa copulatrix, which is ribbed and weakly
sclerotized, except at membranous anterior extremity.

Measurements. 4 39-42mm.; 2 39-40 mm.

Related to the preceding species, having in the male genitalia a similar, but less
extensively developed, broad gnathus ; the scobination on the valve, sparse in
lacrymata, is extensively developed ; the process arising from the sacculus is
reduced and unadorned but similar in outline to that of lacrymata.

Externally poorly characterized ; seen in a series, the combination of pinkish buff
suffusion and bister pattern of the upperside of the wings and the usually dense
brown suffusion of the underside is striking, but individual specimens are more
surely determined by the genitalia. The broad, scobinate gnathus, the broad
triangular, spined cucullus and the process arising from the sacculus in the male
and the shape of the sterigma and colliculum in the female are diagnostic.

Distribution (Map 3). Fernando Po; Cameroun; Congo (Leopoldville) ;
Uganda ; Malawi.

Holotype g. UGANDA: Entebbe, ili—v.1895 (Jackson).

Paratypes : FERNANDO Po : 650 ft., end of wet season (Rev. W. Cooper), 1 9.
[CamMEROUN]: Afriq. Occid., Station Kamerun, Johann-Albrechts Hohe, 1896
(P. Conradt), 1 §. [Conco (Leopoldville)] : Upper Uelle distr., Dungu, v., 2 3;
Yakusu (K. Smith), 1 3, all in British Museum (Natural History) ; Uele, Paulis,
18.i111.1956 (Dr. M. Fontaine), 1 g ; Lusambo, 16.vi.1949 (Dr. M. Fontaine), 1 g ; ibid.,
II.vil.1950, Ig ; Sankaru, Katako-Kombe, I1.ix.1952, 1 g ; Kafakumba, xii.1932

REVISION OF AFRICAN SPECIES OF CLEORA 47

(F. G. Overlaet), I 3, all in Musée Royal de l'Afrique Centrale ; W. Kivu, south Lowa
Distr., Lowowo Valley, 4000 ft., mountain forest, 111.1924, wet season (T. A. Barns),
Ig; Escarpment west of Semliki Valley, 20 mls. S.W. of Boga, 3500-4000 ft.,
vil.1924, borders of tropical forest and long grass country (7. A. Barns), 2 3.

55

Fics. 54-56. C. echinodes genitalia. 54,3; 55, aedeagus ; 56, 9.

48 DiS. SVE CHB

UGANDA : Bwamba, iv—v.1g940 (T. H. E. Jackson), 2g ; ibid., x.1942, 1g ; Budongo
v—-vi.1939 (IT. H. E. Jackson), 1 3 ; Budongo (G. W. Jeffery), 1 § ; Kampala,
25.Vil.1925, I g. [MALAw!] NyAsALAnpD: Mt. Mlanje, 15.11.1913 (S. A. Neave),
Ig abids.47 1 1o Ne ae

Cleora raphis sp. n.

(Text-figs. 57-59 ; Pl. 5, figs. 227-230 ; Map 3)

g. Vestiture white, irrorate with drab and bister, patagia edged dorsally with bister.
Fore wing white, densely irrorate with drab ; antemedial fascia ill-defined, postmedial fascia

Fics. 57-59. C.vaphis genitalia. 57,5; 58, aedeagus ; 59, Q.

REVISION OF AFRICAN SPECIES OF CLEORA 49

sharply defined, bister. Hind wing similar, but antemedial fascia wanting ; medial fascia
present, bister. Veins on both wings paler, tinged with cartridge buff (Pl. 5, fig. 227). Under-
side : apex of fore wing and termen of each wing, between veins M3 and Cw, white ; remainder
of wings suffused with fuscous, lightly round discal spots, densely elsewhere (Pl. 5, fig. 228).

In a comparatively fresh specimen (date 1955), the drab and bister colours are replaced by
smoke grey and iron grey and the cartridge buff veins are more conspicuous ; possibly these
colours fade to brown tones with age.

3g. Genitalia (Text-figs. 57, 58). Scobinate medial plate of gnathus as broad as cucullus,
which is membranous and dilate at apex, extending slightly beyond ventral margin of valve ;
scobinate area on valve medio-ventrally ; arm of sacculus aculeate, two-thirds as long as dorsal
margin of valve ; vesica with a short, slender scobinate band equal in length to width of aedeagus
and two stout tapered cornuti, one slightly greater than, one slightly less than one-third as long
as aedeagus.

. (Pl. 5, figs. 229, 230). Similar to male externally.

®. Genitalia (Text-fig. 59). Lamella postvaginalis narrowly sclerotized medially with
concentric ridges ; lamella antevaginalis ribbed and folded ; colliculum one-half as long as
broad ; bursa copulatrix cylindrical, posterior extremity and anterior half membranous,
remainder moderately sclerotized and ribbed, the ribbing extending weakly into anterior half.

Measurements. 4 42-44mm.; 40mm.

The species has a broad scobinate medial plate to the gnathus, similar to that
found in the preceding species lacrymata and in echinodes and a basically similar
sacculus structure ; the scobinate medial area of the ventral margin of the valve
differs from that found in echinodes in being less coarse and completely separate
from the cucullus.

Externally poorly characterized. Structurally the needle-like process arising
from the sacculus and the ornamentation of the vesica in the male and the shape of
the sterigma in the female are diagnostic.

Distribution (Map 3). Congo (Leopoldville).

Holotype g. [Conco (Leopoldville)] : Kassai district (Taymans), genitalia slide
Geometridae No. 2308 in British Museum (Natural History).

Paratypes : [Conco (Leopoldville)| : Tumbulungu, 8.ix.1930 (G. F. de Witte),
I 2; Coquilhatville, 9.1x.1955 (Dr. M. Fontaine), 1 §; Fl. Congo, Bolombo
[3°59"S. 21°24’”E.], vii.1938 (J. Ghesquiére), 1 g ; Uele, Bambesa, 20.ix.1933 (J.
Leroy), 1° ; Sankaru, Lusambo, 6.viii.1950 (Dr. M. Fontaine), 1 3 ; ibid., 8.viii.1950,
I 6; Bena-Dibele, xi.1g21 (L. Verlaine), 1 g, all in Musée Royal de 1|’Afrique
Centrale.

Cleora aculeata sp. n.
(Text-figs. 60-62 ; Pl. 6, figs. 243, 244, 247, 248 ; Map 3)

6. Vestiture white, sparsely irrorate with bister, patagia edged dorsally with bister. Wings
white ; antemedial fascia on fore wing, medial facia on hind wing and postmedial fascia on each
wing bister, slender and sharply marked ; apex, termen between veins M3 and Cz, and medial
area of fore wing and proximal two-thirds of hind wing sparsely irrorate, remainder of wings
more densely irrorate with bister or a tone paler (Pl. 6, fig. 243). Underside white, suffused
and patterned with fuscous (PI. 6, fig. 247).

50 DD. S. HE MeCH ER

6. Genitalia (Text-figs. 60, 61). Scobinate medial plate of gnathus one and one-third times
as broad as cucullus ; scobinate area on valve medio-ventrally ; arm of sacculus aculeate,
one-half as long as dorsal margin of valve, posterior edge slightly serrate and setose ; vesica with
two cornuti fused at base, one tapered and one-fourth as long as aedeagus, the other one-third
as long as aedeagus and tipped with four spines, and with a slender scobinate band equal in
length to shorter cornutus.

Q. (Pl. 6, figs. 244, 248). Similar to male externally.

@. Genitalia (Text-fig. 62). Lamella postvaginalis weakly sclerotized with concentric,
semicircular medial ridges ; lamella antevaginalis sclerotized laterally ; colliculum evenly
sclerotized ; bursa copulatrix cylindrical, anterior fourth membranous, remainder ribbed and
weakly sclerotized at right side.

Measurements. ¢38mm.; 940mm.

Fics. 60-62. C. aculeata genitalia. 60, aedeagus ; 61,3; 62, 9.

REVISION OF AFRICAN SPECIES OF CLEORA 51

Closely related to the preceding species, raphis ; externally well characterized
by the very clear white ground colour of the wings, especially prominent at apex
and mid-termen of fore wing and by the sharply marked bister pattern ; dis-
tinguished structurally by the shorter, slightly serrate process arising from the
sacculus and by the cornuti on the vesica in the male and by the degree of
sclerotization of the sterigma, colliculum and bursa copulatrix in the female.

Distribution (Map 3). Nigeria.

Holotype g. [NicEerIA] : Warri, vi.1897 (Dr. Roth), genitalia slide Geometridae
No. 2217.

Paratypes : data as holotype, 1 § ; Agberi, Niger, 10.v.1g01 (Ansorge), I 9.

Cleora panarista sp. n.
(Text-figs. 63-65 ; Pl. 6, figs. 249-252 ; Map 3)

6. Vestiture white, irrorate with smoke grey and black ; patagia banded with black ;
abdominal crest on first segment immaculate, second and third segments black dorsally. Wings
white, irrorate with smoke grey and black and, in some examples, suffused with cartridge buff
proximad of postmedial fasciae ; transverse fasciae and lateral streaks in discal area black,
as illustrated ; antemedial fascia toothed distad in submedial fold, medial fascia toothed
proximad on vein A, (Pl. 6, fig. 249). Underside white, patterned with fuscous black (Pl. 6,
fig. 250).

6. Genitalia (Text-figs. 64, 65). Minutely serrate medial plate of gnathus just subequal
to greatest width of cucullus ; very coarsely scobinate area on valve medio-ventrally ; arm of
sacculus smooth, tapered and arcuate, a little longer than greatest width of valve ; vesica with
three cornuti, one rod-like and one-half as long as aedeagus, one one-fifth as long as aedeagus
and tipped with one long and several shorter spines and one one-fourth as long as aedeagus,
arcuate and sharply tapered.

9. (PI. 6, figs. 251, 252). Similar to male externally.

®. Genitalia (Text-fig. 63). Lamella antevaginalis funnel-shaped ; lamella postvaginalis
sclerotized with concentric, semicircular ridges medially ; colliculum evenly sclerotized, rather
broader than long ; posterior two-thirds of bursa copulatrix sclerotized posteriorly and ribbed,
the ribbing extending weakly into the anterior third which is quite membranous.

Measurements. ¢ 40-45mm.; 9%“ 49 mm.

A species strikingly distinct in colour and pattern. Related closely to both
vaphis and aculeata ; distinguished structurally by the coarsely scobinate area on
the valve, the arcuate, tapered process on the sacculus and the ornamentation of the
vesica in the male and by the structure of the sterigma in the female.

The specimens from Malawi are browner than the other material examined ; the
smoke grey of the type is nearer cinnamon brown and the black pattern of the type
is bister. The difference may be geographical but is more probably due to the
considerably greater age of the specimens.

Distribution (Map 3). S.E. Congo (Leopoldville) ; Rhodesia ; Tanzania ;
Malawi.

Holotype g. S. RHopEsiA : Vumba Mts., ii.1956, genitalia slide Geometridae
No. 5224.

52

Fics.

63-65.

DS. PREELCHER:

C. panarista genitalia. 63,9; 64,3; 65, aedeagus.

REVISION OF AFRICAN SPECIES OF CLEORA 53

Paratypes : S. RHopEsIA : Vumba Mts., ii.1956, 1 g ; ibid., Umtali, i-ii.1960,
1 9. [ZAmBiA] N. Ruopesia : Mwinilunga, v.1g61, 1 2 in National Museum of
Rhodesia. [Conco (Leopoldville)] : Elisabethville, 29.11.1935 (Ch. Seydel), I 3;
ibid., xii.1935, I g in Musée Royal de |’Afrique Centrale. [MALAw1] NYASALAND :
Mt. Mlanje, 7.111.1913 (S. A. Neave), 192 ; ibid., 14.iv.1916, I g.

Cleora quadrimaculata (Janse) comb. n.
(Text-figs. 66-68 ; Pl. 7, figs. 253-257 ; Map 3)

Neocleova quadrimaculata Janse, 1932 : 269, pl. 8 : 13 ; fig. roo.

6. Vestiture tilleul buff to pinkish buff, irrorate with cinnamon buff to cinnamon ; patagia
and posterior dorsal margins of abdominal segments edged with fuscous black. Wings pinkish
buff, varyingly suffused with cinnamon and lightly irrorate with bister ; veins warm buff ;

67

68
Fics. 66-68. C. quadvimaculata genitalia. 66, 3 ; 67, aedeagus ; 68, 9.

54 D. S. FLETCHER

transverse fasciae slender and bister (Pl. 7, fig. 253). Underside of wings tilleul buff, weakly
suffused with pinkish buff and patterned with fuscous (PI. 7, fig. 254).

6. Genitalia very weakly sclerotized (Text-figs. 66, 67). Minutely scobinate medial plate
of gnathus semi-circular ; valve rhomboid, apical half of ventral margin minutely setose ;
cucullus very shortly spined ; arm of sacculus a broad-based, tapered process with a serrate-
edged, setose apex curved ventrad through 90° ; vesica with two stout, tapered cornuti fused at
base, each two-thirds as long as aedeagus.

®. Wings tilleul buff to pinkish buff, irrorate with bister, lightly proximad of postmedial
fasciae ; some warm buff irroration, especially on veins, proximad of antemedial fascia on fore
wing and posteriorly distad of postmedial fascia on each wing (PI. 7, figs. 255-257).

Q. Genitalia (Text-fig. 68). Lamella antevaginalis funnel-shaped, similar to that of pana-
vista ; lamella postvaginalis evenly sclerotized with small angular projection medio-posteriorly ;
colliculum shorter than broad ; bursa copulatrix cylindrical, anterior tip membranous, remainder
ribbed and sclerotized, more heavily anteriorly.

Measurements. (4 31-37mm.; 9 32-42 mm.

A small distinctively, cinnamon-coloured species displaying a range of variation
in pattern similar to that of transversaria. The genitalia, however, show no obvious
affinity with tvansversaria ; in the male the form of the sacculus is comparable with
that of vaphis, panarista and aculeata, but the valve lacks the area of coarse
scobination found in each of those species ; in the female the form of the sterigma
is similar to that of panarista.

The limited distribution of this Madagascan species along the south east coast of
Africa, combined with the apparent lack of subspeciation, suggests that it is probably
a recent arrival there.

Distribution (Map 3). S. Africa ; Madagascar.

Holotype g. S. ArricA: Pondoland, Port St. Johns, I~7.x.1931 (Janse), in
Transvaal Museum. Not seen.

Material examined. S. ArricA: Pondoland, Port St. Johns, 10—22.1.1955
(A. J. T. Janse), 13, 19, in Transvaal Museum ; Port St. Johns, 12—30.vi.1923
(R. E. Turner), 1 3 ; ibid., v.1924, 1 2 ; Natal, Umkomaas, 25.1.1913 (L. Hargreaves),
19. MapaGascar : Diego Suarez, 5.i1.1917 (G. Melou), 1 3 ; ibid., 10—30.iv.1917,
93,19; ibid., 15-20.vi.1917, 5 5, 3 2; ibid., 16—-29.viil.1917, 1g, 29; Sakaramy,
16.u.1917 (G. Melou), 1 9; Tananarive, 1 g; Tananarive (Stichel), 2 3;
Analalava, I 9.

Cleora boetschi (Herbulot) comb. n.
(Text-figs. 69-71 ; Pl. 7, figs. 258-261 ; Map 3)
Neocleova boetschit Herbulot, 1961 : 495, fig. 3.

6. Frons and thorax tilleul buff, irrorate with drab and bister ; patagia edged with bister.
Abdomen white to tilleul buff ; first segment white and immaculate, remaining segments with
a pair of bister spots medio-dorsally. Fore wing white, varyingly suffused with drab and bister ;
cubital and anal veins scaled with cinnamon buff ; ante- and postmedial fasciae bister. Hind
wing white, patterned as illustrated with bister ; a broad shade distad of postmedial fascia ;
medial and cubital veins scaled with cinnamon buff (Pl. 7, fig. 258). Underside of wings white,
densely suffused and patterned with fuscous (PI. 7, fig. 259).

REVISION OF AFRICAN SPECIES OF CLEORA

Ay)
a

Fics. 69-71. C. boetschi genitalia. 69,3; 70, aedeagus ; 71, 9.

55

56 D. S. FLETCHER

6. Genitalia (Text-figs. 69, 70). Gnathus one and one-half times as broad as greatest width
of cucullus, broadly incurved medio-ventrally and minutely scobinate ; a longitudinal, coarsely
scobinate band in distal half of valve ; arm of sacculus spatulate, of even width, scobinate at
tip and extending to mid-cucullus ; a setose digitate process medially at one-third valve ; vesica
with three cornuti, one stout, arcuate and tapered, one very slender and setose at apex, each
equal in length to greatest width of aedeagus, and one shorter cornutus with a curved, coarsely
serrate apex.

2. Wings white, lightly irrorate and patterned with bister (Pl. 7, fig. 260) ; hind wing with
weakly marked snuff brown shade distad of postmedial fascia ; both wings lightly irrorate with
warm buff, especially on medial and cubital veins. Underside of wings white, patterned with
fuscous (Pl. 7, fig. 261).

9. Genitalia (Text-fig. 71). Lamella postvaginalis slenderly produced, posteriorly narrowly
rounded ; lamella antevaginalis heavily sclerotized and bilobate posteriorly, anterior edge with
two folds sclerotized in a wrinkled and reticulate pattern ; colliculum rather longer than broad
with diamond-shaped sclerotized area at mid-anterior margin ; bursa copulatrix of almost
even width, posterior four-fifths sclerotized and ribbed, with a more heavily sclerotized disc
medio-anteriorly, anterior fifth membranous and slightly dilate.

Measurements. ¢ 44-52 mm.; 2 46-52 mm.

The species is characterized externally by its large size and in the male by the
densely suffused, dark underside of the wings. Structurally, the shape of the arm
of the sacculus and the ornamentation of the vesica in the male genitalia and the
structure of the sterigma in the female genitalia are diagnostic.

Though the sterigma in the female genitalia is considerably modified, the
scobination of the valve, the broad, scobinate gnathus and the long arm of the
sacculus in the male genitalia suggest an affinity with aculeata and with raplis.

Distribution (Map 3). Guinea ; Ghana ; Nigeria ; Cameroun ; Congo (Leopold-
ville) ; Uganda.

Material examined. Holotype g. CAMEROUN: N’long [Ngoulemakong], 18.1.
1953 (Rk. P. Boetsch), genitalia slide No. C.H.4028 in coll. Herbulot, Paris.

GUINEA : Soundedou, Nr. Macenta, 1600 ft., 13.v.1926 (C. L. Collenette), 2 3.
GHANA: Wassaw dist. 45 miles inland from Sekondi, 1 g. S. NIGERIA: Ilesha
(L. E. H. Humfrey), 23. CAMEROUN : Batouri, iii.1958, 1 J in National Museum of
S. Rhodesia ; Bitje, Ja River, x., wet season (G. L. Bates), 2g and Johann-Albrechts
Hohe, 1898 (L. Conradt), 1 g in British Museum (Natural History). CoNnco
(Leopoldville) : Coquilhatville, 9.1x.1955 (Dr. M. Fontaine), 1 g; Equateur,
Bokuma, 20.xi.1911 (Rev. P. Hulstaert), 1 g ; Eala, 14.1v.1936 (J. Ghesquiére), 1 3 ;
Uele, Paulis, 2-4.1v.1956 (Dr. M. Fontaine), 2 g ; ibid., 13.xi.1956, 1 9; ibid.,
5.vil.1958, 1 9; Uele-Itimbiri, La Kulu, 15.iv.1930 (J. Van den Branden), i 9 ;
Sankuru, Katako-Kombe, 25.iv.1952 (Dr. M. Fontaine), 1 3 ; ibid., 18.vi.1952, 14 ;
ibid., 6.1x.1952, I g¢ ; Lusambo, 17.vi-5.vii.1949 (Dr. M. Fontaine), 1 3; ibid.,
2.Vili—3.1X.1950, 3g, 3 9; Kasai, Luluabourg, 18.vi.1953 (Dr. M. Fontaine), i 3, all
in Musée Royal de l’Afrique Centrale ; W. Kivu, Upper Lowa Valley, Nr. Masisi,
5000-6000 ft., 11.1924, wet season (T. A. Barns), I § in British Museum (Natural
History).

UGANDA: Masaka, Katera, Sango Bay, x.1g60 (R. H. Carcasson), I 3, in
Coryndon Museum.

REVISION OF AFRICAN SPECIES OF CLEORA 57

Cleora bicornis sp. n.
(Text-figs. 72-74 ; Pl. 7, figs. 262-265 ; Map 9)

3d. Vestiture white to tilleul buff ; abdomen, except first segment, lightly irrorate with
snuff brown. Fore wing weakly suffused with snuff brown distad of postmedial fascia ; ante-
and postmedial fasciae snuff brown and very slender, the former preceded by and the latter
followed by a band of cinnamon brown irrorate with light buff ; discal spots white outlined with
snuff brown. Hind wing similarly marked, but antemedial fascia and band wanting ; in some
examples a snuff brown medial fascia extends from discal spot to anal margin (PI. 7, fig. 262).
Underside of wings white, patterned with snuff brown (PI. 7, fig. 263).

6. Genitalia (Text-figs. 72, 73). Medial plate of gnathus smoothly rounded ; sacculus with
two tapered processes, the tips of each curved through 90° ; a weakly sclerotized lip just
basad of cucullus ; margin of cucullus extended basad, parallel with dorsal margin of valve,
pustulate and setose ; aedeagus a little narrowed apicad ; vesica with one tapered cornutus
equal in length to width of aedeagus at middle.

74

Figs. 72-74. C. bicornis genitalia. 72,4; 73, aedeagus ; 74, 9.

58 D. S. FLETCHER

9. Similar to male, but underside of wings with broader and darker terminal bands, which
are fuscous (PI. 7, figs. 264, 265).

9. Genitalia (Text-fig. 74). Lamella antevaginalis consisting of two broad, tongue-like
plates overlapping slightly medially ; colliculum shorter than broad ; posterior half of bursa
copulatrix ribbed and very weakly sclerotized ; remainder membranous, globular anteriorly.

Measurements. ¢ 2 37-44 mm.

In external appearance closely similar to C. oculata, but differing in the clearly
defined ground colour proximad of the postmedial fascia. In the male genitalia the
two tapered processes on the sacculus and in the female genitalia the structure of
the sterigma are diagnostic.

The close similarity between oculata and bicornis in colour and pattern is not
reflected in either the male or the female genitalia of the two species. Structurally
bicornis appears to have no close relative and is for the present placed arbitrarily
with those species in which there are well developed processes on the sacculus in
the male genitalia.

Distribution (Map 9). Ivory Coast ; Nigeria ; Congo (Leopoldville).

Holotype 3. W. Arrica : Lagos, genitalia slide Geometridae No. 2281.

Paratypes : Ivory Coast: Bingerville, I-5.viii.1g15 (G. Melou), 1 g; ibid.,
5-7.Vili.I915, 1g. NIGERIA: data as type, 12 4, 3 9 ; Ilesha, (L. E. H. Humfrey),
I 9. [ConcGo (Leopoldville)] : Stanleyville, 9.vi.1948 (Drv. Faniel), 1 g in Musée
Royal de l'Afrique Centrale.

Cleora dargei (Herbulot) comb. n.
(Text-figs. 75-77 ; Pl. 7, figs. 266-269 ; Map 4)
Neocleova dargei Herbulot, 1961 : 493, fig. 2.

6. Vestiture : first abdominal segment white, remainder tilleul buff irrorate with bister ;
patagia edged with bister ; twin bister spots medio-dorsally on posterior edge of each abdominal
segment except first. Wings tilleul buff, suffused with drab and irrorate and patterned with
bister ; antemedial fascia on fore wing and postmedial fascia on each wing bister, the former
preceded, the latter followed by broad, snuff brown fascia ; veins warm buff distad of post-
medial fasciae, cubital and anal veins interruptedly warm buff to base (Pl. 7, fig. 266). Under-
side white, suffused and patterned with bister (Pl. 7, fig. 267).

6. Genitalia (Text-figs. 75, 76). Scobinate medial plate of gnathus narrowly rounded
medially ; juxta sclerotized in a Y-shaped pattern ; sacculus extending to one-half ventral
margin of valve with a short, tapered, apical projection and a setose, medial ridge ; when
closed valves viewed from ventral position, the marginal projections of the sacculus appear
triangular in form ; apex of aedeagus evenly rounded ; vesica with two elaborately spined
cornuti fused at base, one about one-half as long, the other one-third as long as aedeagus ;
the spining varies somewhat individually, but the longer cornutus is usually truncate apically ;
the shorter one is dilate at base and has the apex tapered and curved through 45° and bearing
usually 2—5 spines.

9. Vestiture similar to that of male, but ground colour white. Wings white, patterned
similarly to male, but drab suffusion and bister irroration very light (PI. 7, fig. 268). Underside
white, suffused and patterned with bister (Pl. 7, fig. 260).

9. Genitalia (Text-fig. 77). Lamella postvaginalis with posteriorly rounded medial plate ;

REVISION OF AFRICAN SPECIES OF CLEORA 59

colliculum as broad as long ; anterior half of bursa copulatrix membranous, posterior half
ribbed and sclerotized with a slight shoulder-like projection at one side posteriorly.

Measurements. 4 34-44mm.; @ 38-45 mm.

A long series of both sexes displays little variation in either colour or pattern and
no tangible external diagnostic character. In the male genitalia the shape of the
sacculus and of the cornuti and in the female genitalia the form and structure of the
bursa copulatrix are diagnostic.

Fies. 75-77. C. dargei genitalia. 75,35; 76, aedeagus ; 77, 9.

60 DS. BVP CHER

Biology. Moths have been reared from larvae found feeding on Eucalyptus
camaldulensis Dehnh. in Nigeria and from Entandrophragma angolense de Candolle,
Pinus patula Schiede and Cupressus sp. in Uganda.

Distribution (Map 4). Sierra Leone; Ivory Coast; Ghana; Nigeria ;
Cameroun ; Gabon ; Angola ; Congo (Brazzaville) ; Congo (Leopoldville).

Material examined. Holotype g. CAMEROUN: N’long [Ngoulemakong], 18.1.
1953 (Rk. P. Boetsch), genitalia slide No. C.H.4027 in coll. Herbulot, Paris.

SIERRA LEONE, I g, 19; ibid. (A. B. Frere), 2 3,22; ibid. (Major Bainbridge),
I g. Ivory Coast, 1 g; Bafing river, S of Touba, 1200 ft., 4.vii.1926 (C. L.
Collenette), 1 3 ; Bingerville, xi.1913 (Gaston Melou), 2 3, 29; ibid., vi, vili-x.1915,
9d, 11 9. GHANA: N. Territories, Kete-Krachi (A. W. Cardinall), 3 9 ; Kumasi,
Odumase Swamp, 1913 (Smeed), 195 ; Kumasi, ii-ili.1923 (N. E. Bell), 1 3 ; Kumasi,
Spurrell), 1 8 ; Ashanti, Goaso (G. N. Gibbs), 1 g. NiGErRIA: Ilesha (L. E. H.
Humfrey), 23 ; Lagos, 1g, 6 9 ; Warri, 1 9. CAMEROUN: Lolodorf, 1894-1895
(L. Conradt), 1 § ; Johann-Albrechts Hohe, 1898 (L. Conradt), 1 9. GABON :
Tchibanga (P. Rougeot), I g. CoNGo (Brazzaville) ; Brazzaville, viii.1948 (P.
Rougeot), I g. Conco (Leopoldville) : Bopoto (Kenred Smith), 3 9 ; Yakusu (K.
Smith), 1 g, all in British Museum (Natural History) ; Leopoldville, 11.xi.1953
(Dr. M. Fontaine), 1 3 ; Leopoldville, Binza, 11.viii.1953 (Dr. M. Fontaine), 1 3 ;
Ubangi, Gemena, 1.1936 (J. Ghesquiére), 1 9 ; Equateur, Flandria, 24.11i-20.iv.1932
(R. P. Hulstaert), 5 3 ; Equateur, Bokota, 1927 (R. P. Hulstaert), 1 g ; Bokela,
17.1v.1940 (R. P. Hulstaert), 1g; Eala, 20.x.1917 (R. Mayné), 19; Eala, xi.1936
(J. Ghesquiére), 1 3 ; Ifuta, 21.x.1g21 (Verlaine), 1 9 ; Luebo, x.1930 (J. P. Colin),
Ig ; Stanleyville, vi.1929 (J. Colin), 1g ; allin Musée Royal de l'Afrique Centrale ;
Upper Uele Dist., Dungu, v, I 9, in British Museum (Natural History) ; Uele,
Rungu, 30.iv.1960 (Dr. M. Fontaine), 1 3; Uele, Paulis, i,iv,ix.1956 (Dr. M.
Fontaine), 1 3, 2 9; Sankuru, Lusambo, 25-29.v1.1949 (Dr. M. Fontaine), 2 3 ;
ibid., 14—20.x.1949, 2g ; ibid., iv,viiix.1950, 5g, 4 9, allin Musée Royal de l’Afrique
Centrale ; Kasai, Lusambo, ix—x.1919, 4 ¢ in British Museum (Natural History) ;
Sankuru, Djeka, 17.xii.1952 (Dr. M. Fontaine), 1 § ; Sankuru, Katako-Kombe,
ili, Viil,1X—xil.1951-53 (Dr. M. Fontaine), 9 3, 4 9; Kivu, Rwankwi, 9.vili.1947
(J. V. Leroy), 1g; ibid., 6-7.x.1947, 1 § ; Rutschuru, 1.1928 (Ch. Seydel), 1 g, all
in Musée Royal de l'Afrique Centrale ; Lindi River, Bafwasende, 2000 ft., vii.1g21
(I. A. Barns), I $, in British Museum (Natural History). UGanpa : Bundibugyo,
3440 ft., 22.vili-3.1x.1952 (D. S. Fletcher), 1 2 ; Mpanga Forest, [Kampala], 21.iv.
1961, ex Entandrophragma angolense (W. K. Brown), 1 2 ; Mpanga Forest, 21.x.1960,
ex Cupressus sp. (W. K. Brown), 1 9 ; Kaweri, 21.xii.1963, ex Pinus patula (W. K.
Brown), I 3.

Cleora dactylata sp. n.
(Text-figs. 78-80 ; Pl. 7, figs. 270-273 ; Map 4)
36 2. Externally closely similar in both sexes to davgez and appears to be separable only by

reference to the genitalia. In one example the medial area of the hind wing and the posterior
half of the medial area of the fore wing are densely irrorate with bister.

REVISION OF AFRICAN SPECIES OF CLEORA 61

6. Genitalia (Text-figs. 78, 79). Gnathus and juxta similar to those of dargei ; sacculus
extending to one-half ventral margin of valve with inner margin setose and with a short, minutely
scobinate, digitate projection from apex ; apical fourth of aedeagus irregularly shaped (Text-
fig. 79) ; vesica with two cornuti fused at base, the apical one comb-like at one side and about
one-third as long as aedeagus, the other one-sixth as long as aedeagus, tapered and curved through
45° from a bulbous base.

9. Genitalia (Text-fig. 80). Lamella antevaginalis shallow and broad ; lamella post-
vaginalis with a triangular, sclerotized pattern medially ; colliculum apparently not developed ;
slightly more than anterior half of ductus bursae membranous, remainder sclerotized and ribbed
with dilate, membranous additional sac at left side posteriorly.

Measurements. 4 31-42mm.; 9 43-46 mm.

Closely related to darget with which it occurs, examples of each species having
been taken by Dr. Fontaine at Katako-Kombe on consecutive days. Differs

80

Fics. 78-80. C. dactylata genitalia. 78,3; 79, aedeagus ; 80, 9.

62 D. Ss. PLETCHER

structurally from dargei in the male genitalia in the slender slightly scobinate,
digitate extension of the sacculus, in the irregular shape of the apex of the aedeagus
and in the shape of the cornuti on the vesica. In the female genitalia the form of
the sterigma, the absence of a colliculum and the development of the dilate, mem-
branous additional sac are diagnostic.

Distribution (Map 4). Congo (Leopoldville) ; W. Uganda.

Holotype g. [Conco (Leopoldville)] ; W. Kivu, Nr. Masisi, Upper Lowa Valley,
5000-6000 ft., forest and long grass, 11.1924, wet season (T. A. Barns), genitalia
slide Geometridae No. 2297.

Paratypes. [Conco (Leopoldville)] : data as holotype, 1g ; Nr. Walikili, 3000-
4000 ft., forest, 11.1924, wet season (T. A. Barns), 2 § all in British Museum (Natural
History) ; Sankuru, Katako-Kombe, 27.xu1.1951 (Dr. M. Fontaine), 1 3 ; ibid.,
23-24.V11.1952, 2 9 ; ibid., 10.ix.1952, Ig ; ibid., 15.xi.1952, 1 9 ; ibid., 11.1.1953,
Ig; Lusambo, 24.vii.1949 (Dr. M. Fontaine), 1 § ; Luluabourg, 20.viii.1955 (Ch.
Seydel), 19; Uele, Paulis, 18.11.1956 (Dr. M. Fontaine), 1 g ; ibid., 12.iv.1956, 2 dg,
IQ; ibid., 3.v.1956, 1g ; Boyenga, 1930 (Delpiérre), 1 § ; Fomlioko, 20.11.1921
(Verlaine), 1 § ; Boende, 1.iv.1940 (R. P. Hulstaert), 1 g, all in Musée Royal de
l'Afrique Centrale. UGANDA : Bwamba, v.1956 (R. Carcasson), I 3, I &.

Cleora thyris sp. n.
(Text-figs. 81-83 ; Pl. 8, figs. 274-280 ; Map 4)

3g. First abdominal segment white, remainder of vestiture white, irrorate with bister. Wings
white, irrorate and patterned with bister ; terminal interneural spots, medial and postmedial
fasciae on each wing, discal streak and antemedial fascia on fore wing sharply marked and a
shade darker ; cubital veins and other veins distad of postmedial fasciae streaked with warm
buff (Pl. 8, fig. 274). Underside tilleul buff, suffused and patterned with bister (Pl. 8, fig. 275).

6. Genitalia (Text-figs. 81, 82). Gnathus with semi-circular scobinate medial plate, equal
in diameter to two-fifths width of cucullus, the scobination extending posteriorly along sub-
scaphium ; process from sacculus spatulate and incurved, outer surface of apex and inner surface
of base pustulate and setose ; vesica with two cornuti, one of even width, serrate at one side and
scobinate on ventral and dorsal surfaces apically, five-eighths as long as aedeagus, the other
tapered sharply, scobinate at one side apically and one-fourth as long as aedeagus.

9. Vestiture white, very lightly irrorate with bister, abdominal segments, except first, with a
pair of bister spots medio-posteriorly. Wings white, patterned as in male, but irroration very
light and sparse ; buff colouring on veins paler and weakly marked. (PI. 8, figs. 276, 277).

®. Genitalia (Text-fig. 83). Medial area of lamella postvaginalis heavily sclerotized in horse-
shoe pattern ; colliculum one and one-half times as long as broad ; ostium bursae broad and
shallow, anterior margin sharply defined ; anterior fourth of bursa copulatrix membranous,
remainder sclerotized and ribbed.

Measurements. ¢38-50mm.; 2 40-50 mm.

The species is variable in colour and pattern, displaying a range of poly-
morphism similar to that of twlbaghata and nigrisparsalis ; in examples of both sexes
the area proximad of the antemedial fascia on the fore wing is ochraceous buff and
distad of the postmedial fascia on each wing there is a broad band of the same colour,
comparable with tulbaghata ab. flavipleta ; in another example the ochraceous buff

REVISION OF AFRICAN SPECIES OF CLEORA 63

is replaced by bister (Pl. 8, fig. 278). There are further examples in which the
medial area of each wing is suffused with fuscous black comparable with tulbaghata
ab. fumata (Pl. 8, fig, 279) ; discal spots may be heavily ringed with or be entirely
fuscous black (Pl. 8, fig. 280). The genitalia afford the most certain means of
identifying ‘hyvis ; in the male the process on the sacculus and in the female the
broad, shallow ostium bursae together with the form and proportions of the mem-
branous and sclerotized parts of the bursa copulatrix are diagnostic.

Distribution (Map 4). Ethiopia; Kenya; N.W. Congo (Leopoldville) ;
Tanzania ; Malawi.

Holotype g. B.E.A. [Kenya] : Kyambu, 18.vii.1g19 (W. Feather).

Fics. 81-83. C. thyvis genitalia. 81, aedeagus ; 82,4; 83, 9.

64 D. S. FLETCHER

Paratypes ; ETHIOPIA: Negelli, v.1962 (S. Chojnacki), 2g. Kenya: N.F.D.,
Marsabit, 4500 ft., 11.1956 (J. G. Williams), I 3, 3 2; ibid., vi.1955, I g in Coryndon
Museum ; Mt. Marsabit, 4500 ft., 1.1946 (T. H. E. Jackson), 13 ; Mt. Elgon, vii.1951
(T. H. E. Jackson), 1 ; Kitale, xi.1953 (C. R. Howard), 1 § ; Nakuru, 14.iv.1952
(A. L. H. Townsend), 1 9; Nyeri, vii.t950 (A. L. H. Townsend), 1 9; Nyeri
(H. B. Kettlewell), 2 9 ; data as holotype, 12 J, 12 9; Nairobi, April-June (van
Someren), 2 3, ibid., 27.vi.1g18, I g, all in British Museum (Natural History) ;
Nairobi, Karura Forest, 111.1949 (EF. Pinhey), 1 2 ; Nairobi, iv—vi. (van Someren), 2 3 ;
ibid., 27.vi.1918, 1g ; ibid., x.1923, 12 ; Nairobi, Thika [Road], v.1950 (E. Pinhey),
I g, all in Coryndon Museum ; Meru District, Mt. Kenya, ix.1930 (Mrs. H. Young),
Ig; Mt. Meru, Kenya distr., 5600 ft., xii.1g20 (W. N. van Someren), 1 9, all in
British Museum (Natural History). [Conco (Leopoldville)] : Kibali-Ituri, Mt.
Rowa, 5.xii.1952 (J. Hecq), I J in Musée Royal de l'Afrique Centrale. [TANZANIA]
Ig; Amani, 28.xi.1934 (Mrs. Editha Dalton), i 3, both in British Museum (Natural
History) ; Makoa, 7-27.1.1959 (Lindner), 25,19; ibid., 6-25.i1.1959, 2d, 2 Qin
Zoological Museum, Stuttgart. [MALAwl] NyASALAND: Mlanje, 30.iv.1913, I 3,
in British Museum (Natural History).

Cleora nigrisparsalis (Janse) comb. n.
(Text-figs. 84-86 ; Pl. 8, figs. 281-286 ; Map 4)

Neocleora nigrisparsalis Janse, 1932 : 270, pl. 8: 5 ; fig. roo.

g. Vestiture white, irrorate with fuscous black, lightly on first abdominal segment. Wings
white, densely irrorate and patterned with fuscous black (Pl. 8, fig. 281). Underside of wings
white, patterned with fuscous black (PI. 8, fig. 282).

6. Genitalia (Text-figs. 84, 85). Scobinate medial plate of gnathus semicircular, equal in
diameter to two-fifths width of cucullus ; a small spatulate, digitate process from apex of
sacculus ; vesica with two cornuti fused at base, one of even width, apex serrate at one side,
dorsal and ventral surfaces scobinate apically, one-half as long as aedeagus, the other tapered
sharply apicad and one-seventh as long as aedeagus.

2. Similar in pattern to male ; upperside of wings less heavily, but more evenly irrorate
with fuscous black (Pl. 8, figs. 283, 284).

2. Genitalia (Text-fig. 86). Anterior margin of lamella postvaginalis irregularly shaped ;
colliculum one and one-quarter times as long as greatest width, slightly broadened posteriorly,
posterior margin sharply defined ; anterior three-sevenths of bursa copulatrix membranous,
remainder sclerotized and ribbed with a shoulder-like projection at one side of posterior margin.

Measurements. ¢g40-44mm.; Q 41-52 mm.

The range of variation in colour and pattern in migrisparsalis is similar to that
found in both thyvis and tulbaghata ; the similar forms in each species are virtually
indistinguishable except by genitalia. Typical forms of migrisparsalis (Pl. 8, figs.
281-284) are recognisable externally by the dense fuscous black irroration of the
upperside of the wings and by the sharply contrasted and well-defined fuscous black
and white pattern of the underside. Structurally the small digitate process on the
sacculus and the shape and proportions of the cornuti on the vesica in the male
genitalia and the shape of the sterigma, the sharply defined posterior margin of the

REVISION OF AFRICAN SPECIES OF CLEORA 65

colliculum and the shape and proportions of membranous and sclerotized parts of
the bursa copulatrix are diagnostic.

Biology. Larvae have been found damaging the foliage of coffee (C. arabica) at
Ruiru in Kenya.

Fics. 84-86. C. nigvisparsalis genitalia. 84, 3; 85, aedeagus ; 86, 9.

66 D. S. FLETCHER

Distribution (Map 4). Natal; Transvaal ; Mozambique ; Malawi ; Zambia ;
Rhodesia ; Tanzania ; Kenya ; Burundi ; Congo (Leopoldville) ; Angola.

Holotype g. S. Ruopesia, Umtali, in Transvaal Museum, Pretoria ; not
examined.

Material examined. NatTaL: Pietermaritzburg, Malta, i-ii.1927-8 (G. van Son),
8 g, 2 2 in British Museum (Natural History), 6 g in Transvaal Museum ; Natal
National Park, Mont-aux-Sources, 30.1.1954, Ig ; Muden, 15 i1.1955 (H. Cookson),
I 9 in Transvaal Museum. TRANSVAAL : White River, 11.1909 (A. T. Cooke), 1 g ;
Barberton (Harrison), I$ ; Barberton (A. A. Williams), 1.1933, 1 3 ; Marieps Mtn.,
6-14.1 (G. van Son), 3 3, I 9, in British Museum (Natural History), 2 g in Transvaal
Museum ; Pretoria, i.1913 (Gladstone), I § ; Tshakoma, Zoutpansberg, xi.1931
(G. van Son), 1 § ; Magoebaskloof, iv.1933 (Rev. D. P. Murray), 1 29; Erasmus
Reserve, Pilgrim’s Rest District, 30.1v.1960 (f°. Neubecker), 3 3 ; Kowyn’s Pass,
Pilgrim’s Rest District, 22.11.1962 (Vari & Leleup), 1 9, all in Transvaal Museum.
MoZAMBIQUE : Dondo, ix.1954 (H. Cookson), 1 9. Matawi: Zomba, 3000 ft.,
iv.1913 (E. Ballard), 1 § ; Zomba, 1.1921 (H. Barlow), 2 3, 1 2; ibid., v.1g20, I Q,
all in British Museum (Natural History). Ruoprsia: Nr. Bulawayo, Khami,
111.1956, 2 ¢ ; Marandellas, ii,x,xi1.1960, 3 J, 1 2; Vumba Mts., Umtali, iii,xi,xu,
446; Umtali District, inn, 2 9, all in National Museum of Rhodesia ; Umtali,
i-ili,vi, I 9, in Transvaal Museum, 2 § in British Museum (Natural History).
Salisbury, iii,iv,v,vii, 5 g in National Museum of Rhodesia, 2 3 in British Museum
(Natural History) ; Inyanga, iv.1961, 1 g in National Museum of Rhodesia ;
Penalonga (O. A. Kidwell), 1 3 ; Mt. Selinda, 17—31.1.1959 (G. van Son), 1 f ; Mt.
Selinda, 8-9.iii.1954 (H. Cookson), 1 9, all in Transvaal Museum ; Victoria Falls,
10.iv.1927 (R. H. R. Stevenson), 1 3 ; Shamva, ii.1g21 (O’New), 1g ; Mountain Inn,
Melsetter, xi.1950 (H. B. Kettlewell), 1 3,19. ZAMBIA: Kashitu, 14.v.1915 (H. C.
Dollman), 1 3 ; N’dola, 4.1.1923, I g, all in British Museum (Natural History) ;
TANZANIA : Makoa, 7—-27.1.1959 (Lindner), 1 2 in Zoological Museum, Stuttgart ;
Mamboia (Dr. Baxter), I Qin British Museum (Natural History). Kenya : Nairobi,
v.1927 (D. M. Hopkins), 1 9 ; ibid., 1.1953 (E. Pinhey), 1 9 in Coryndon Museum ;
ibid., 8.ix.1927 (Mrs. D. M. Hopkins), 1 3, 1 2; ibid., x.1920 (W. N. van Someren),
IQ; ibid., 1.1928, 1 2 ; ibid., 8.iv.19g12 (C. Montague Smyth), 1 3 ; ibid., bred 1937
(R. H. Simmonds), 1 3 ; ibid., 5500 ft., 111.1939 (MaclInnes), 1 9 ; Nairobi Plains,
Kikuyu, 5.v.1g00 (R. Crawshay), 1 § ; Makueni, Machakos [District], vi.1958, 1 g
in Coryndon Museum ; Ruiru, vi.1963 (P. E. Wheatly), 3 3, 3 29; Mt. Kenya,
vii.1930 (E. Barns), 5 9; Ndarugu, 18-19.vi.1917 (W. Feather), 2 3 ; Suna, S.
Kavirondo, 1.1932 (W. Feather), 4 3, I 9, all in British Museum (Natural History).
Burunpi : Kitega, i,vi-x.1962 (Dr. M. Fontaine), 35 3, 17 9. Conco (Leopold-
ville) : Kibali-Ituri, Nioka, vi,vii,x,xi (J. Hecq), 2 3, 4 9 all in Musée Royal de
) Afrique Centrale ;. Elisabethville, iciv, x-xu (Ch. Seydel), 9 3 in Musée Royal de
l Afrique Centrale, 7 g, 2 9 in British Museum (Natural History). ANGoLa: Bihé,
X1i.1934-1.1935 (E. Braun), 2 3.

REVISION OF AFRICAN SPECIES OF CLEORA 67

Cleora tulbaghata (Felder) comb. n.
(Text-figs. 87-89 ; Pl. 9, figs. 289-296, 301 ; Map 4)

Boarmia tulbaghata Felder, 1875, pl. 125: 5.

Chogada acaciavia sensu Warren, 1904 : 474.

Chogada acaciaria ab. flavipleta Warren, 1904 : 474.
Chogada acaciavia ab. fumata Warren, 1904 : 474.
Neocleova munda sensu Janse, 1932 : 267, text-fig. 100.

6. Genitalia (Text-figs. 87, 88). Medial plate of gnathus narrowly rounded at tip, which is
one-third as broad as cucullus ; sacculus moderately sclerotized with a short, membranous,
triangular apical projection at about two-thirds ventral margin of valve ; vesica with two
cornuti fused at base, one one-half as long as aedeagus, the apical fourth dilate, with one side
coarsely serrate and with ventral surface scobinate, the other one-fifth as long as aedeagus and
sharply tapered apicad.

2. Genitalia (Text-fig. 89). Sclerotization of lamella postvaginalis extends posteriorly very
slenderly at each side of short medial plate ; colliculum one and one-third times as long as
broad, without sharply defined posterior margin ; anterior two-sevenths of bursa copulatrix
membranous, remainder sclerotized and ribbed with a shoulder-like projection at one side of
posterior margin.

Measurements. (4 33-49mm.; 2 35-49 mm.

A species variable in colour and pattern. In the commonest form the ground
colour of the male is white, irrorate with drab and fuscous ; the transverse fasciae
are fuscous black and the veins are interruptedly light buff distad of the postmedial
fasciae (Pl. 9, fig. 289) ; the underside of the wings is white, patterned with fuscous
(Pl. 9, fig. 290). In this form the female is similarly patterned, but the irroration,
which is sparse, and the transverse fasciae are bister (Pl. 9, fig. 291).

The holotype (Pl. 9, fig. 296) is white, very lightly suffused with ochraceous and
patterned with fuscous. In marking it remains unique, but is closely approached
by a recurrent form (PI. g. fig. 295) in which the antemedial fascia of the fore wing
and the postmedial fascia on each wing are broadly fuscous, the former edged
proximally, the latter edged distally with snuff brown.

Another recurrent form, ab. flavipleta Warren (Pl. 9, fig. 293), has the area
proximad of the antemedial fascia on the fore wing and broad bands distad of the
postmedial fascia on each wing ochraceous buff to ochraceous orange.

In ab. fumata Warren (Pl. 9, fig. 294) the medial area of the fore wing and the
area proximad of the postmedial fascia on the hind wing are densely irrorate with
fuscous.

A completely black form (Pl. 9, fig. 301) occurs in part of the Fish Hoek Valley in
Cape Province. It is referred to by Dr. H. B. D. Kettlewell (1957 : 9, pl. 5), who
found it resting on the fire-blackened trunks of a species of Acacia. Ina subsequent
personal communication Dr. Kettlewell writes : “ It is one of the few examples I
know of a southern hemisphere Lepidopteron having a clear-cut melanic form, nor
do I know of it anywhere else except in the very local area of the Fish Hoek Valley.
In this locality, the species seems to depend on an introduced Australian shrub,
Rooikrans [Acacia cyclops A. Cunn.]—a sort of fire-resistant myrtle. The Valley is
regularly burned out by the coloureds who lived there and the tree trunks were
completely blackened everywhere. The insects sat on these trunks, where the

FLETCHER

S.

De

68

87, aedeagus ; 88,35; 89, 9.

C. tulbaghata genitalia.

Fics. 87-89.

REVISION OF AFRICAN SPECIES OF CLEORA 69

melanic form appeared to me to have considerable cryptic advantage. I would say
that the melanic form was between 4 and Io per cent. of the population over the
the period I collected .”

The species is closely related to thyris, nigrisparsalis and to the following species,
munda ; in the male genitalia the shapes and sizes of the medial plate of the gnathus
and of the process on the sacculus and the shape of the broadly spatulate cornutus
on the vesica are diagnostic ; in the female genitalia the shape of the sclerotized
parts of the sterigma, the colliculum and the bursa copulatrix are diagnostic.

Distribution (Map 4). Kenya ; Zambia ; Mozambique ; Transvaal; Natal ;
Cape Province.

Material examined. Holotype g¢ (without abdomen). Knysna, C. W., ex coll.
Felder.

KeENnyA : Suna, S. Kavirondo, iv.1930 (W. Feather), 1 3; ibid., 1.1931, I gf;
Kibwezi, 11.1.1917 (W. Feather), 1 9. ZAMBIA: (Gimson), 23,19. MOZAMBIQUE :
Lorenzo Marques (Distant Coll.), 1g ; Lorenzo Marques, 3.vii.1949 (H. B. Kettlewell),
Ig; Magude, x.1918 (C. J. Swierstra), 1 9; Delagoa Bay (Monteiro), 2 9.
TRANSVAAL : Barberton (Harrison), 2 2 ; Three Sisters, 28.ii.1g11 (A. J. T. Janse),
I 9. ZuLULAND: (G. F. Leigh), r g, 1 2 in British Museum (Natural History) ;
Eshowe, 1.1956 (D. A. Swanepoel), 1 9 in Transvaal Museum. Nata: Durban,
Umhlanga, 13-20.vii.1950 (H. B. D. Kettlewell), 2 § ; Umhlanga Rocks, 6.i1.1955
(C. G. C. Dickson), t 2 in Transvaal Museum ; Durban (G. F. Leigh, Col. Bowker,
Stella Bush, C. G. C. Dickson), 26 3, 18 9 in British Museum (Natural History), 3 3,
I Qin Transvaal Museum ; Port Natal (Guenzius), 3 3,22 ; Pinetown (G. F. Leigh),
26,1292; Krantzkloof, 18.1.1918 (L. Hargreaves), I § ; Umkomaas, 25.xii.1g1o (L.
Hargreaves), i § ; ibid., 8.ii.1913, I g ; Port Shepstone, 12.i.1924 (Lt. Col. G. R.
Oakes), I g. CAPE PROVINCE: E. Griqualand (Kendall), 1 g, 1 9; E. Cape,
Kamanassie, ix.1949 (H. Bridges), 1 § in Transvaal Museum ; Pondoland, Port St.
Johns, i-v.1924 (R. E. Turner), 5 3, 7 @ ; ibid., 23-26.viii.1950 (H. B. D. Kettlewell),
24,19; ibid., 10-22.1.1955 (A. J. Janse), 3 3, 1 9 in Transvaal Museum ; W.
Pondoland, Nggeleni, 17.11.1904 (H. H. Swinny), 1 g ; East London, 19.11.1913
(G. F. Leigh), 23,19 ; Grahamstown, 1 9; Port Elizabeth, ii,iii,iv (C. G. C. Dickson)
I g, 2 2 in Transvaal Museum ; Groot River Pass, 22—23.1.1955 (A. J. T. Janse),
I g in Transvaal Museum ; Knysna, C.W. (ex coll. Felder), 3 J, 1 2; Wilderness,
Knysna Forest, 20.x—-8.xi.1952, at M.V. light (H. B. D. Kettlewell), 23 3, 12 9;
ibid., 6—20.11.1934, I gd, 4 9; Knysna, 10-20.1.1958 (A. J. T. Janse), 1 3, 2 9 and
24.v1.1950 (C. G. C. Dickson), I 9 in Transvaal Museum ; Belvedere, 10.i1-3.iv.19g21
(Dr. H. G. Brewer), 1 3,29 ; Harkerville, 9.iii.1g21 (Dr. H. G. Brewer), 1 3 ; Mossel
Bay, 15.ili-20.iv.1932 (R. E. Turner), 1 3 ; Cape Town, ix.1877, 1 ¢ ; ibid., 1.1871,
Ig; ibid., v.1g12 (Lord Gladstone), 1 3 ; ibid., 14—21.1.1961 (C. G. C. Dickson), I 3,
I 9 in Transvaal Museum ; Cape Town, Camps Bay, i,li,viii,ix,xi,xii (A. J. Duke),
54, I Qin Transvaal Museum ; Cape Town, Seapoint, 13.x.1953 (A. J. Duke), 1 J in
Transvaal Museum ; Mowbray, ii.1950 (A. J. Duke), 1 g in Coryndon Museum ;
Cape Town, Kirstenbosch, 5—29.x11.1954 (A. T. J. Janse), 2 3, 1 2 and Clovelly,
li,iv,xil (D. A. Swanepoel), 3 3, I 9 in Transvaal Museum ; Simonstown, (H. G.

7oO

Fics. 90-92.

DoS. PREDCHER

92

C. plax genitalia. 90, $ ; 91, aedeagus ; 92, 9.

REVISION OF AFRICAN SPECIES OF CLEORA 71

Hunt), 3 9 ; Fish Hoek, x—xi.1949 (H. B. D. Kettlewell), 2 3 ; ibid., 1949-1950, 6 3,
IQ; ibid., 28.ii1.1950, 23 fg, 5 2; ibid., 15.iv.1950, I gf ; ibid., 1-iii.1950, at light,
225,792; Fish Hoek Valley, 10.xi.1952-3.il1.1953, M.V. light (H. B. D. Kettlewell),
236,92; Fish Hoek-Kommetje, 1951 (H. B. D. Kettlewell), 1 3.

Cleora plax sp. n.
(Text-figs. go-92 ; Pl. 8, figs. 287, 288 ; Map 4)

6. Vestiture white, irrorate with drab and bister ; patagia edged with fuscous. Wings
white, lightly irrorate with bister ; broad snuff brown fascia proximad of antemedial fascia on
fore wing ; similar fascia distad of postmedial fascia on each wing ; cubital veins and remaining
veins distad of postmedial fascia on each wing interruptedly warm buff ; other markings fuscous
in holotype, (a very fresh specimen), bister in remainder of series (Pl. 8, fig. 287). Underside of
wings white, suffused with fuscous (Pl. 8, fig. 288).

6. Genitalia (Text-figs. 90, 91). Scobinate medial plate of gnathus shallow and broad,
equal in width to cucullus ; apex of sacculus truncate with short, digitate process inclined
towards mid-valve ; apex of aedeagus narrowly rounded and inclined to one side ; vesica with
two cornuti fused at base, one slightly sinuous with apical fourth dilate and spatulate and with
one side coarsely serrate and ventral surface scobinate, the whole about one-half as long as
aedeagus, the other tapered sharply apicad, one-sixth as long as aedeagus.

2. Upperside of wings patterned similarly to that of male, but irroration, especially in basal
area of fore wing and medial area of each wing, greatly reduced. Fuscous pattern of underside
restricted to discal spots and terminal bands, narrow on hind wing.

®. Genitalia (Text-fig. 92). Anterior margin of lamella postvaginalis sinuous and produced
medially ; colliculum slightly longer than broad, posterior margin sharply defined ; anterior
third of bursa copulatrix membranous, remainder sclerotized and ribbed with a shoulder-like
projection at one side of posterior margin.

Measurements. ¢ 40-41 mm. ; 2 46 mm.

Similar externally to forms of thyris, tulbaghata and munda. Male genitalia
similar to those of nigrisparsalis and tulbaghata in structure of sacculus and similar
to tulbaghata in shape of cornuti ; the broad, shallow scobinate medial plate of the
gnathus is distinctive. In the female genitalia the posterior margin of the lamella
postvaginalis is diagnostic.

Distribution (Map 4). N.E. Congo (Leopoldville) ; W. Uganda.

Holotype g. UGanpa: Masaka, Katera, Sango Bay, x.1960 (R. H. Carcasson),
genitalia slide Geometridae No. 5289.

Paratypes : [ConGco (Leopoldville)] : Escarpment, west of Semliki Valley, 20
mls. south-west of Boga, 3500—4000 ft., vii.1924, borders of tropical forest and long
grass country (7. A. Barns), genitalia slide Geometridae No. 2299, 1 g in British
Museum (Natural History) ; Mongbwalu (Kilo), 1937, (Me. Harford-Jordens), 1 2 ;
Kilo, 1.1940 (RR. FF. Maristes), 2 3 ; Mambunga via Boma (R. P. Bittremieux),
I 9, all in Musée Royal de |’Afrique Centrale.

Cleora munda (Warren) comb. n.
(Text-figs. 93-95 ; Pl. 9, figs. 297-300, 302 ; Map 4)
Chogada munda Warren, 1899 : 52.
g. Genitalia (Text-figs. 93, 94). Medial plate of gnathus narrowly rounded, as in tulbaghata ;

72 D. S. FLETCHER

sacculus moderately sclerotized but without process ; vesica with two cornuti fused at base,
one five-sixths as long as aedeagus, of even width and with apical sixth coarsely serrate at one
side, the other one-half as long as aedeagus and tapered apicad.

9. Genitalia (Text-fig. 95). Anterior margin of lamella postvaginalis irregularly shaped ;
short colliculum as broad as long ; posterior margin produced and only on this part is the
margin sharply defined ; anterior sixth of bursa copulatrix membranous, remainder ribbed and
sclerotized.

Measurements. 3 36-45 mm. ; 2 42-50 mm.

95
Fics. 93-95. C.munda genitalia. 93,3; 94, aedeagus ; 95, 9.

REVISION OF AFRICAN SPECIES OF CLEORA aS

Closely similar in colour and pattern to the commonest form of tulbaghata, from
which it can seemingly be distinguished only by the structure of the genitalia ;
variation in pattern is markedly less common than in that species. In some
examples the medial area of each wing is densely suffused with fuscous ; one
example has ochraceous buff bands comparable with tulbaghata ab. flavipleta ; the
holotype remains unique in pattern, with sharply defined transverse fasciae on a
white ground colour (PI. 9, fig. 302).

In the male genitalia the simple sacculus and the shape of the cornuti, and in the
female genitalia the form of the short colliculum and the proportions of the mem-
branous and sclerotized parts of the bursa copulatrix are diagnostic.

Distribution (Map 4). Kenya; Tanzania; Malawi; Rhodesia ; Transvaal ;
Natal ; Cape Province.

Material examined. Holotype 9. ZuLULAND: Edukumbaan Hills, v.1895,
genitalia slide Geometridae No. 2145.

KENYA: Kitale, v.1953 (C. R. Howard), 1 g in Coryndon Museum ; Nairobi
(van Someren), 4 3 ; Kenya coast, Shimba Hills, xii.tg61 (R. H. Carcasson), I 3 in
Coryndon Museum. TANZANIA: Usambara, Amani, iv.1952 (C. Howard), I g in
Coryndon Museum ; E. Usambara, Amani, 11.1953 (E. Pinhey), 1 2 in Coryndon
Museum ; Amani, iii-iv.1936 (B. Cooper), 2 3, 2 9 ; Amani, 28.xi.1934 (Mrs. Edith
Dalton),29. Matawt: Mlanje, 10.iv.1913, 1g ; Mlanje Boma, 2400 ft., 27.1v.1g10
(S. A. Neave), 1 9 ; Chintiche (T. H. Lloyd), 1 9. Ruopesta: Mt. Selinda, 17-31.
1.1959 (G. van Son), 1 3 ; ibid., gQ-17.1v.1956 (G. van Son & L. Vari), 1 g, I 2 in
Transvaal Museum ; Umtali District, 16.i1.1933 (P. A. Sheppard), 1 2 ; ibid., 23.1.
1934, I 9, both in National Museum of S. Rhodesia. TRANSVAAL : Zoutpansburg,
Tshakoma, xi.1931 (G. van Son), 1 9 ; Magoebaskloof Rest Camp, 4000 ft., 21.11.1960
(R. F. Lawrence), 1 § ; Cyprus Farm, near Ofcolaco, 24—28.ix.19g61 (L. Vari), 2 2 ;
Mariepskop, 16.iv.1955 (H. Cookson), 1 g, allin Transvaal Museum ; Marieps Mtn.,
5.Xll.1925—6.1.1926 (G. van Son), 5 3, 2 9; Pilgrim’s Rest Distr., Erasmus Reserve,
30.iv.1960 (F. Neubecker), 4 g, 1 9 in Transvaal Museum ; Pilgrim’s Rest Distr.,
Kowyn’s Pass, 22.11.1962 (L. Vari & Leleup), 1 2 in Transvaal Museum ; Louws
Creek, xi.1922 (H. G. Williams), 1 g. NaAtaL: Hluhluwe, 16.iii.1952 (Dr. G. van
Son), I g in Transvaal Museum ; Pinetown, 6.iii.1g10 (G. F. Leigh), 1 § ; Durban,
ii,v,vi,x (C. G. C. Dickson), 7 3, 1 2 in Transvaal Museum ; Pietermaritzburg,
Malta, i-ii.1928 (G. van Son), 4 3, 3 2 in British Museum (Natural History), 2 2 in
Transvaal Museum. CAPE PROVINCE: Port St. Johns, 10-22.1i.1955 (A. J. T.
Janse), 2 3 in Transvaal Museum.

Cleora lamottei (Herbulot) comb. n.
(Text-figs. 96-98 ; Pl. ro, figs. 305-308, 320 ; Map 5)
Neocleova lamottei Herbulot, 1954 : 322, pl. 1 : 4, text-fig. 13.
Neocleora lamottei Herbulot ; Herbulot, 1958 : 103.

6. First abdominal segment white ; remainder of vestiture white, irrorate with bister ;
patagia edged with bister. Wings white, irrorate with bister ; a broad snuff brown band

74 D. S. FLETCHER

proximad of antemedial fascia on fore wing ; a similar band distad of postmedial fascia on each
wing; anal vein on fore wing and other veins distad of postmedial fasciae interruptedly light buff
to ochraceous tawny (PI. ro, fig. 305). Underside of wings white, densely suffused with bister
(RE romice soo):

g. Genitalia (Text-figs. 96,97). Uncus with minute, thorn-like tip ; scobinate medial plate
of gnathus as broad as cucullus ; a sclerotized, spined semicircular dilation arises at mid-dorsal
margin of valve and extends towards mid-valve, the spining continuous with that of cucullus ;
apex of juxta as broad as gnathus ; apex of sacculus in form of a broad, spatulate and slightly
setose blade-like process ; vesica with two tapered cornuti fused at base, one two-fifths as long
as aedeagus and with a minute, but pronounced and more strongly sclerotized tip, the second
one-fifth as long as aedeagus, stout and with two or three short spines at one side below tapered
apex.

9. Differs from male in upperside of wings, in reduced bister irroration and in having less

96

98

Fics. 96-98. C. lamotte: genitalia. 96, aedeagus ; 97,9; 98, 9.

REVISION OF AFRICAN SPECIES OF CLEORA 75

heavily marked discal spots and in reduction of snuff brown bands in posterior half of fore wing
and on hind wing (PI. 10, fig. 307). Differs in underside in reduction of bister suffusion in proxi-
mal half of each wing (PI. 10, fig. 308).

2. Genitalia (Text-fig. 98). Sterigma almost circular with a sclerotized disc at each side ; a
short, thorn-like projection from lamella postvaginalis medio-posteriorly ; colliculum tapered a
little anteriorly ; anterior third of bursa copulatrix membranous and a little dilate, remainder
cylindrical, ribbed and sclerotized.

Measurements. 9 34-39 mm.; 2 37-46 mm.

In a series of nearly 100 males and 40 females, seven males and five females are
comparable with tulbaghata ab. flavipleta, having a broad band of warm buff to
ochraceous tawny proximad of the antemedial fascia on the fore wing and similar
bands distad of the postmedial fascia on each wing (PI. Io, fig. 320) ; two males and
one female are comparable with tulbaghata ab. fumata, in which the medial area of
the fore wing and the hind wing, proximad of the postmedial fascia, are suffused
with fuscous ; one male combines the characters of both forms.

Tentatively placed after tuwlbaghata and munda on account of the similar range of
variation in pattern shown by the moths ; the form of the female genitalia suggests
an affinity with the following species, toulgoetae, but otherwise appears an isolated
species.

In the male genitalia the form of the gnathus, juxta and sacculus and the pro-
portions of the two cornuti and in the female genitalia the development of the
sterigma are diagnostic.

Distribution (Map 5). Sierra Leone ; Guinea ; Ghana; Nigeria ; Cameroun ;
Gabon ; W. Congo (Leopoldville) ; Principe I. ; Sao Thomé I.

Material examined. Holotype g. GUINEA: Monte Nimba, 30.vili.1g51 (M.
Lamotte & R. Roy) in Muséum national d’Histoire naturelle, Paris.

ST. PRINCIPE : 1500-2000 ft., iv-v.1928 (T. A. Barns), 105,59. SIERRA LEONE :
Ig. Ivory Coast: Bingerville, v,vi,viii-xii (G. Melou), 38 3, 18 9; Agboville,
1-8.vi.1915 (G. Melou),1 9. GHANA: A’koon, 4.i.1919 (G. Harrison), 1g ; Ashanti
(Mrs. D. Houston), 2 3 ; Ashanti, Goaso (G. H. Gibbs), 1 g ; Bibianaha [Bibiani}
70 miles N.W. of Dimkwa, 700 ft., ii,v (H. G. F. Spurrell), 2 3, 2 2 ; Coomassie
[Kumasi] (H. Whiteside), 3 3, 29; Ashanti, Juaso (G. H. Gibbs), 1 g ; Abossi (Dr.
J.J. Wilson), 12; Tarkwa, 3 g,292,; Wassaw distr., 45 miles inland from Sekondi,
Ig; Sekondi, xii.1g21 (N. E. Bell), 1 g. Nicrerta: Lagos, 2 gf; Ilesha (Capt.
L. E. H. Humfrey), 1 3, 1 2; Warri, iv—vili.1897 (Dr. Roth), 4g, 29; Degama
[Degema] (Dr. Ansorge), 1 g ; Ikom, 18.v.1930 (EF. Haig), 1 § ; Old Calabar, 2 3,
29. CAMEROUN : Johann-Albrechts Hohe, 1898 (L. Conradt), 7 3, 2 2; Lolodorf,
1894-1895 (L. Conradt), 6 3; Bitje, Ja River, x, wet season (G. L. Bates), 1 Q ;
Epulan [Efulen], 2.iv.19g26 (G. Schwab), 1 §. GABON: Ogowe, 1890-1894 (L.
Gazengel), 23 ; Lastourville (P. C. Rougeot), 3 3 ; Tchibanga, iv,xi (P. C. Rougeot),
6 g. Conco (Leopoldville) : Kitobola, 1911 (Rovérve), 2 g ; Equateur, Bokote,
19.1.1926 (R. P. Hulstaert), x $ and Tshuapa, Flandria, 18.x.1947 (Rev. P. Hulstaert),
I 9, in Musée Royal de 1’Afrique Centrale.

76 D. Ss. PLEDCHER

Cleora toulgoetae (Herbulot) comb. n.
(Text-figs. g9-101 ; PI. 10, figs. 309-312 ; Map 5)

Neocleorva toulgoetae Herbulot, 1961 : 493, fig. 1.

3g. First abdominal segment white ; remainder of vestiture white, irrorate with pale drab
and bister ; patagia edged with bister. Wings white ; sub-basal area of fore wing and bands
distad of postmedial fascia on each wing pinkish cinnamon to cinnamon ; remainder of wing
patterned and irrorate with bister ; cubital veins and other veins distad of postmedial fasciae
streaked with warm buff (Pl. 10, fig. 309). Underside of wings white, suffused and patterned
with fuscous (Pl. ro, fig. 310).

g. Genitalia (Text-figs. 99, 100). Scobinate medial plate of gnathus almost semicircular,
base as broad as cucullus, scobination extending posteriorly along subscaphium ; apex of juxta
sclerotized broadly ; apical third of sacculus dilate and scobinate, more heavily on inner surface
scobination extending to cover slender, over-curved, spatulate apex ; vesica with two tapered
cornuti fused at base, one one-half and one one-third as long as aedeagus ; the longer cornutus is

10!

Fics. 99-101. C. toulgoetae genitalia. 99, aedeagus ; 100, g ; Io1, 9.

REVISION OF AFRICAN SPECIES OF CLEORA ae

lightly and minutely scobinate on one surface just apicad of middle, the shorter one usually
smooth, but occasionally one or two adpressed spines are situate mid-way along one surface.

9. Upper surface of wings differs in reduction of darker irroration and in better definition of
pattern, especially of the cinnamon transverse bands (PI. 10, fig. 311). Under surface less
suffused and pattern more sharply defined (Pl. 10, fig. 312).

©. Genitalia (Text-fig. 101). Lamella antevaginalis semicircular ; lamella postvaginalis
with angular projection medio-dorsally ; colliculum twice as long as broad, posterior margin
sharply defined ; anterior third of bursa copulatrix ovate and membranous, remainder ribbed
and sclerotized.

Measurements. ¢ 9 32-40 mm.

Variation in the degree and extent of the dark irroration on the upperside of the
wings occurs in the male ; in one female the hind wing, proximad of the medial
fascia, and the posterior half of the medial area of the fore wing are suffused with
fuscous, comparable with tulbaghata ab. fumata. Structurally the form of the female
genitalia suggest an affinity with the preceding species, /amottet. Weakly irrorate
specimens are externally distinct, especially the females, in their small size and in
their cinnamon to pinkish cinnamon transverse bands. Structurally the shape of
the sacculus in the male genitalia and that of the sterigma in the female genitalia
are diagnostic.

Distribution (Map 5). Guinea ; Sierra Leone ; Ivory Coast ; Ghana ; Nigeria ;
Fernando Po ; Cameroun ; Gabon ; Angola ; Congo (Leopoldville) ; Burundi.

Material examined. Holotype g. GuINEA: Tondon, ii.1956 (Mme. M. L. de
Toulgoet), genitalia slide No. C.H.4027 in coll. C. Herbulot, Paris.

SIERRA LEONE : Moyamba (D. Cator),1 9. Ivory Coast: 29,29; Bingerville,
1915 (G. Melou), 1 3, 1 9; Agboville, 1-8.vi.1g15 (G. Melou), 1 9. GHANA:
Bibianaha [Bibiani], 70 miles N.W. of Dimkwa, 700 ft., 19.x.1g10 (H. G. F. Spurrell),
Enchi (Capt. B. D. Peake), 1 9 ; Manso, N.E. of Cape Coast, ili-iv.1922 (N. E. Bell),
Ig. NiceriA: Lagos1d; Ilesha (L. FE. H. Humfrey), 1 2 ; Old Calabar (F. C.
Pudney), 1 9. FERNANDO Po: (W. Cooper), I J. CAMEROUN: xi. (G. Schwab),
IQ; Johann-Albrechts Hohe, 1898 (L. Conradt), 1 9; Bitji, Ja River, 2000 ft.,
wet season, iv—v.1909 (G. L. Bates), 1 g ; ibid., ix—xi.1g1I, 1g. GaBon : Lastour-
ville (P. C. Rougeot), 1 g. ANGOLA: Quicolungo, 120 km. W. of Lucala, 800 m.,
iv.1936 (R. Braun), 1 3. Conco (Leopoldville) : Eala, ix.1936 (J. Ghesquiére), I 2
in Musée Royal de l'Afrique Centrale. Dungu, Upper Uele distr., iv, 1g ; Middle
Lowa Valley, Nr. Walikali, 3000-4000 ft., forest, ii.1924, wet season (T. A. Barns),
Ig. Burunpi : Usumbura, goo m., 8.xii.1g61 (Dr. M. Fontaine), I 3.

Cleora rostella sp. n.
(Text-figs. 102-104; Pl. 10, figs. 313-319; Map 5)

6. Head and thorax white, irrorate with drab, patagia edged with fuscous. Abdomen :
first segment white, second segment fuscous, remaining segments white, lightly irrorate with
drab, posterior margin of each segment with a pair of fuscous spots medio-dorsally. Wings

DS. PREM Crk

78

0

102

102, 2; 103,¢; 104, aedeagus.

C. rostella genitalia.

Fics. 102-104.

REVISION OF AFRICAN SPECIES OF CLEORA 79

white, irrorate with drab and fuscous and, distad of postmedial fasciae, with pale smoke grey ;
transverse fasciae fuscous ; some warm buff irroration proximad of antemedial fascia on fore
wing and distad of postmedial fascia on each wing, well-defined on fore wing between veins M1
and M3; cubital veins and other veins distad of postmedial fasciae interruptedly warm buff ;
a lateral fuscous streak between veins Wz and M3 on fore wing, extending from postmedial
fascia to termen (PI. 10, fig. 313). Underside white patterned with fuscous (PI. Io, fig. 314).

3g. Genitalia (Text-figs. 103, 104). Tip of uncus tapered and depressed ; scobinate medial
plate of gnathus slender and tapered ; apex of juxta broadly Y-shaped and rugose ; apical
process on sacculus short and digitate ; two minute, setose digitate processes arise at mid-valve ;
a further similar, but slightly larger process is situate midway between first pair and ventral
margin of valve ; apex of aedeagus incised ; vesica with two stout, tapered cornuti fused at base,
one one-half and one three-eighths as long as aedeagus.

9. Pattern similar to that of male, but drab and fuscous irroration of ground colour greatly
reduced ; the slender and sharply marked transverse fasciae and the pale smoke grey irroration
distad of the postmedial fascia on each wing and the warm buff irroration and fuscous lateral
streak in the discal area of the fore wing distad of the postmedial fascia are conspicuous (PI. 10,

figs. 315, 316).

9%. Genitalia (Text-fig. 102). LLamella postvaginalis strongly sclerotized medially ; posterior
margin of colliculum sharply defined ; anterior sixth of bursa copulatrix dilate and membranous,
remainder cylindrical, ribbed and sclerotized.

Measurements. ¢g 36-42 mm. ; 2 44-49 mm.

In addition to the most common form described as the typical, this polymorphic
species displays a range of forms similar to those found in tulbaghata ; forms occur
in which the antemedial fascia on the fore wing and postmedial fascia on each wing
are broad and heavily marked (PI. 10, fig. 317) ; in another form the proximal
third of the fore wing and the distal third of each wing are densely fuscous (PI. 10,
figs. 318, 319) ; forms comparable with tulbaghata abs. flavipleta and fumata also
occur.

In the typical form the pale smoke grey irroration distad of the postmedial fascia
on each wing, the warm buff irroration and the fuscous lateral streak in the discal
area distad of the postmedial fascia on the fore wing are distinctive. In the male
genitalia the depressed tip of the uncus, the rugose juxta, the structure of the sacculus
and the incised tip of the aedeagus and in the female the shape and development of
the sterigma are diagnostic.

Distribution (Map 5). Rhodesia ; Malawi; Tanzania; Uganda; Sudan ;
Congo (Brazzaville) ; Burundi ; Congo (Leopoldville) ; Angola.

Holotype g. [Martawi] NyasaLanp : Limbe, ix—x.1926 (H. Barlow), genitalia
slide Geometridae No. 2336.

Paratypes : S. RuopesiA : Chirundu, on Zambesi, 22.11.1950 (N. Mitton), 2 J in
Transvaal Museum : Salisbury, 17.iv.1g16, I g in National Museum of Rhodesia.
[ZAMBIA] N. Ruopesia: N’kana (L. Ellison), 1 § ; Chisorwe [14°52”S. 29°05"E.],
Luano Valley, 11.11.1928 (M. Burr), 1 § ; Mpeta, Loangwa R., aff. of Zambesi,
xi-xl.1895, beginning of rainy season (Coryndon), 3 g. [MALAWI] NYASALAND :
Zomba, 3000 ft., iv.1913 (E. Ballard), 13 ; Zomba, vi.1923 (H. Barlow), 1 2 ; Mlanje
11.1925 (H. Barlow), 1 g ; Magunda Estate, Luchenza (F. Nisbet), 1 9. [TANZANIA]
TANGANYIKA : Old Shinyanga, 29.1.1954 (E. Burtt), 1 9. UGANDA: Kampala,
15.11.1932 (H. Hargreaves), 15. SUDAN : Tambura, Southern Bahr-el-Ghazal, 4 g.

80 Dp. S. FLETCHER

[Conco (Brazzaville)] Conco Repusiic: Fort Crampel, 2 g. BURUNDI:
Usumbura, 900 m., 6—7.vil.1961 (Dv. M. Fontaine), 23,19 ; ibid., 13.viil.1961, 1 Q ;
ibid., 1.x1.1961, 1 g 3 ibid., 11x. 1961, 1 5 abid., 31.x1i-19061, 1°; ibid, goto
1962, 1 6, 1 2; Kitega, 5.v.1962 (Dr. M. Fontaine), i 9, all in Musée Royal de
l Afrique Centrale. [Conco (Leopoldville)] BELGIAN Conco: Region de M’Pala
(R. P. Guilleme), 1 2; E. Lake Kivu, ix.1g19 (T. A. Barns), I 9 ; 150-200 miles
W. of Kambove, 3500-4500 ft., 29.x.1907 (S. A. Neave), 1 gf ; Katanga Distr.,
Kafakumba, i.1927, 1 g'; S.E. Katanga, 4000 ft., 24.xi.1907 (S. A. Neave), 1 g, all in
British Museum (Natural History) ; Elisabethville, 26.ix.1932 (Ch. Seydel), 1 3;
ibid., 27.x.1937, 16 ; ibid., ¥6.xi1.1940, 1 2 ; ibid., x.1948, 14,29 > ibid, 20 xaemg4g,
IQ; ibid., 9.iv.1950, 1 2; ibid., 15.1.1951, I 9, all in Musée Royal de l’Afrique
Centrale; Elisabethville, 23.x.1951 (Ch. Seydel), 1 $ ; ibid., 24-29.xi.1951, 1g, 19 ;
ibid., 23.xil.1953,°1 9°; ibid., 14.x01.1954, 1 9 ; ibid., 71.1956, 1g; ibid, 2a amma5e,
I 9; thid, 2.x1.1956, 1 9 > ibid, 22.x11.1956, 1 Ga ibids 54.1057, sees
28.1.1957,1 9, allin British Museum (Natural History) ; Elisabethville, r. Lubumbashi,
II-24.vi.1914 (Overlaet), I J, 1 2 ; Lubumbashi, xi.1926 (Ch. Seydel), 1 § ; Katanga,
Muteba, xii.1922 (Ch. Seydel), 1 §; Katanga, Luashi, 1933 (Freyne), 2g; ibid., 1935,
1g ; Katanga, Luashi, 3.vii.1924 (Ch. Seydel), 2g ; Lulua, Kapanga, 13—-14.xi.1932
(F. G Overlaet), 4 3 ; ibid., 1.1933, 1 ¢ ; ibid., vil.1933, 1 ¢ ; Lulua, Sandoa, iv.1932
(F. G. Overlaet), 1 J, allin Musée Royal de 1’ Afrique Centrale ; Kasai, Lusambo, 3-10.
ix.I9gI9, I gd, 1 9; Upper Kasai Distr. (P. Landbeck), 1 g, all in British Museum
(Natural History) ; Kasai, Luluabourg, 12.vi.1952 (Dr. M. Fontaine), x 2 ; ibid.,
6.vi.1953, I 6 ; Luluabourg, 24.x.19g21 (Verlaine), 1 § ; Lusambo, 8.1.1926 (Ch.
Seydel), 23 ; Sankuru, Katako-Kombe, 15.xi.1951 (Dr. M. Fontaine), 13 ; Sankuru,
Dimbelenge, 19.v.1953 (Dr. M. Fontaine), 1 § ; Kilo 300 de Kindu (Dr. Russo),
Ig; Kilo 345 de Kindu (Dr. Russo), 1 3 ; Leopoldville, 12.vii.1953 (P. Jobels), 1 @ ;
ibid., 6.x11.1955, I g ; Leopoldville, 29.xi.1953 (Dr. M. Fontaine), 1 3, all in Musée
Royal de l'Afrique Centrale ; Upper Uele distr., Dungu, ix, 1 g in British Museum
(Natural History). ANGoLA: Gamba, Bihé, xii.1934 (R. Braun), 7 3; ibid.,
1.1935, 346; Talala, Benguella, I xii.1905 (Dr. Ansorge), I 3.

Cleora legrasi (Herbulot) comb. n.
(Text-figs. 105-107 ; Pl. 11, figs. 325-329 ; Map 5)

Neocleova legvasi Herbulot, 1955 : 40, fig. 2.

6. Genitalia (Text-figs. 105, 106). Uncus very slightly curved and tapered with a hooked
tip ; scobinate medial plate of gnathus slender and produced ventrad ; apex of juxta rugose ;
cucullus and apical two-thirds of dorsal margin of valve densely clothed with very long setae
extending well beyond dorsal margin of valve ; sacculus without ventral process ; two or three
short, setose, digitate processes arise at mid-valve ; two or three similar, but smaller processes
are situated midway between first series and ventral margin ; vesica with two, stout, tapered
cornuti fused at base, one three-sevenths as long and one one-third as long as aedeagus.

©. Genitalia (Text-fig. 107). Lamella postvaginalis heavily sclerotized medially in a sharply
defined, semi-ovate pattern ; anterior third of bursa copulatrix dilate and membranous, remain-
der ribbed and weakly sclerotized.

Measurements : ¢ 34-44 mm. ; 9 38-44 mm.

REVISION OF AFRICAN SPECIES OF CLEORA 81

Superficially legrasi is closely similar to acaciaria, as Herbulot pointed out in his
original description ; little variation is shown among the series in the British Museum
(Natural History) ; seven specimens in a series of 148 are suffused with fuscous
(Pl. 11, fig. 329) and comparable with tulbaghata ab. fumata, two are similar to
tulbaghata ab. flavipleta, and one combines the features of both.

The hooked tip to the uncus, the rugose juxta and the development of the sacculus

107

Fics. 105-107. C. legvasi genitalia. 105, 3g; 106, aedeagus ; 107, 9.

82 Dp: S. PLEDCHER

reduced to short, setose, digitate processes in the male genitalia and the shape of the
bursa copulatrix and development of the sterigma in the female genitalia relate
legrast closely to the preceding species, vostella. The straighter uncus, the very long
setae on the cucullus, the lack of a process on the ventral margin of the sacculus in
the male genitalia and the pattern of sclerotization of the sterigma in the female
genitalia are diagnostic.

Distribution (Map 5). Madagascar.

Material examined. Holotype g. MaApDAGAscAR centre, Tananarive, Parc de
Tsimbazaza, alt. 1200 m., 3.11.1952 (P. Vvette), in Muséum national d’Histoire
naturelle, Paris.

MapDAGaAscaR:: Diego Suarez, ii,lli,iv,vii.ig17 (G. Melou), 9 fg, 1 9; N.E.
Madagascar, base de la montagne Tsaratanana (ex Lamberton, 1923), I g; E.
Madagascar, Marcantsetra, 2 § ; Central Madagascar, 2500 ft. (fF. B. Pratt), 2 3,
iQ; Tananarive, 1 9; Station Perinet, 149 km. east of Tananarive, i,1i,i1,x,xi,xi
(NV. & G. Olsoufief), 78 3, 18 2 ; Ambinanindrano, 50 km. west of Mahonoro, i,v
(G. K. Kendall-Cornish), 23,192 ; Mananjary, 1918 (G. Melou), 83,12 ; Mananjara,
xi.1918, 44 ; Ankarampotsy, near Fianarantsoa, col. de Tantamaly, xii.1933-1.1934
(R. Catala), 5 5, 1 2; Betsileo, 3000-4000 ft., 1881 (Deans Cowan), I g; S.
Madagascar (Me. Lamberton), 2 g.

Cleora angustivalvis (Herbulot) comb. n.
(Text-figs. 108-110 ; Pl. 10, figs. 321-324 ; Map 5)

Neocleora angustivalvis Herbulot, 1965 : 121.

6. First abdominal segment white, remainder of vestiture white, irrorate with bister ;
patagia edged with bister. Wings white, irrorate with drab and bister and patterned with
bister (Pl. 10, fig. 321). Underside white, patterned with fuscous (PI. Io, fig. 322).

6. Genitalia (Text-figs. 108, 109). Uncus stout and blunt to tip, which is slender and depressed ;
scobinate medial plate of gnathus narrowly rounded and produced ventrad ; apex of juxta
Y-shaped, smoothly sclerotized ; cucullus with normal short setae, ventral edge with four setose,
tubercle-like projections ; sacculus weakly developed and without projecting process ; short,
setose, digitate processes arise near mid-valve, two on right valve, four on left valve, but prob-
ably varying in number individually, vesica with two tapered cornuti fused at base, one one-half
and one slightly longer than one-half as long as aedeagus.

9. Differs from male in sparse irroration and more conspicuous patches of bister distad of
postmedial fascia on fore wing, the larger extending from costa to vein Mj, the smaller between
veins M2 and Cz ; veins on both wings interruptedly warm buff distad of postmedial fascia
(Pl. 10, fig. 323). Underside differs in the reduction of fuscous pattern, especially on hind wing
(Pl. 10, fig. 324).

2. Genitalia (Text-fig. 110). Sterigma weakly developed ; lamella postvaginalis sclerotized
in mitre-shaped pattern ; colliculum tapering anteriorly, three times as long as mean width ;
slightly more than posterior half of bursa copulatrix weakly sclerotized and ribbed, remainder
membranous.

Measurements : ¢ 35 mm.; 2 38-44 mm.

REVISION OF AFRICAN SPECIES OF CLEORA 83

The structure of the uncus and of the sacculus indicate a close affinity with the
two preceding species, vostella and legrasi. Externally the male is similar to a small
pale form of acaciaria, but distinguished from it by the less heavily patterned
underside of the wings ; the female is distinct in the bister patterning of the fore
wing distad of the postmedial fascia and in the very lightly marked underside of
the hind wing.

110

Fics. 108-110. C. angustivalvis genitalia. 108, aedeagus ; 109, 3 ; I10, 9.

84 D. S. FLETCHER

The species is distinguished externally from both rothkirchi insularum and
transversaria, the other species of Cleora occurring in the Comoro Islands, by its
larger size ; in the male the much sparser brown irroration gives angustivalvis a
grey appearance by comparison with males of tvansversaria and in the female the
pattern of both upper and under surfaces of the wings distinguishes angustivalvis.

Structurally the cucullus and the development of the sacculus in the male
genitalia and the form of the sterigma and bursa copulatrix in the female genitalia
are diagnostic.

Distribution (Map 5). Comoro Islands ; probably endemic.

Material examined. Holotype §. Grande Comore, 1884 (L. Humblot).

Paratypes : Grande Comore, 2 g, 11 9; ibid., 1894 (L. Humblot), 6 2 ; Grande
Comoro, vill.191I (G. F. Leigh), 1 9.

Cleora serena sp. n.
(Text-figs. 111-113 ; Pl. 11, figs. 337-340 ; Map 5)

6. Vestiture white, lightly irrorate with drab and bister, except for first abdominal segment
which is immaculate ; patagia lightly edged with bister. Wings white, irrorate with drab and
bister and patterned with bister ; a broken snuff brown band, marked with light buff on veins,
situate immediately distad of postmedial fascia on each wing ; proximal half of subterminal
fascia on each wing irrorate with pale smoke grey (PI. 11, fig. 337). Underside white, patterned
with fuscous (Pl. 11, fig. 338).

6. Genitalia (Text-figs. 112, 113). Uncus with short, thorn-like tip ; scobinate medial plate
of gnathus semi-circular, little broader than uncus ; juxta broadly Y-shaped at apex ; projec-
tion from apex of sacculus minute ; a slightly larger, setose, digitate process at mid-valve,
sometimes asymmetrical and bilobate on one side ; vesica with two short, tapered cornuti
fused at base ; one one-half as long and one two-fifths as long as aedeagus.

9. Similar to male, but drab and bister irroration much reduced, allowing a sharper defini-
tion of pattern, especially of the broad, snuff brown band marked with light buff on the veins,
situate just distad of the postmedial fascia on the upperside of each wing ; pale smoke grey
irroration on subterminal fascia well defined and extending to termen in discal area on fore wing
and in distal half of hind wing (PI. 11, figs. 339, 340).

@. Genitalia (Text-fig. 111). Lamella postvaginalis bell-shaped ; lamella antevaginalis
consisting of a strongly sclerotized lateral band ; colliculum of even width, twice as long as
broad ; anterior two-ninths of bursa copulatrix membranous and globular with a slight, bulbous
projection, remainder cylindrical, sclerotized and ribbed, the ribbing extending very weakly into
membranous part.

Measurements. 4 34-38 mm. ; 2 34-44 mm.

The pattern of the type series is uniformly marked, similar to that of angustivalvis
from the Comoro Islands, to which serena is also closely related in structure of the
genitalia of both sexes. Related closely also in these structures to vostella, with
which it is probably sympatric in Angola. Externally the broad snuff brown band
distad of the postmedial fascia on each wing, well defined in the female, is distinctive ;
structurally the development of the sacculus in the male genitalia and the sterigma
and colliculum in the female genitalia are diagnostic.

Distribution (Map 5). Congo ; Angola.

REVISION OF AFRICAN SPECIES OF CLEORA 85

Holotype g. ANGOLA: Fazenda Congulu, Amboim district, 7-800 m., 17-22.iv.
1934 (Dr. K. Jordan), genitalia slide Geometridae No. 2274.

Paratypes : type locality, 7-11.iv.1934, Ig, 69; ibid., 12-16.iv.1934, 1g, 1 9;
ibid., 17-22.iv.1934, I g, I 9 ; Quirimbo, 75 km. E. of P. Amboim, 300 m., 7—12.v.
1934 (Dr. K. Jordan), 1 g, 11 2; N’Dalla Tando, 2700 ft., 21.xi.1908 (Dr. W. J.
Ansorge), 1 9. CONGO: 1 9.

Fics. 111-113. C. sevena genitalia. 111, 2; 112,g; 113, aedeagus.

86 D. S. FLETCHER

Cleora oligodranes (Prout) comb. n.
(Text-figs. 114-116 ; Pl. 11, figs. 341-344 ; Map 5)
Chogada oligodvanes Prout, 1922 : 358.
Neocleova oligodvanes (Prout) Janse, 1932 : 271, pl. 8: 8; text-fig. Ioo.

6. Vestiture : first abdominal segment white, remainder pinkish cinnamon, irrorate with
fuscous. Wings pinkish cinnamon, irrorate and patterned with fuscous (Pl. 11, fig. 341).
Under surface of wings tilleul buff evenly, but very sparsely irrorate and patterned with fuscous
(Pl. 11, fig. 342).

116

Fias. 114-116. C. oligodvanes genitalia. 114, 9; 115,4; 116, aedeagus.

REVISION OF AFRICAN SPECIES OF CLEORA 87

6. Genitalia (Text-figs. 115, 116). Uncus with minute thorn-like tip ; scobinate medial plate
of gnathus about two-thirds as wide as cucullus ; apex of juxta broadly Y-shaped ; sacculus
setose at apex, sometimes minutely produced ; a slender, setose, digitate process at mid-valve,
longer on left valve than right, and another shorter one, midway between first and ventral
margin ; vesica with two tapered cornuti fused at base, one three-fifths as long, and one one-
half as long as aedeagus, the shorter cornutus sparsely spined along one side.

®. Differs from male in the vinaceous buff ground colour of the wings and in reduction of the
fuscous irroration ; a broad snuff brown fascia is situated proximad of the antemedial fascia on
the fore wing and a similar fascia is situated distad of the postmedial fascia on each wing
(Pl. 11, fig. 343). Underside differs in the greyer ground colour of the wings and less well
defined pattern (Pl. 11, fig. 344).

9. Genitalia (Text-fig. 114). Lamella postvaginalis almost circular, weakly sclerotized except
medially ; anterior fourth of bursa copulatrix membranous, remainder cylindrical, ribbed and
very weakly sclerotized.

Measurements. ¢ 32-37 mm. ; 2 35-41 mm.

Externally variable in degree of development of transverse fascia. Variable also
in neuration ; in the fore wings of six of the 15 examples studied Sci and Scez are
stalked ; in a further example they are coincident, and in two other examples Scg

anastomoses with the stalk of Sc3-s.

The genitalia of both sexes also show a degree of variation. In the male of the
type series the short digitate processes at mid-valve are asymmetrical in two
examples and absent in a third, they are absent also from a Vredendal specimen.
In the female from Ceres the medial sclerotized area of the lamella postvaginalis is
tapered posteriorly.

Externally similar to some forms of guadrimaculata, but distinguished by the
lightly marked under surface of the wings. Structurally the male genitalia indicate
a Close affinity with angustivalvis and serena, but are distinguished by the scobinate,
shorter cornutus on the vesica. In the female genitalia the weakly sclerotized
sterigma is diagnostic.

Distribution (Map 5). Orange Free State ; Bechuanaland ; Cape Province.

Material examined. Holotype g. [ORANGE FREE STATE]: Thaba’nchu, 1.1915
(G. Edelston), genitalia slide Geometridae No. 2138.

ORANGE FREE STATE : type data, 6g, 1 9. BECHUANALAND : Upington, 14.iii.
1950 (H. B. Kettlewell), 1 9. CAPE PROVINCE : Vredendal, 23—30.vii.19g27 (G. van
Son), 1g, 1 2; Karoo, Richmond, 16.iv.1949 (H. B. D. Kettlewell), 1 g ; Ceres,
11.1925 (R. E. Turner), 1 9 ; Kokstad, 27.i11.1952 (C. G. C. Dickson), 1 3, I 9 in
Transvaal Museum ; Pondoland, Port St. Johns, 7—13.vili.1923, I g.

Cleora macracantha (Herbulot) comb. n.
(Text-figs. 117-119 ; Pl. 11, figs. 330-334 ; Map 5)
Neocleova macracantha Herbulot, 1959 : 39, fig. I.

¢g. First abdominal segment white ; remainder of vestiture white, irrorate with snuff brown
and bister ; patagia edged with bister ; posterior margins of abdominal segments, except first
edged dorsally with bister, sometimes as paired spots. Wings white, irrorate with pinkish buff,
snuff brown and bister and patterned with bister ; antemedial fascia of fore wing and postmedial

88

Fics. 117-119.

D. S. FLETCHER

C. macracantha genitalia. 117,92; 118,3; 119, aedeagus.

REVISION OF AFRICAN SPECIES OF CLEORA 89

fascia of each wing sharply defined ; discal spots heavily ringed with bister ; a broad, ill-defined
band of pinkish buff to cinnamon brown proximad of antemedial fascia on fore wing and a
similar band distad of postmedial fascia on each wing ; cubital veins and other veins distad of
postmedial fasciae interruptedly light buff (Pl. 11, fig. 330). Underside of wings white, heavily
patterned with fuscous (Pl. 11, fig. 331).

3g. Genitalia (Text-figs. 118, 119). Uncus short and stout with a thorn-like tip ; scobinate
medial plate of gnathus of even width and rounded at tip ; apex of juxta broadly Y-shaped and
rugose ; sacculus strongly sclerotized, fused with and extending to three-fourths ventral margin
of valve then extending to a tapered tip, level with seven-eighths ventral margin ; a short,
setose, digitate process at mid-sacculus and two similar but minute processes nearer mid-valve;
aedeagus long and slender, one and one-half times as long as sacculus ; vesica with two stout
tapered cornuti fused at base, one one-half as long, the other slightly less than one-half as long as
aedeagus.

Q. Similar to male, but upperside of wings usually less densely irrorate (Pl. 11, figs. 333, 334).

Q. Genitalia (Text-fig. 117). Sterigma asymmetrical and contorted ; posterior medial area
weakly sclerotized ; anterior fourth of bursa copulatrix membranous with two small projections;
remainder of bursa ribbed and sclerotized.

Measurements. ¢g 30-40 mm. ; 2 39-50 mm.

Variation in pattern similar to that of legrasi ; among a series of 160 specimens
are nine examples (PI. 11, fig. 332) comparable with tulbaghata ab. fumata, one male
has the band distad of the postmedial fascia and the posterior half of the fore wing,
except terminal sixth, suffused with warm buff and one female combines the
characters of both forms. Distinguished externally from other known Madagascan
species of Cleora by the clear white ground colour and sharply contrasted and heavily
marked pattern on the underside of the wings. In the male genitalia the develop-
ment of the sacculus and in the female genitalia the asymmetrical sterigma are
diagnostic.

The strongly developed, rugose juxta, the development of the sacculus and the
long smooth cornuti indicate a close affinity with rostella ; the asymmetrical
sterigma, however, remains a unique curiosity in the genus.

Distribution (Map 5). Madagascar ; endemic.

Material examined. Holotype g. MapaGascaR centre, Tananarive, Parc de
Tsimbazaza, alt. 1200 m., 2.ii.1952 (P. Vvette), in Muséum national d'Histoire
naturelle, Paris.

MADAGASCAR : Diego Suarez, ii-ix. (G. Melou), 52 3, 32 9; Kulau (G. Melou),
24; Tamatave et foréts d’Alahakato (Edouard Perrot), 1 3 ; Imarina, 1892 (R. P.
Camboué), 1 g ; Ambinanindrano, 50 km. W of Mahonoro, iii,v,vili,x,xui (G. K.
Kestell-Cornish), 5 3, 3 2; Tananarive, 1933, I g ; Mananjary, 1918 (G. Melou),
129,89; Mananjara, xi.1918, 37 5, 42 ; S. Madagascar, iv.1922 (Lamberton), I 9.

Cleora derogaria (Snellen) comb. n.
(Text-figs. 120-122 ; Pl. 12, figs. 345-348 ; Map 5)

Boarmia derogaria Snellen, 1872 : 73, pl. 6 : 36.
Boarmia obsitaria Mabille, 1890 : 47. Syn.n.
Chogada acaciaria sensu Warren, 1898 : 248.

go

Fics. 120-122.

DiS. PLE CHET:

C. deyogaria genitalia.

120, 29; 121,63; 122, aedeagus.

REVISION OF AFRICAN SPECIES OF CLEORA gI

Chogada acaciaria ab. inusitata Warren, 1898 : 248. Syn.n.
Chogada subspurcata Warren, 1898 : 248.
Boarmia derogaria Snellen ; Swinhoe, 1904 : 532 [Synonymy].

6. First abdominal segment white ; remainder of vestiture buffy brown, irrorate with bister.
Wings buffy brown, irrorate and patterned rather obscurely with bister (Pl. 12, fig. 345). Under-
side of wings tilleul buff densely suffused and heavily patterned with fuscous (Pl. 12, fig. 346).

dg. Genitalia (Text-figs. 121, 122). Uncus stout with a minute thorn-like tip ; scobinate
medial plate of gnathus almost rectangular, one-half as broad as cucullus ; apex of juxta
rugose ; apex of sacculus, triangular in section with one ridge setose, extends to about two-
thirds ventral margin of valve ; a cluster of minute setose, digitate processes at mid-valve
varying in number from four to six ; vesica with two stout, tapered cornuti fused at base, one
two-fifths and one three-fifths as long as aedeagus.

9. Lighter in colour and with better defined pattern than male, ground colour of vestiture
and upperside of wings tilleul buff (Pl. 12, fig. 347). Ground colour of underside of wings white
and much less suffused with fuscous than male (Pl. 12, fig. 348).

@. Genitalia (Text-fig. 120). Lamella postvaginalis with heavily sclerotized medial lobe ;
lamella antevaginalis with medial fold fused with colliculum ; anterior two-fifths of bursa
copulatrix ovate and membranous with two slight bulbous projections, remainder cylindrical,
ribbed and sclerotized.

Measurements. ¢ 35-40 mm. ; 2 39-45 mm.

The tone of the ground colour in the male varies from drab in the paler form to
buffy brown in darker examples ; the female varies in the extent of the dark
irroration. The darker form of the male and the well contrasted paler forms of the
female are readily recognisable by their colour and pattern.

The genitalia of both sexes are similar in general pattern to those of rostella ; those
of the male differ in the shorter, stouter uncus, the broader medial plate of the
gnathus, the detail of the apex of the juxta and in the development of the sacculus ;
those of the female differ in the detail of the sterigma and in the proportionately
longer membranous part of the bursa copulatrix.

Distribution (Map 5). Gambia; Sierra Leone ; Ivory Coast ; Nigeria ; W.
Congo (Leopoldville) ; N. Angola.

Material examined. Holotype g of Boarmia derogaria Snellen (without abdomen):
Banana, Afrika, v. Woerden, in Rijksmuseum van Natuurlijke Historie, Leiden.

Lectotype 3 of Boarmia obsitaria Mabille. From two male syntypes I select as
LECTOTY PEthe specimen labelled : Landana, Boarmia obsitaria Mabille, ex. Musaeo
P. Mabille 1923. Oberthtir Coll. Brit. Mus. 1927-3, genitalia slide Geometridae No.
2135.

Lectotype 2 of Chogada subspurcata Warren. From a series of 5 ¢ and 4 9
syntypes I select as LECTOTYPE the female specimen labelled : Warri, ix.1897
(Dr. Roth), Chogada subspurcata Warren type 2, Rothschild Bequest B.M. 1939-1.

Holotype 9 of Chogada acaciaria ab. inusitata Warren : Warri, vi.1897 (Dr. Roth),
genitalia slide Geometridae No. 2152.

GAMBIA (A. Moloney), 2 3, 3 9. SIERRA LEONE (C. R. Bartlett), 1 3. Ivory
Coast : Bingerville, 1-7.vii.1915 (G. Melou), 1g. At SEA: Between Old Calabar
and Accra (G. E. Bergman), 1 9. NIGERIA: Warrti, iv,vii-ix (Drv. Roth), 63,6;
Degema (Ansorge), 35,29; Niger Coast, 1 2 ; Opobo, 1 ; Rio del Ray, 12.x.1931
(D. R. Rosevear), 1 9. N.ANGoLta : Landana, I g.

92 D. S. FLETCHER

Cleora pavlitzkiae (Fletcher) comb. n.
(Text-figs. 123-135 ; Pl. 12, figs. 349-362 ; Map 6)

Neocleova pavlitzkiae Fletcher, 1958a : 139, pl. 2 : 8, to, pl. 4 : 26-28.

3. Underside of wings white, patterned with fuscous ; postmedial fascia on fore wing usually
well defined from two-thirds costa to lower angle of cell ; costa of fore wing tinged with light
buff ; specimens in which the upper surface is densely irrorate with fuscous black have the
white area of the underside mottled.

Upperside of wings variable in colour and pattern in both sexes.

3g. Genitalia. Uncus evenly stout, apex rounded with a minute thorn-like tip ; scobinate
medial plate of gnathus varies geographically in shape and extent of scobination ; apex of juxta
broadly Y-shaped, sometimes minutely spiculate or rugose ; sacculus with short apical process
projecting at about one-half ventral margin of valve, the shape and extent of the projection
varying geographically ; vesica with two stout, tapered cornuti fused at base, varying slightly
in proportions, but the larger scobinate on ventral surface from one-fourth to seven-eighths.

9. Underside of wings white, very sparsely patterned with fuscous ; in examples where the
upperside is comparable with tulbaghata ab. fumata the fuscous patterning of the underside is
similar to that of the male.

@. Genitalia. Sterigma and colliculum vary geographically ; anterior third of bursa
copulatrix globular and membranous, sometimes with one or two slight bulbous projections
from opposite sides ; remainder of bursa cylindrical, ribbed and sclerotized.

The species may be distinguished externally by the mottled pattern of the
underside of the wings in the male and structurally in the male genitalia by the
larger of the two cornuti being scobinate on the ventral surface for the greater part
of its apical half, only the apex remaining smooth, and in the female genitalia by
the proportions and form of the bursa copulatrix.

Subspecies Jamella, occurring up to an elevation of about 7000 ft. in Kenya and
Tanzania, may have had a wider distribution in Africa ; a single male from Musake at
6350 ft. on Mt. Cameroon has male genitalia closely similar to those of lamella,
differing only in the development of the scobinate medial plate of the gnathus and
probably now represents a distinct subspecies ; a male from the Zomba Plateau in
Malawi and a male from Tsumeb in the northern part of S.W. Africa are similar.
The genitalia of the female from Tsumeb are apparently identical with those of the
nominate subspecies.

Three males and a female from between 3700 and 4500 ft. in Central and Western
Uganda (Kampala, Entebbe and Kilembe) and in E. Congo (Leopoldville),
Mutwanga, are intermediate between subsp. Jamella and subsp. saltuensis ; further
material will probably shew them to represent a distinct subspecies.

Distribution (Map 6). S.W. Africa ; Rhodesia ; Malawi ; Tanzania ; Kenya ;
Ethiopia ; Uganda ; Burundi ; Rwanda ; E. Congo (Leopoldville) ; Cameroun.

Cleora pavlitzkiae pavlitzkiae (Fletcher)
(Text-figs. 123-125 ; Pl. 12, figs. 349-354 ; Map 6)

3 2. Inthe nominate subspecies the ground colour of the upper surface of the wings is white,
usually weakly irrorate but very rarely with a shade darker than bister (PI. 12, figs. 349-352) ;
forms occur that are comparable with tulbaghata ab. flavipleta and a variation of the flavipleta
form has the medial area of the fore wing and the proximal half of the hind wing almost immacu-

REVISION OF AFRICAN SPECIES OF CLEORA 93

late (Pl. 12, fig. 353) ; a female from Vumba has the upper surface of the wings patterned with
fuscous black (Pl. 12, fig. 354).

3g. Genitalia (Text-figs. 123, 124). Scobinate medial plate of gnathus semicircular, at base
three-fifths as broad as cucullus ; projection of sacculus beyond ventral margin of valve blunt,
slightly incurved and setose but not scobinate, two-fifths as long and one-eighth as broad as
width of cucullus ; two cornuti, one three-sevenths and one four-sevenths as long as aedeagus.

Fics. 123-125. C. pavlitzkiae pavlitzkiae genitalia. 123, aedeagus ; 124,45; 125, 9.

94 D, S. PLEDCHER

Q. Genitalia (Text-fig. 125). Central area of lamella postvaginalis sclerotized in bell-shaped
pattern. Colliculum strongly developed with sharply defined posterior margin ; twice as long
as mean width, tapered anteriorly.

Measurements. ¢ 37-45 mm. ; 92 41-45 mm.

Distribution (Map 6). Rhodesia ; Malawi.

Material examined. Holotype g. S. RHopEsIA: Vumba, 7.xi.1936 (J. E.
Drysdale), genitalia slide Geometridae No. 2171.

S. RHopEsIA : Vumba Mts., Umtali, ii,xui, 7 g, 1 9 and Vumba, 16.iii.1925
(P.A.S.), I g, in National Museum of Rhodesia ; Vumba, ii,xi (G. van Son), 2 4,
2 9 and Mt. Selinda, xii.1935 (G. van Son), 2 3, in Transvaal Museum ; type data,
49. Marawr: Port Herald, Nr. Zambezi, vi.1926 (H. Barlow), 1 g.

Cleora pavlitzkiae lamella ssp. n.
(Text-figs. 126, 127 ; Pl. 12, figs. 355, 356; Map 6)

36°. Wings in most examples patterned and densely irrorate with bister to fuscous black

(Pl. 12, figs. 355, 356).

3g. Genitalia (Text-fig. 127). Medial plate of gnathus as in nominate subspecies ; projection
of sacculus beyond ventral margin of valve spatulate, straight and setose, three-fifths to four-
fifths as long as width of cucullus ; proportions of cornuti as in nominate subspecies.

2. Genitalia (Text-fig. 126). Similar to those of the nominate subspecies, but with a slightly
broader posterior margin to the colliculum.

Measurements. ¢ 38-50 mm. ; 9 44-46 mm.

Distribution (Map 6). Kenya ; Tanzania.

Holotype g. Kenya: Nanyuki, x.1961 (R. H. Carcasson), genitalia slide
Geometridae No. 5288.

Paratypes : Kenya: Nairobi, v.1905 (Jackson), 1 g ; Nairobi, Ngong, vii.1954
(Fowler & Coulson), 1 g ; Mt. Kenya, Naro Moru, 7000 ft., 20.vi.1949 (J. A. Riley),
Ig; Mt. Kenya, Meru Distr., ix.1930 (Mrs. H. Young), 1 § ; Between Nanyuki
and Meru (FE. Barnes), 1 § ; Nakuru, bred (A. L. H. Townsend), ix.1936, 1 g ; ibid.,
1.1937, 1 6% bids, vil.r937, © 2 ; ibid) 1%.1937, 2-6 ; abid., 1.1938, 1 Seine
9.1.1943,'-1-¢°3 ibid, 131.1944 1 Gg; Mt. Elgon, 1.1053 (7. BE? jackson) aes:
ibid., 1.1959, I g in Coryndon Museum.

Other material. [TANZANIA] TANGANYIKA : Morogoro, Kibuku, 1 9 ; Njombe,
6000-7000 ft., 19.x.1947 (W. Peters), 1g ; ibid., 19.x.1951, 1g ; Mufindi (P. Burdon),
246,292; Marangu, I-20.iii.1959 (Lindner), I g in Zoological Museum, Stuttgart.

Cleora pavlitzkiae etesiae ssp. n.
(Text-figs. 128-130 ; Map 6)

$69. Externally similar to paviitzkiae lamella in having the upper surface of the wings densely
irrorate and patterned with bister.

g. Genitalia (Text-figs. 128, 129). Scobinate medial plate of gnathus as in nominate sub-
species ; projection of sacculus spatulate and incurved through ninety degrees, one-fourth as

REVISION OF AFRICAN SPECIES OF CLEORA 95

broad as cucullus, minutely scobinate and extending beyond ventral margin of valve ; two
cornuti, one one-third and one three-fifths as long as aedeagus.

2. Genitalia (Text-fig. 130). Lamella postvaginalis similar in form and depth to those of the
preceding subspecies, but more heavily sclerotized medio-posteriorly ; well defined posterior
margin of colliculum fused with strongly sclerotized anterior margin of lamella antevaginalis.

Measurements. ¢ 38-50 mm. ; 2 44-48 mm.

126

Fics. 126-127. C. pavlitzkiae lamella genitalia. 126, 2; 127,64.

96

Fics. 128-130.

D. S. PEE TDCHER

C. pavlitzkiae etesiae genitalia. 128, aedeagus ; 129,45; 130, 9.

REVISION OF AFRICAN SPECIES OF CLEORA 97

Only one specimen, a female, shows any marked variation in pattern ; the bister
irroration on the upper surface of the wings is reduced, the antemedial fascia on
the fore wing and the postmedial fascia on each wing are broad and heavily marked.
This example is similar in appearance to the form of the nominate subspecies
illustrated on Pl. 12, fig. 353.

The incurved, spatulate and minutely scobinate projection of the sacculus and the
proportionately shorter cornuti in the male genitalia and the strongly sclerotized
margin of the lamella antevaginalis in the female genitalia are diagnostic.

Distribution (Map 6). Ethiopia.

Holotype g. [EruHiop1A] ABYSSINIA : Harar, 4.vi.1939 (R. E. Ellison), genitalia
slide Geometridae No. 5322.

132

Fics. 131-132. C. pavlitzkiae oriadelpha genitalia. 131, 3; 132, 9.

98 D. Sv FPLBICHER

Paratypes ; [ETHIOPIA] ABYSSINIA: Harar, 18.iii.1939 (R. E. Ellison), 1 9;
ibid., 9.vi.1939, Ig ; ibid., 1x=x.1930, 1g, 14-x.1939; I 9, 30.x.1938, I gy 26:mEags,
I g, 5-xil.1937, I ¢ ; Addis Ababa, 21.iii.1939 (T. Wikely), 1 g ; Addis Ababa,
20.11.1948 (K. M. Guichard), 1 g ; ibid., 7.vii.1948, 2 g.

Cleora pavlitzkiae oriadelpha ssp. n.
(Text-figs. 131, 132 ; Pl. 12, figs. 357, 358 ; Map 6)

6 2. Variable in pattern, but fairly densely irrorate with fuscous as in the subspecies lamella
and etesiae (Pl. 12, figs. 357, 358).

3d. Genitalia (Text-fig. 131). Scobinate medial plate of gnathus as in nominate subspecies ;
projection from sacculus spatulate and minutely scobinate, apex obtusely incuved one-third as
broad as cucullus and with less than one-half of the width extending beyond ventral margin of
valve ; two cornuti, one one-third and one five-eighths as long as aedeagus, closely similar in
proportion to those of subspecies efesiae.

9. Genitalia (Text-fig. 132). Lamella postvaginalis shallow, posterior margin with a heavily
sclerotized, raised tip medially ; lamella antevaginalis twice as deep as lamella postvaginalis and
evenly sclerotized ; colliculum of even width, three times as long as broad, posterior margin
sharply defined.

Measurements. ¢ 46-50 mm. ; 2 40-45 mm.

The few known males appear externally distinct in their consistently large size, a
not unexpected character in view of the high elevation at which they occur.
Structurally the shape of the projection from the cucullus and the proportions of
the cornuti in the male genitalia and the form of the sterigma in the female genitalia
are diagnostic.

Probably confined to the high ground of the Mau Escarpment. Two specimens
labelled “‘ Nakuru, bred”’ suggest an area of overlap between subspecies lamella
and subspecies oriadelpha ; there is, however, no additional data to indicate either
the elevation or the precise source of the early stages from which these adults were
reared. Since Nakuru is so close to the south-eastern extremity of the Mau
Escarpment, it is possible that the early stages came from that region.

Distribution (Map 6). Kenya, Mau Escarpment.

Holotype 6. KEnya: Kaptagat, [7867 ft.], 26.xi.1948 (Walker).

Paratypes : Kenya: Kaptagat [7867 ft.], iii.1948 (Walker), 1 § ; ibid., 31.vii.
1948, 1 2; Mile 478 on Uganda Rly. [Molo, 8065 ft.], 14.xi.1g00 (C. S. Betton), 1 3 ;
Nakuru, bred, 27.vii.1948, 1 9 ; ibid., 18.vii.1948, 1 9.

Cleora pavlitzkiae saltuensis ssp. n.
(Text-figs. 133-135 ; Pl. 12, figs. 359-362 ; Map 6)

369. Externally intermediate in the male between the nominate subspecies and subsp.
lamella, being more densely irrorate with bister than in the former, but not so darkly coloured as
the latter ; veins interruptedly but prominently warm buff to ochraceous orange distad of the
postmedial fascia in most examples (Pl. 12, figs. 359, 360). The female differs from that of the
nominate subspecies in the light, but even fuscous irroration of the upperside of the wings
(Pl. 12, figs. 361, 362).

REVISION OF AFRICAN SPECIES OF CLEORA 99

6. Genitalia (Text-figs. 133,134). Scobination of medial plate of gnathus broad andextending
posteriorly along subscaphium ; projection from sacculus spatulate and minutely scobinate at
apex and along inner edge, three-fourths as broad as cucullus ; two cornuti, one one-half and
one one-third as long as aedeagus, the scobination on the ventral surface of the larger cornutus
reduced to a short patch in apical half, equal in length to one-fourth of the cornutus ; shorter
cornutus bears one or two adpressed spines in some examples.

Fics. 133-135. C. pavlitzkiae saltuensis genitalia. 133, aedeagus ; 134, 3; 135, 9.

100 D.*S: MVE DRCHER

@. Genitalia (Text-fig. 135). Lamella antevaginalis and lamella postvaginalis of about even
depth, the latter evenly sclerotized, slightly produced and tapered medio-posteriorly ; colli-
culum of even width, three times as long as broad.

Measurements. ¢ 36-46mm.; 2 41-44 mm.

There is little variation in colour and pattern in the series examined ; one male
example is similar to the form of the nominate subspecies illustrated on PI. 12, fig.
353, with heavily marked transverse fasciae, and two males are comparable with
tulbaghata ab. fumata, the medial area of the fore wing and the proximal half of the
hind wing being suffused with fuscous. Structurally the extensive scobination of
the medial plate of the gnathus and subscaphium and the form of the projection
from the sacculus in the male genitalia, and the even depth of the lamellae and the
more evenly sclerotized lamella postvaginalis are diagnostic.

Biology. Adults have been reared from larvae found on Pinus patula and
Cupressus lusitanica in W. Uganda.

Distribution (Map 6). Burundi ; Rwanda ; Central Eastern Congo (Leopold-
ville) ; S.W. Uganda, Kigezi.

Holotype §. Burunpi: Kitega, 17.vii.1962 (Dr. M. Fontaine), in Musée Royal
de l'Afrique Centrale.

Paratypes: BurunbDi: Kitega, g—21.11.1962 (Dr. M. Fontaine), 2 2; ibid.,
17.1v.1962, I 4, 8.vi.1962, I 9, 13-28.vil.1962, 7 J, 2 9, I-29.vili.1962, I0 g, 5 Q,
Q-I5.X.1962, 2 g, allin Musée Royal de l'Afrique Centrale. [RwaAnpa] : Lake Kivu,
Rugege Forest, 7000 ft., x.1g21 (IT. A. Barns), I § in British Museum (Natural
History). [Conco (Leopoldville)] BELGIAN ConcGo : Rutschuru, 1.1928 (Ch. Seydel),
1g: Kivu, Ters. Lubero, Mulo;.o.x.19054 (R. P. M. J. Celts), 159 ; N. Laer
Rwankwi, iv—v.1948 (J. V. Leroy),23,1 ; ibid., 16—29.vill.1947, Ig, I 9, 6.1x.1947,
I g, 19.1x.1947, I g, I.x1.1947, I g, xil.1947, 1g ; Kivu, Nyamunyunga (Mulungu),
v.1960 (J. Hecq), 1 3, allin Musée Royal de l’Afrique Centrale ; Mt. Hoyo, 31.x.1956
(Ch. Seydel), 1 g. UGANDA: Kigezi, Mafuga Forest, il.1952 (T. H. E. Jackson),
292; Mafuga Forest, 17.vii.1961, ex Pinus patula (W. K. Brown), 1 3; ibid.,
I4.vil.1962, I g, 23.vii.1962, 1 9; Mafuga, g.vill.1g62, ex Cupressus lusitanica
(W. K. Brown), t 9.

Cleora lima sp. n.
(Text-figs. 136, 137 ; Pl. 12, figs. 363, 364 ; Map 8)

3d. Vestiture and upper surface of wings closely similar to pavlitzkiae saltuensis. Under
surface of wings white, suffused and patterned with fuscous (PI. 12, fig. 364) ; the dark suffusion
is markedly more extensive than that found in pavlitzkiae saltuensis.

6. Genitalia (Text-figs. 136, 137). Uncus stout with minute thorn-like tip ; scobinate
medial plate of gnathus semi-circular ; apex of juxta slightly rugose ; apical projection of
sacculus cylindrical, cygnate and scobinate one and one-fourth times as long as width of
cucullus ; vesica with two tapered cornuti fused at base, one one-half and one one-third as long
as aedeagus, the longer cornutus scobinate on ventral surface from one-half to three-fourths of
its length, the shorter one smooth.

2. Not known.

Measurement. ¢ 39-42 mm.

REVISION OF AFRICAN SPECIES OF CLEORA 101

Closely related to and sympatric with pavlitzkiae saltuensis ; distinguished from
it externally in the more darkly suffused under surface of the wings and structurally
in the less extensively scobinate medial plate of the gnathus, the differently
developed apex of the juxta and strikingly in the shape of the sacculus.

Distribution (Map 8). E. Congo (Leopoldville) ; Burundi ; Rwanda.

Holotype g. [Conco (Leopoldville)] : Rwankwi, iv.1948 (J. V. Leroy), in Musée
Royal de |’Afrique Centrale.

Paratypes : Burunpi: Kitega, 25.viii.1g62 (Dr. M. Fontaine), 1 3 in Musée
Royal de l'Afrique Centrale. [RwaNnpbDA]: Lake Kivu, Rugege Forest, 8000 ft.,
xil.Ig2t (IT. A. Barns), 1 g. [Conco (Leopoldville)]: N.W. Kivu, Upper Oso River,
4000 ft., forest with some grass, ii.1924, wet season (7. A. Barns), I 3.

Fics. 136-137. C. lima genitalia. 136, ¢; 137, aedeagus.

Cleora scobina sp. n.
(Text-figs. 138-140 ; Pl. 9, figs. 303, 304 ; Pl. 11, figs. 335, 336; Map 5)

g. Vestiture white, lightly irrorate with snuff brown and fuscous, except on first abdominal
segment ; patagia edged with bister. Wings white, lightly irrorate with cinnamon drab and
bister ; a broad fascia proximad of antemedial fascia on fore wing and a similar fascia distad of

102 D2 S. FEET CHER

postmedial fascia on each wing cinnamon drab ; other transverse fasciae bister tinged with
tawny ; cubital veins and other veins distad of postmedial fasciae streaked light to warm buff,
streak on My, of fore wing dilate to large spot. (Pl. 11, fig. 335). Under surface of wings
white, sharply patterned with bister (Pl. 11, fig. 336).

6. Genitalia (Text-figs. 138, 139). Uncus slender with minute thorn-lke tip ; scobinate
medial plate of gnathus one and one-fourth times as broad as cucullus, slightly produced medio-

\
‘

Fics. 138-140. C. scobina genitalia. 138, aedeagus ; 139, ¢ ; 140, 9.

REVISION OF AFRICAN SPECIES OF CLEORA 103

ventrally ; scobination extending posteriorly to subscaphium for distance equal to one-half of its
width ; juxta slightly rugose at apex ; projection of sacculus tapered, spatulate, scobinate and
slightly setose, equal in length to uncus ; vesica with two tapered cornuti fused at base, one
three-tenths and one two-fifths as long as aedeagus, each scobinate on one surface in apical half.

Female patterned similarly to male on upper surface of wings but dark irroration sparse ;
cinnamon drab bands reduced and broken ; transverse fasciae very slender, bister to fuscous,
lacking tawny tinge of male (PI. 9, figs. 303, 304).

®. Genitalia (Text-fig. 140). Sterigma ovate ; lamella postvaginalis with a raised, tapered
lobe medio-posteriorly ; anterior fourth of bursa copulatrix membranous and ovate, remainder
cylindrical, ribbed and sclerotized and obtusely bowed.

Measurements. ¢ 37-42 mm. ; 2 38-44 mm.

Specimens from E. Congo (Leopoldville) are rather more bister in general tone
than the pinker brown examples from Kampala and other localities in Western
Uganda, but in other respects the series shows little variation.

A short series of five specimens from the Ivory Coast is provisionally associated
with scobina, though excluded from the type series. The specimens are rather
smaller and a little more heavily marked, but agree structurally with scobina in the
genitalia of both sexes. Closely allied to pavlitzkiae and to lima, differing externally
from the former in the pattern of the under surface of the wings and from both in
the warmer pinker or redder brown tone of the markings of the upper surface of the
wings. In the male genitalia the form of the scobinate medial plate of the gnathus
and the form of the sacculus and in the female genitalia the form of the sterigma are
diagnostic.

Biology. Adults have been reared from larvae found on Cupressus lusitanica
and Eucalyptus torelliana in Western Uganda.

Distribution (Map 5). Uganda ; Burundi ; Congo (Leopoldville) ; Ivory Coast.

Holotype g. UGANDA: Kampala, 22.vili.1932 (H. Hargreaves), genitalia slide
Geometridae No. 22091.

Paratypes : UGANDA: Kampala, 7.vii.1927 (H. Hargreaves), 1 $ ; ibid., pupated
13.V.1931, emerged 22.v.1931, I 9; ibid., 11.vii.1932, 1 ¢ ; Kampala, 29.vii.1925,
Ig ; Kampala, 26.xi.1929 (G. H. E. Hopkins), 1 3 ; Entebbe, 1.iv.1g00 (Capt. H. B.
ex Cupressus lusitanica (W. K. Brown), 1 2; Mpanga Forest, 21.vii.1962, ex
Eucalyptus torelliana (W. K. Brown),1 9. Burunpi : Usumbura, goo m., 8.vil.1g61
(Dr. M. Fontaine), 1 3, in Musée Royal del’ Afrique Centrale. [Conco (Leopoldville)]
BELGIAN ConGo : Ruwenzori, 18—20.v.1950 (H. B. D. Kettlewell), 3 3,292; Takalu,
W. of Lake Albert, iv, r g ; E. Ituri Valley, 30 miles south of Irumu, 3000 ft.,
vii.1924 (T. A. Barns), 1 g ; Upper Uelle distr., Dungu, iv,v,vii, 3 ¢ ; Upper Uelle
distr., Sabuni, v, 1 3, all in British Museum (Natural History) ; Kibali-Ituri, Nioka,
I0.vli.1953 (J. Hecg), 1 2 ; Haut-Uelé, Moto, 1921, 2 ¢ ; Rutschuru, 2.1.1928 (Ch.
Seydel), 1g ; N. Lac Kivu, Rwankwi, 4.ix.1947 (J. V. Leroy), 1 g ; ibid., iv.1948,
2; ibid., v.1948, 1g ; Lulua Kapanga, xii.1932 (F. G. Overlaet), 1 3, all in Musée
Royal de |’Afrique Centrale.

Other material. Ivory Coast: Bingerville, 1915 (G. Melou), 1 g, 1 9; ibid.,
I-7.Vi.1915, I 2 ; ibid., LI-14.viii.I9g15, Ig, I Q.

104 D. S$: FLETCHER

Cleora radula sp. n.
(Text-figs. 141-143 ; Pl. 13, figs. 365-379 ; Map 7)

6. Colours of vestiture and of upper and under surfaces of wings vary geographically.

6. Genitalia (Text-figs. 141, 142). Umncus with minute thorn-like tip ; scobinate medial
plate of gnathus semicircular, diameter equal to one-half width of cucullus ; apex of juxta
tugose ; arm of sacculus spatulate, tapered and scobinate, subequal in length to uncus, varying
a little individually in width ; vesica with two cornuti fused at base, one one-third and one
one-sixth as long as aedeagus, the longer one densely and quite coarsely scobinate in apical

two-thirds and usually very little tapered, the shorter one tapered and usually completely
smooth.

aay,
Ls APA be

Pa 143

Fics. 141-143. C.vadula genitalia. 141, aedeagus ; 142, 4; 143, 9.

REVISION OF AFRICAN SPECIES OF CLEORA 105

9. Vestiture white, lightly irrorate with bister ; patagia edged with bister. Wings white ;
fore wing with a slender snuff brown band proximad of antemedial fascia ; similar band distad
of postmedial fascia on each wing, that on the fore wing broadening to subterminal at costa and
failing at inner margin, that on the hind wing failing at costa ; remainder of wing very lightly
irrorate and patterned with bister (Pl. 13, fig. 367). Underside of wings white, patterned with
bister (Pl. 13, fig. 368).

9%. Genitalia (Text-fig. 143). Sterigma with large, clearly defined ostium bursae ; colliculum
twice as long as broad with a short projection medio-posteriorly ; anterior fourth of bursa
copulatrix membranous ; remainder of bursa cylindrical, ribbed, sclerotized and scobinate on
inner surface ; posterior margin of seventh sternite sclerotized and shallowly concave medially.

The scobinate and spatulate projection from the sacculus in the male genitalia
and the structure of the sterigma in the female genitalia suggest an affinity with
scobina and lima. From these species vadula may be distinguished in the male
genitalia by the less extensive scobination of the gnathus and by the dense scobin-
ation and form of the larger of the two cornuti ; in the female genitalia the large,
clearly defined ostium bursae in the sterigma serves to separate rvadula from scobina ;
the female of /ima is not yet known.

Distribution (Map 7). Guinea ; Ghana ; Nigeria ; Fernando Po ; Cameroun ;
Angola ; Congo (Leopoldville) ; Uganda ; Kenya.

Cleora radula radula ssp. n.
(Pl. 13, figs. 365-369 ; Map 7)

6. Vestiture drab, irrorate with bister ; patagia edged with bister. Wings tilleul buff,
densely irrorate with snuff brown and fuscous ; sub-basal fascia on fore wing and a band distad
of postmedial fascia on each wing snuff brown ; cubital veins and other veins distad of post-
medial fasciae streaked with light buff ; other transverse fasciae and discal spots fuscous (PI. 13,
fig. 365). Under surface tilleul buff near discal spots, remainder of wings suffused and patterned
with fuscous (Pl. 13, fig. 366).

Q. (PI. 13, figs. 367, 368). See under description of species.

Measurements. 4 38-44 mm. ; 2 34-44 mm.

Fifteen of the 43 male examples are darker than the type, bister replacing snuff
brown and fuscous black replacing fuscous ; in such specimens the transverse
fasciae in the distal third of each wing are edged with a sparse white irroration. In
six other males the fore wing is suffused with light buff between the submedial fold
and vein Ai, from base to subterminal fascia, and the bands distad of the post-
medial fascia on each wing, snuff brown in the type, are similarly suffused ; in these
examples the hind wing is suffused with light buff especially distad of the postmedial
fascia (Pl. 13, fig. 369). The darker male examples are externally similar to
derogaria, but have a more sharply defined pattern ; the genitalia are, however,
quite distinct.

Eleven males from Angola, apparently intermediate between typical radula and
the following subspecies, are provisionally associated with the nominate series, but
excluded from the type series ; the ground colour of the wings is paler, clearly
visible distad of the discal spot on the fore wing and the dark irroration is less dense.

Distribution (Map 7). Equatorial forest of Gabon and Congo (Leopoldville) ;
Angola.

106 DES PIE CEE

Holotype g. [Conco (Leopoldville)] : Lusambo, 29.vi.1949 (Dr. M. Fontaine),
in Musée Royal de l'Afrique Centrale.

Paratypes : GABON: Lastourville, 1.1959 (P. Rougeot), 1 J, in British Museum
(Natural History). [Conco (Leopoldville)] : Leopoldville, Binza, 29.ix.1954 (Dr.
M. Fontaine), 1 9; Stanleyville, 9.vi.1948 (Dr. Faniel), 1 § ; Sankuru, Djeka,
17.xil.1952 (Dr. M. Fontaine), 1 § ; Sankuru, Komi, vi.1939 (J. Ghesquiére), I ¢ ;
Sankuru, Lusambo, 7.v.1950 (Dr. M. Fontaine), 1 f ; ibid., 14.vi.1949, If ; 17.vi.
1949, 13,19; 20.vi.1949, 1d ; 2.vil.1949, 1g ; 14.VUl.1950, 1g ; 18.vil.1950, 14 ;
23.VU.1950, 14,19 ; 2.vili.1950, I 9 ; O6.vilil.1950, 1g ; 7.vVill.1950, 2¢ ; 8.vill.1950,
Ig} 9Q-Vill.1950, 1 ¢ ; 14:Vill.1950, 1d ; 31-vill.1950, 1G > 1%.1950, 1d 3 ees,
I4.ix., each Ig; xX.1950, Ig; II.x.1950, I g ; Sankuru, Katako-Kombe, 16.iii.
1952 (Dr. M. Fontaine), 1 3 ; ibid., 19.11.1952, 15 ; 6.1x.1952, 1g ; 15.1X.1952,1¢ ;
8.x.1952, 1g 3 14.xi.1952, 1 g; Uele, Paulis, 19.11.1956 (Dr. M. Fontame) maa
ibid., 2.11.1956, I 9; 4.iv.1956, 1g; 8.iv.1956, Ig; 12.iv.1956, If ; 4.vii.1956,
Ig; 28.xi.1952,1¢ ; I.xii.1956,1¢ ; ibid., 28.xii.1955, 1 9 ; Ituri, Nduye-Makara,
x.19g2I (A. Pilette), 1 3, all in Musée Royal de 1’Afrique Centrale ; Kondolola
District, Lindi Valley, 1600-1700 ft., v.1g21 (IT. A. Barns), 2 § in British Museum
(Natural History).

Other material. ANGOLA: Quicolungo, 120 km. N. of Lucala, 800 m., iv.1936
(R. Braun), 11 3.

Cleora radula leptopasta ssp. n.
(PL. 13, figs: 370,371 ¢ Map 7)

6. Differs from nominate subspecies in colour. Ground colour of wings white ; antemedial
fascia on fore wing, medial and postmedial fasciae, terminal interneural spots and outline of
discal spots on each wing bister ; remaining transverse fasciae snuff brown ; light irroration
snuff brown and bister. (Pl. 13, fig. 370). Under surface of wings white, suffused and patterned
with bister (Pl. 13, fig. 371).

Measurements. 4 38-43 mm.

Externally closely similar to darget with which it occurs and only reliably
determined by reference to the genitalia.

Distribution (Map 7). Ghana ; Nigeria.

Holotype g. [NIGERIA]: Lagos, genitalia slide Geometridae No. 5469.

Paratypes : [GHANA] GoLD Coast : Coomassie [Kumasi] (H. Whiteside), 2 3 ;
Nsuaem, 1.1922 (NV. E. Bell), 1g. NiGeria: Lagos, 1g ; Iesha (L. E. H. Humfrey),
IG. 5. Nigeria ig.

Specimens from the localities listed below are associated provisionally with subsp.
leptopasta, but excluded from the type series. The males from Fernando Po and
from Lolodorf and Johann-Albrechts Hohe in Cameroun are more densely irrorate
with bister ; no male from Guinea is available for study.

REVISION OF AFRICAN SPECIES OF CLEORA 107

GUINEA : Massadou near Macenta, 1000 ft., 13-17.v.1926 (C. L. Collenette), 1 9.
FERNANDO Po: (W. Cooper), 2 g, 1 9. [CAMEROUN]: Afriq. Occid., Station
Kamerun, Lolodorf, 1894-1895 (L. Conradt), 1 3 ; Johann-Albrechts Hohe (L.
Conradt), 23,19; Bitje, Ja River, 1 3.

Cleora radula arenosa ssp. n.
(Pl. 13, figs. 372-376 ; Map 7)

g. Ground colour of upper surface of wings white ; sub-basal band on fore wing and band
distad of postmedial fascia on each wing russet to ochraceous tawny and often diffuse ; streaking
on veins ochraceous tawny (PI. 13, figs. 373, 374). In three examples medial area of fore wing
posterior of cubitus and hind wing proximad of postmedial fascia suffused with fuscous black
(El 03, fe. 372).

9. Only specimen known possibly aberrant ; area proximad of medial fascia on upperside of
each wing suffused with bister ; strongly marked bister spot on fore wing between veins MW,» and
Cu; and between postmedial and subterminal fasciae (Pl. 13, figs. 375, 376).

Measurements. ¢ 38-41 mm.; 9 43 mm.

The male is strikingly distinct in the presence of the conspicuous russet to
ochraceous tawny sub-basal band on the fore wing and the similarly coloured
postmedial bands on each wing.

Distribution (Map 7). E. Congo (Leopoldville), Mt. Hoyo and Kivu ; W. Uganda.

Holotype g. UGANDA : Bwamba, vi.1956 (R. Carcasson).

Paratypes : [Conco (Leopoldville)] BELGIAN Conco : Mount Hoyo, 10.ii.1956
(Ch. Seydel), 1 3; ibid., 1.111.1956, 1 g ; Ituri Forest, 3800-4000 ft., i1-iv.1930,
beginning wet season (Lord Howard de Walden Exp.), 1g; Escarpment West
Semliki Valley, 20 mls. S.W. of Boga, 3000-4000 ft., vii.1g24, borders of tropical
forest and long grass (T. A. Barns), 1 g ; N. Lac Kivu, Rwankwi, iv.1948 (J. V.
Leroy), 2 § and 19.vill.1947, I g in Musée Royal de |’Afrique Centrale ; Takulu, W.
of Lake Albert, 1 ¢ ; W. Kivu, south side middle Lowa Valley, south of Walikali,
3000 ft., forest, ili.1924, wet season (T. A. Barns), 1 § ; W. Kivu, Upper Lowa
Valley, nr. Masisi, 5000-6000 ft., forest and long grass, ii.1924, wet season (T. A.
Barns), 2 3; W. Kivu, Lowowo Valley, South Lowa District, 4000 ft., mountain
forest, 111.1924, wet season (7. A. Barns), 1 g. UGANDA: Bwamba, v.1956 (R.
Carcasson), 33 ; ibid., vi.1956, 19 ; Kigezi District, Impenetrable Forest, Kanungu,
4500 ft., v.1952 (J. A. Burgess), 1 3.

Cleora radula eumelana ssp. n.
(Pl. 13, figs. 377-379 ; Map 7)

6. Ground colour of wings on upper surface cartridge buff, weakly suffused with russet and
irrorate with black ; veins streaked with ochraceous tawny ; remainder of pattern black
(Pl. 13, fig. 378). Under surface tilleul buff, suffused and patterned with fuscous black (Pl. 13,
fig. 379).

Measurements. 4 40-41 mm. ; 2 40 mm.

108 D. Ss. PLETCHER

The subspecies is distinct in the male in the russet suffusion, the strong black
irroration and the strongly marked black pattern of the upper surface of the wings.

The single female is probably a striking aberration ; it has the medial area of the
fore wing black, edged proximally and distally with russet ; the proximal half of
the hind wing is black, edged distally with russet ; remainder of wings ochraceous
buff, irrorate and patterned with black and, in the apical area of the fore wing, with
russet (Pl. 13, fig. 377). Under surface of wings light buff, suffused and patterned
with fuscous black.

Of the type locality R. H. Carcasson of the Coryndon Museum writes: “ The
Malaba Forest was once part of the Kakamega Forest, which is in Western Kenya
on the escarpment above Lake Victoria. The altitude is from 5000-6000 ft.,
Malaba itself being nearer 5000 ft.

The Kakamega area has a very interesting relic Congo fauna and flora and many
of the birds are endemic races of Congo species ; the same pattern is repeated in
some of the small mammals and in numerous Lepidoptera. The distributional
pattern of your Cleora is by no means unusual.”

Distribution (Map 7). W. Kenya, Kakamega.

Holotype g. W. Kenya: [Kakamega], Malaba Forest, vi.1957 (C. R. Howard),
genitalia slide Geometridae No. 5360.

Paratypes : data as holotype, I g, I 9 in British Museum (Natural History),
2 6 in Coryndon Museum.

Cleora rothkirchi (Strand) comb. n.
(Text-figs. 144-146 ; Pl. 14, figs. 380-391 ; Map 8)

Boarmia rothkivcht Strand, 1914 : 44.
Boarmia acaciavia sensu Hampson, 1903 : 330.

36%. Colour and pattern of upper and under surfaces of wings vary geographically and are
described under subspecific headings ; all subspecies have discal spots fuscous to fuscous black
with smoke grey centres, contrasting sharply with respective medial areas.

3d. Genitalia (Text-figs. 144, 145). Uncus with thorn-like tip ; scobinate medial plate of
gnathus narrowly rounded ; apex of juxta Y-shaped ; sacculus dilate towards mid-valve in
apical third ; apex of sacculus minutely produced and sparsely setose ; vesica with two stout,
tapered cornuti fused at base, one five-eighths as long, one one-half as long as aedeagus, the
shorter tipped with a cluster of spines or spined at one side in apical eighth.

9. Genitalia (Text-fig. 146). Lamella postvaginalis weakly sclerotized medially ; colliculum
narrowed, anterior width equal to one-half of length, posterior margin sharply defined ; anterior
fourth of bursa copulatrix membranous, a little dilate at one side, remainder cylindrical, ribbed
and lightly sclerotized.

A species widely distributed throughout tropical Africa and the Mascarene region,
varying geographically in colour of vestiture and wings, and apparently without
especially close affinities with any other known species. In the male genitalia the
shape of the sacculus and the shape and proportions of the two cornuti, the shorter
one being scobinate at one side near apex, and in the female genitalia the shape of
the sterigma are diagnostic.

REVISION OF AFRICAN SPECIES OF CLEORA 109

Distribution (Map 8). Ivory Coast ; Ghana ; Nigeria ; Cameroun ; Angola ;
Congo (Leopoldville) ; Rwanda; Burundi; Uganda; Kenya; Tanzania ;
Mozambique ; Rhodesia ; Socotra ; Comoro Is ; Madagascar.

Cleora rothkirchi rothkirchi (Strand)
(Pl. 14, figs. 380-385 ; Map 8)

Boarmia rothkirvchi Strand, 1914 : 44.

6. Vestiture tilleul buff, vinaceous buff or pinkish buff and, except first abdominal segment,
irrorate with bister ; patagia edged with bister. Wings tilleul buff, vinaceous buff or pinkish
buff irrorate and patterned with bister or fuscous ; fore wing with a snuff brown to bister band

144

Fics. 144-146. C. vothkirchi genitalia. 144, aedeagus ; 145,43; 146, 9.

IIo DiS) PMEMGERE AR

proximad of antemedial fascia ; a similar band distad of postmedial fascia on each wing, these
bands weakly defined ; veins interruptedly light to warm buff distad of postmedial fasciae,
streak on vein M2 of forewing dilate to form spot (PI. 14, fig. 380). Under surface of wings white,
patterned with bister (Pl. 14, fig. 381).

9. Pattern similar to that of male but less strongly defined and dark irroration usually
sparse (Pl. 14, fig. 382). Underside white, tinged with light buff on veins and margins and pat-
terned with bister (Pl. 14, fig. 383).

Measurements. 4 33-38 mm. ; 2 34-40 mm.

The series includes forms comparable with tulbaghata abs. flavipleta and fumata ;
a male from W. Kenya has the proximal third of the fore wing and distal half of
each wing densely snuff brown.

Distribution (Map 8). Ivory Coast ; Ghana; Nigeria ; Cameroun ; Angola ;
Congo (Leopoldville) ; Uganda ; Kenya ; Tanzania ; Mozambique ; Rhodesia.

Material examined. Holotype 9. CAMEROUN: Duala, 22.x. (v. Rothkirch), in
Deutsches Entomologisches Institut.

Ivory Coast: Bingerville, 1915 (G. Melou), 29. Guana: N. Territorie Ketes,
Krachi (A. W. Cardinall), 2 9. Niceria: R. Niger, 7 miles S. of Baro, 7.1i1.1911
(G. B. Simpson), 1 9 ; Bonny, 3.1.1902 (Ansorge), 19. ANGOLA: Quirimbo, 75 km.
E. of P. Amboim, 300 m., 7—12.v.1934 (Dr. K. Jordan), 1 3, I 9, allin British Museum
(Natural History). Conco (Leopoldville) : Stanleyville a Coquilhatville, xi.1921
(L. Verlaine), 1 2 ; Yumbi [1°53"S. 16°34’E.], 1.x.1929 (A. J. Bredo), x 9 ;-Bolombo
[3°59"S. 21°24"E.], vii.1938 (J. Ghesquiére), 1 9 ; Rutschuru, 1.1928 (Ch. Seydel),
Ig; N. Kivu, Ngesho, ix.1937 (J. Ghesquiére), 1 29 ; Lusambo, vii,vii (Dr. M.
Fontaine), 3 9; Katanga, R. Lufira, 14.x.1925 (Ch. Seydel), 1 g ; Elisabethville,
iii,v (Ch. Seydel), 13, 2 9. Rwanpba: Kisenyi, 20.iv.1957 (Dr. M. Fontaine), i 9.
BurunvD1 : Usumbura, 900 m., vi,vii,viii (Dv. M. Fontaine), 3 9, all in Musée Royal
de l'Afrique Centrale. UGANDA: Kampala, 26.vii.1925, I g ; Entebbe (F. /.
Jackson), 3 9. Kenya: (G. W. Jeffery), 1 3 ; Kibwezi, v.1922 (W. Feather), 1 9.
Suna, Kavirondo, iv.1932 (W. Feather), 1 g. TANZANIA: Shinyanga, Mwandui,
11.1952 (Capt. Croft), 1 g. MozAMBIQUE: Lorenzo Marques (Distant coll.), 1 3g.
S. Ruopesta : Victoria Falls, rain forest, 22.vii.1949 (H. B. D. Kettlewell), 1 3.

Cleora rothkirchi amydropa ssp. n.
(Pl. 14, figs. 388-391 ; Map 8)
Boarmia acaciavia sensu Hampson, 1903 : 330.

39. Differs from the nominate subspecies and from subsp. zmsulavuwm in both sexes in the
generally grey and not pinkish buff or ochraceous buff colour of the vestiture and upper surface of
the wings (Pl. 14, figs. 388, 390). The under surface of the wings differs in being white,
patterned with fuscous black (Pl. 14, figs. 389, 391).

Measurements. 4 34-38 mm. ; 2 38-43 mm.

Forms comparable with abs. flavipleta and fumata of tulbaghata also occur in this
subspecies. Five specimens from the northern part of the Kenya coast are closely

REVISION OF AFRICAN SPECIES OF CLEORA ELE

similar to those of the type series from the island of Socotra and they have been
tentatively associated with this subspecies, though excluded from the type series.
Distribution (Map 8). Socotra ; ? N. Kenya Coast.

Holotype g. Socotra: Deneghan, 300 ft., 14.111.1953 (G. Popov), genitalia slide
Geometridae No. 2194.

Paratypes : data as holotype, 6 J, 3 2; Ahdo Dimellus, 3000 ft., 8—15.11.1899
(W. R. O. Grant), I 3, 2 &.

Other material. KENYA: Mombasa Island, I-7.x.1903 (Ff. J. Jackson), 1 3;
Mombasa, vi.1g916 (van Someren), 1 9 ; Diani, 15 mls. S. of Mombasa, iv.1953 (N.
Mitton), 13, all in British Museum (Natural History) ; Kenya coast, Shimba Hills,
xii.1961 (R. H. Carcasson), 1 2 ; Kenya coast, Gazi Forest, xii.1g61 (R. H. Carcasson)
I 9, both in Coryndon Museum.

Cleora rothkirchi insularum ssp. n.
(Pl. 14, figs. 386, 387 ; Map 8)

6. Differs from the nominate subspecies in both sexes in its smaller size and in the generally
more ochraceous colour of the upper surface of the wings (Pl. 14, figs. 386, 387).
Measurements. ¢ 30-35 mm.; 2 33-35 mm.

Five specimens from the Comoro Islands are provisionally associated with sub-
species insularum, but are excluded from the type series. One male from Anjouan,
a male from Grande Comore and a male from Moheli agree well with the type series
from Madagascar. The female from Mayotte (38 mm. wing-span) is similar to the
nominate subspecies and the second male from Anjouan is similar to a small subsp.
amydropa.

Distribution (Map 8). Madagascar ; ? Comoro Is.

Holotype g. Mapacascar: Diego Suarez, ii-iv (G. Melou), genitalia slide
Geometridae No. 2181.

Paratypes : MADAGASCAR: Diego Suarez, 2I-23.iv.1917, I gd, 4 2; ibid., 3.v.
1917, 14 ; ibid., 8-24.vu.1917, 14, 29; ibid., 23.viii.1g17, 1g ; Lambomakandro,
Sakaraha, Tulear, ili.1935 (R. Catala), 1 9 ; Mananjara, xi.1918, 6 J.

Other material. Comoro Is.: Anjouan I., 26—-30.vi.1g1r (G. F. Leigh), 1 3;
ibid., iv—vii.19g51 (Behagel), 1 § ; Grande Comore, 1884 (L. Humblot), 1 3, 1 2;
Moheli, Fomboni, 10 m., 1x.1958 (P. Griveaud), 1 g ; Mayotte, Chingoni, 70 m.,
x.1958 (P. Griveaud), I 9.

II2

D.-S. PFRETCHER

CHECK LIST OF SPECIES INCLUDED IN CLEORA

CLEORA, Curtis, 1825
Cerotricha Guenée, 1857
Barsine Meyrick, 1883
Meyrickia Butler, 1884
Chogada Moore, 1887
Carecomotis Warren, 1896
Neocleova Janse, 1932

cinctavia cinctavia (Denis & Schiffermiiller,

1775)
pascuaria (Brahm, 1791)

consimilavia (Duponchel, 1829)
cinctavia bowesi Richardson, 1952
cinctavia insolita (Butler, 1878)
sublunaria (Guenée, 1857)

tvansfixavia (Walker, 1860)

atvolinearia Hulst, 1888

aveataria Broadwell, 1907
projecta (Walker, 1860)

manitoba (Grossbeck, 1911) syn. n.

nigvofasciavia (Leech, 1897)
leucophaea (Butler, 1878)
elegans (Oberthiir, 1884)
pagina (Wileman, 1911)
venustaria (Leech, 1891)
alienaria alienaria (Walker, 1860)
alienaria gelidaria (Walker, 1863)
alienaria vasanaria (Swinhoe, 1915)
alienaria fumipennis Prout, 1929
fraterna (Moore, 1888)
determinata (Walker, 1860)
yakushimana (Inoue, 1956)
minutaria (Leech, 1891)
hermaea Prout, 1929
subbarbaria Prout, 1929
sevocata Prout, 1929
hospita (Prout, 1916)
ilustraria illustraria (Walker, 1863)
ulustvavia anestiayvia (Swinhoe, 1915)
tlustraria crina Prout, 1929
illustvavia aequivoca Prout, 1929
apista Prout, 1929
meceoscia Prout, 1929
mecistoscia Prout, 1929
paepalima West, 1932
tella (West, 1929)
pendleburyi Prout, 1929
xanthorrvhages Prout, 1929
myjoebergi Prout, 1926
neomenia Prout, 1932
vepetita (Butler, 1882)
inflexaria (Snellen, 1881)

epistictis (Meyrick, 1889)

proletavia (Swinhoe, 1915)
lipoteva West, 1920
processaria (Walker, 1860)

nigvonotavia (Wileman, 1911) comb. n.

injectavia tnjectavia (Walker, 1860)
sublectavia (Walker, 1863)
compactaria (Walker, 1863)

injectaria fuliginosa (Warren, 1894)

injectaria vittata (Warren, 1899)

injectavia dobboensis Prout, 1929

injectavia anidryta Prout, 1929

idiocvossa Turner, 1918

concentraria concentraria (Snellen, 1877)

? invalidaria (Snellen, 1895)
concentrarvia praia Prout, 1928
concentravia inobeda Prout, 1929
godeffroyi (Butler, 1886)
psectva Fletcher, 1957
samoana (Butler, 1886)
collenetter Prout, 1929
esotevica Prout, 1929

f. pusillanimis Prout, 1935
myyvmidonaria (Guenée, 1857)
leucostigma Prout, 1929
tongaica (Butler, 1886)
vitensis (B.-Baker, 1905)
psychastis (Meyrick, 1886)
immemorata (Walker, 1863)

lichenina (Butler, 1877)
munditibia Prout, 1927
stenoglypta Prout, 1929
licorvnaria Guenée, 1857
dodonaeae Prout, 1929
cheesmanae Prout, 1929
nausort (B.-Baker, 1905)
perstricta Prout, 1934
scviptaria (Walker, 1860)

panagrata (Walker, 1862)

stigmaticata (Walker, 1862)

menanaria (Walker, 1863)

antipodaria (Felder, 1874)

avenacea (Butler, 1879)

desiccata (Butler, 1879)
perlepidaria (Warren, 1900)
compectinata (Warren, 1906)
displicata (Walker, 1860)
pheucta Prout, 1937
goldfincht Prout, 1937
diphasia diphasia Prout, 1937
diphasia vefota Prout, 1937

REVISION OF AFRICAN SPECIES OF CLEORA 113

vhadia vhadia Prout, 1929
vhadia frigescens Prout, 1937
perbona Prout, 1937
lacteata (Warren, 1897)
decisarvia (Walker, 1866)
amphidoxa Prout, 1937
callicrossa (Meyrick, 1889)
nigristigma nigristigma Prout, 1937
nigvistigma talaseensis Prout, 1937
hemiopa hemiopa Prout, 1928
hemiopa quirvosi Prout, 1929
hemiopa ecdees Prout, 1929
ictuibasis Prout, 1937
cucullata cucullata (Fletcher, 1953)
cucullata fusconebulata (Fletcher, 1953)
monodactyla (Fletcher, 1953)
purissima (Fletcher, 1953)
batillata (Fletcher, 1953)
onycha onycha (Fletcher, 1953)
onycha amplissima (Fletcher, 1953)
onycha hastata (Fletcher, 1953)
onycha cultvata (Fletcher, 1953)
onycha sevvata (Fletcher, 1953)
onycha limitata (Fletcher, 1953)
onycha acuata (Fletcher, 1953)
taprobana (Fletcher, 1953)
kalist (Fletcher, 1953)
biclavata (Fletcher, 1953)
inornata (Fletcher, 1953)
fasciata (Fletcher, 1953)
olivata (Fletcher, 1953)
cornaria (Guenée, 1857)
invectavia (Walker, 1860)
properata (Walker, 1860)
vepulsaria (Walker, 1860)
propulsaria propulsaria (Walker, 1860)
propulsaria fieldi (Fletcher, 1953)
olivomaculata (Fletcher, 1953)
vostvata vostvata (Fletcher, 1953)
vostvata montliata (Fletcher, 1953)
falculata (Fletcher, 1953)
tenebrata tenebrata (Fletcher, 1953)
tenbrata acutiovata (Fletcher, 1953)
tenebrata avcuata (Fletcher, 1953)
tenebrata buruensis (Fletcher, 1953)
tenebrata fumata (Fletcher, 1953)
tenebrata parviorata (Fletcher, 1953)
inoffensa inoffensa (Swinhoe, 1902)
inoffensa cinereomarginata (Fletcher, 1953)
inoffensa glaucata (Fletcher, 1953)
inoffensa celebesa (Fletcher, 1953)
inoffensa minorata (Fletcher, 1953)
inoffensa exsilata (Fletcher, 1953)

indigna (Fletcher, 1953)
costiplaga costiplaga (Fletcher, 1953)
costiplaga flaviorata (Fletcher, 1953)
costiplaga ferrata (Fletcher, 1953)
costiplaga eichhorni (Fletcher, 1953)
costiplaga umbrata (Fletcher, 1953)
pupillata pupillata (Walker, 1860)
pupillata fuliginata (Fletcher, 1953)
pupillata luzonensis (Fletcher, 1953)
buxtoni (Fletcher, 1953)
perfumosa extendata (Fletcher, 1953)
perfumosa perfumosa (Warren, 1896)
braeckeli Debauche, 1941
forficulata (Fletcher, 1953)
levata (Fletcher, 1953)
albobvunneata albobyunneata (Fletcher, 1953)
albobrunneata cordata (Fletcher, 1953)
sabulata sabulata (Fletcher, 1953)
sabulata inconspicuata (Fletcher, 1953)
contiguata contiguata (Moore, 1867)
contiguata brooksi (Fletcher, 1953)
contiguata imparata (Fletcher, 1953)
contiguata bigladiata (Fletcher, 1953)
acaciaria (Boisduval, 1833)
tvansversavia (Pagenstecher, 1907)
betularvia (Warren, 1897)

funesta (Warren, 1905)
flavivenata sp. n.
melanochorda (Fletcher, 1958)
papillifer Prout, 1934
cancey sp. 0.
tamsi sp. Nn.
viettei (Herbulot, 1958)
oculata sp. n.
prosema Prout, 1927
anacantha sp. n.
epiclithva sp. n.
caycassoni sp. 0.
herbuloti herbuloti (Fletcher, 1958)
herbuloti phaea ssp. n.
acevata sp. N.
subcincta subcincta (Warren, 1901)
subcincta longifibulata (Fletcher, 1958)
lacrymata sp. n.
echinodes sp. n.
vaphis sp. n.
aculeata sp. n.
panarista sp. n.
quadrimaculata (Janse, 1932)
boetscht (Herbulot, 1961)
bicornis sp. 0.
dargei (Herbulot, 1961)
dactylata sp. n.

II4

thyvis sp. n.

nigvisparsalis (Janse, 1932)

tulbaghata (Felder, 1875)

plax sp. n.

munda (Warren, 1899)

lamottei (Herbulot, 1954)

toulgoetae (Herbulot, 1961)

vostella sp. 0.

legvasi (Herbulot, 1955)

angustivalvis (Herbulot, 1965)

sevena Sp. N.

oligodvanes (Prout, 1922)

macracantha (Herbulot, 1959)

devogaria (Snellen, 1872)
obsitaria (Mabille, 1890)

D. SS: FLETCHER

subspurcata (Warren, 1898)
pavlitzkiae pavlitzkiae (Fletcher, 1958)
pavlitzkiae lamella ssp. n.
pavlitzkiae etesiae ssp. n.
pavlitzkiae oviadelpha ssp. n.
pavlitzkiae saltuensis ssp. n.
lima sp. n.
scobina sp. n.
vadula vadula sp. n.
vadula leptopasta ssp. n.
vadula avenosa ssp. nN.
vadula eumelana ssp. 0.
vothkirchi vothkirchi (Strand, 1914)
vothkirvcht amydyropa ssp. 0.
vothkivcht insularum ssp. n.

SPECIES TO BE MOVED FROM CLEORA WHEN GENERA ARE AVAILABLE

Cleova albitrigonis Prout, 1927, Trans. ent. Soc. Lond. 75 : 195, pl. 20 : 13.
Cleova argicerauna Prout, 1929, Tveubia 7 : 448, pl. 9 : 14.
Cleova atriclava Prout, 1926, Novit. zool. 33 : 183.
Cleova bathyscia Turner, 1917, Proc. Linn. Soc. N.S.W. 42 : 371.
Cleova cnephaea Prout, 1915, Novit. zool. 22 : 359.
Cleova cryptogonia Prout, 1927, J. Bombay nat. Hist. Soc. 31 : 941, pl. 2 : 7.
Cleova discipuncta Joicey & Talbot, 1917, Ann. Mag. nat. Hist. (8) 20: 74, pl. 4: 8.
Cleova euboliaria (Walker), Turner, 1917, Proc. Linn. Soc. N.S.W. 42 : 372.
Tephrosia? euboliavia Walker, 1860, List Lep. Ins. B.M. 21 : 4109.
Cleova euplates Prout, 1925, Tvans. ent. Soc. Lond. 1925 : 315, pl. 36 : 26.
Cleora expleta Prout, 1932, Jl. fed. Malay St. Mus. 17 : 105, pl. 11 : 25.
Cleova fenestvata Prout, 1916, Novit. zool. 23 : 52.
Cleova flaccida (Warren), Joicey & Talbot, 1917, Ann. Mag. nat. Hist. (8) 20 : 74.
Alcis flaccida Warren, 1903, Novit. zool. 10 : 388.
Cleora flaccida constricta Joicey & Talbot, 1917, Ann. Mag. nat. Hist. (8) 20 : 74, pl. 4 : 16.
Alcis cinnamomea Rothschild, 1915, Lepidoptera Brit. Ornith. Union Exped. S. Dutch New
Guinea, 86.
Cleova gypsochroa Turner, 1947, Proc. R. Soc. Od 58 : ot.
Cleora hoplogaster Prout, 1916, Novit. zool. 23 : 51.
Cleova inaequipicta Prout, 1921, Bull. Hill Mus. Witley 1 : 150, pl. 18 : 14.
Cleova incompletavia (Guenée), Vinson, 1938, Bull. Maurit. Inst. 1(4) : 38.
Boarmia incompletaria Guenée, 1862, in Maillard, Notes sur l’tle de la Réunion, Annexe G, 27.
Cleova nesiotis Turner, 1926, Pap. Proc. R. Soc. Tasm. 1925 : 99.
Cleova ovygaria (Guenée), Vinson, 1938, Bull. Maurit. Inst. 1(4) : 38.
Neocleora ovygaria byunneata (Warren) Herbulot, 1956, Naturaliste malgache 8 : 252.
Xylopteryx brunneata Warren, 1912, Novit. zool. 9 : 523.
Cleova polymiges Prout, 1926, Novit. zool., 33 : 22.
Cleova proemia Prout, 1917, Novit. zool. 24 : 435.
Neocleova vefulgens Herbulot, 1965, Bull. Soc. ent. Fr. 69 : 258.
Boarmia (Cleova) rhamphoides Wehrli, 1943, in Seitz, Macrolepidoptera of the World 4, Suppl., 494.
Cleova yvussoi Prout, 1932, Rev. Zool. Bot. afr. 21 : 244.
Cleova scripta Joicey & Talbot, 1917, Ann. Mag. nat. Hist. (8) 20 : 74, pl. 3: 7.
Cleovra semidiscata (Warren) Warren, 1907, Novitt. zool. 14 : 172.
Chogada semidiscata Warren, 1906, Novit. zool. 13 : 141.

REVISION OF AFRICAN SPECIES OF CLEORA 115

Cleora subnigrata (Warren) Prout, 1927, J. Bombay nat. Hist. Soc. 31 : 942, pl. 2: 7.
Scotopteryx? subnigvata Warren, 1901, Novit. zool. 8 : 34.

Cleora tova Prout, 1926, Novit. zool. 33 : 183.

Cleora trigvapta Prout, 1927, J. Bombay nat. Hist. Soc. 31 : 940.

Cleora trisinuata Warren, 1898, Novit. zool. 5 : 248.

Cleova vatia Prout, 1927, J. Bombay nat. Hist. Soc. 31 : 941.

SPECIES MOVED FROM CLEORA TO OTHER GENERA

Cleora aargostigma Prout, 1927, J. Bombay nat. Hist. Soc. 31 : 939 moved to Alcis. Comb. n.
Cleora aeglophanes Prout, 1926, Sarawak Mus. J. 3(2) : 203 ; 1928, op. cit., pl. 16 : 9, moved to
Alcis. Comb. n.
Cleta albipunctaria Schaus, 1901, Trans. Am. ent. Soc. 27 : 241, treated by Prout as Cleora,
moved to Stellidia in the Noctuidae. Comb. n.
Cleova amictozona Prout, 1932, Novit. zool. 38 : 114 moved to Alcis. Comb. n.
Cleova antelmaria (Mabille) ; Vinson, 1938, Bull. Maurit. Inst. 1(4) : 38.
Boarmia antelmaria Mabille, 1893, Ann. Soc. ent. Belg. 37 : 64.
Chogada marmorata Warren, 1897, Novit. zool. 4 : 247 moved to Ascotis. Comb. n.
Cleova athola Prout, 1926, Novit. zool. 33 : 183 moved to Alcis. Comb. n.
Cleova bianquita Schaus, 1901, Tvans. Am. ent. Soc. 27 : 182 moved to Vinemina. Comb. n.
Cleova chionospila Turner, 1947, Proc. R. Soc. Qd 58 : 90 moved to Parathemis and placed in the
synonymy of Tephrosia externaria Walker, 1866. Syn. n.
Cleova cockaynei Prout, 1916, Novit. zool. 23 : 53 moved to Alcis. Comb. n.
Cleora colorifera Prout, 1916, Novit. zool. 23 : 53 moved to Alcis. Comb. n.
Cleora coniozona Prout, 1927, J. Bombay nat. Hist. Soc. 31 : 940, pl. 2 : 3 moved to Alcis. Gomb.
n.
Cleova decussata Moore, 1867, Proc. zool. Soc. Lond. 1867 : 628, pl. 33 : 4 moved to Alcis. Comb.
n.
Cleora derivata Prout, 1926, Sarawak Mus. J. 3(2) : 203 moved to Alcis. Comb. n.
Cleora dolichoptila Turner, 1947, Proc. R. Soc. Qd 58 : 91 moved to Chlenias. Comb. n.
Boarmia (Cleora) flavolinearia (Leech) ; Wehrli, 1954, in Seitz, Macrolepidoptera of the World 4,
Suppl., 720.
Boarmia flavolinearia Leech, 1891, Entomologist 24, Suppl., 47 moved to Alcis. Comb. n.
Boarmia (Cleorva) fortunata (Blachier) ; Wehrli, 1943, in Seitz, Macrolepidoptera of the World 4,
Suppl., 494.
Boarmia fortunata Blachier, 1887, Fewille jeun. Nat. 17 : 103.
Boarmia obscura B.-Baker, 1891, Trans. ent. Soc. Lond. 1891 : 218.
Boarmia wollastoni B.-Baker, 1891, Tvans. ent. Soc. Lond. 1891 : 217 moved to Ascotis.
Comb. n.
Boarmia buechlei Kilian, 1897, Societas ent. 12 : 41.
Cleora glaucotoxa Prout, 1927, Tvans. ent. Soc. Lond. 75 : 195, pl. 20: 12 moved to Ascotis.
Comb. n.
Cleova godmani Druce, 1892, Biologia cent.-am., Zool., Lep. Het. 2: 72, pl. 48 : 9 moved to
Semiothisa. Comb. n.
Cleova gracilis Warren, 1904, Novit. zool. 11 : 107 moved to Ividopsis. Comb. n.
Cleova hemichroma Turner, 1947, Proc. R. Soc. Qd 58 : 89 moved to Chlenias. Comb. n.
Cleova hemiphanes Prout, 1925, Novit. zool. 32 : 56 moved to Alcis. Comb. n.
Cleova hypopoecilia Prout, 1928, Bull. Hill Mus. Witley 2 : 151 moved to Alcis. Comb. n.
Cleova imbecilis (Moore) ; Prout, 1927, J. Bombay nat. Hist. Soc. 31 : 939.
Pseudocovemia imbecilis Moore, 1888, Descr. New Indian Lepidopterous Insects in coll.
Atkinson, 241 moved to Alcis. Comb. n.
Cleora ivrita Prout, 1928, Bull. Hill Mus. Witley 2 : 152 moved to Alcis. Comb. n.
Cleova ivvita f. (2sp.) obvuta Prout, 1928, loc. cit.

116 D. S: BEE TORE RK

Cleova latifasciata (Warren) ; Prout, 1927, J. Bombay nat. Hist. Soc. 31 : 938.
Poecilalcis latifasciata Warren, 1893, Proc. zool. Soc. Lond. 1893 : 427.
Boarvmia euvyzona Hampson, 1895, Fauna Br. India, Moths 3 : 281.
Boarmia eurydiscaria Hampson, 1902, J. Bombay nat. Hist. Soc. 14 : 507 moved to Alcis.
Comb. n.
Cleova megaspilavia Moore, 1867, Proc. zool. Soc. Lond. 1867 : 629 moved to Alcis. Comb. n.
Cleora nigridorsavia Guenée, 1857, Histoive naturelle des Insectes, Species généval des Lépidoptéres
9 : 232 moved to Alcis. Comb. n.
Cleova nigriscripta (Warren) ; Prout, 1929, Bull. Hill Mus. Witley 3 : 38, 51.
Poecilalcis nigriscripta Warren, 1903, Novit. zool. 10 : gor.
Cleora nigviscrvipta gavisa Prout, 1929, Bull. Hill Mus. Witley 3 : 38.
Cleova nigriscripta plenimedia Prout, 1929, Bull. Hill Mus. Witley 3 : 51 moved to Alcis.
Comb. n.
Cleova pachydesma Turner, 1947, Proc. voy. Soc. Qd 58 : 90 moved to Symmetroctena. Comb. n.
Cleova periphracta Prout, 1926, Savawak Mus. J. 3(2) : 202 moved to Alcis. Comb. n.
Cleova phaeocala Turner, 1947, Proc. R. Soc. Qd 58 : 91 moved to Chlenias. Comb. n.
Cleova praecisa Turner, 1917, Proc. Linn. Soc. N.S.W. 42 : 372 moved to Syneova. Comb. n.
Cleova praevariegata Prout, 1926, Savawak Mus. J. 3(2) : 202; 1928, 3(3), pl. 16 : 8 moved to
Alcis, Comb. n.
Cleova vufomarginata Moore, 1867, Proc. zool. Soc. Lond. 1867 : 628 moved to Alcis. Comb. n.
Boarmia (Cleora) scortea (Bastelberger) ; Wehrli, 1943, in Seitz, Macrolepidoptera of the World 4,
Suppl., 494.
Boarmia scortea Bastelberger, 1909, Ent. Z. Frank.a.M. 23 : 33 moved to Alcis. Comb. n.
Cleova semiochrea Prout, 1917, Novit. zool. 24 : 315 moved to Alcis. Comb. n.
Cleova semipullata Prout, 1925, Novit. zool. 32 : 57 moved to Alcis. Comb. n.
Cleova sericea (Warren); Prout, 1927, J. Bombay nat. Hist. Soc. 31 : 942.
Apophyga sericea Warren, 1893, Proc. zool. Soc. Lond. 1893 : 418 removed to A pophyga.
Cleora tevebvavia (Guenée) Vinson, 1938, Bull. Maurit. Inst. 1(4) : 38.
Hypopalpis terebraria Guenée, 1862, in Maillard, Notes sur Vile de la Réunion, Annexe G, 29.
Boarmia perfovavia Guenée, 1862, op. cit., p. 30, pl. 23 : 4.
Boaymia rousseli Oberthiir, Etudes Lép. comparées 7 : 275, pl. 169 : 1652, 1653 moved to
Ascotis. Comb. n.
Cleova variegata (Moore); Prout, 1927, J. Bombay nat. Hist. Soc. 31 : 938.
Pseudocoremia variegata Moore, 1888, Descr. New Indian Lepidopterous Insects in coll.
Atkinson, 240.
Cleova nebulosa Swinhoe, 1891, Trans. ent. Soc. Lond. 1891 : 488.
Cleova variegata convariata Prout, 1935, Novit. zool. 39 : 231 moved to Alcis. Comb. n.
Cleova versicolor Prout, 1915, Savawak Mus. J. 2(2) : 181 moved to Alcis. Comb. n.
Cleora venustulavria Walker, 1866, List Lep. Ins. B.M. 35 : 1579 moved to Alcis. Comb. n.

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118 D. SS. FEETCHER

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INDEX

Index to nominal genera and to Ethiopian species and subspecies of Cleova (synonyms in italics)

acaciaria Boisduval, 16 cancer sp. n., 28

acerata sp. n., 41 carcassoni sp. n., 36
aculeata sp. n., 49 Carecomotis Warren, 5, 6, 7
Alcis Curtis, 7 Catoria Moore, 8
amydropa ssp. n., 110 Cerotricha Guenée, 4
anacantha sp. n., 34 Chogada Moore, 5
angustivalvis Herbulot, 82 Cleora Curtis, 4

arenosa ssp. N., 107 Colocleora Prout, 7

Ascotis Hiibner, 7 Cusiala Moore, 8

Barsine Meyrick, 5

betularia Warren, 20 dactylata sp. n., 60
bicornis sp. n., 57 dargei Herbulot, 58
boetschi Herbulot, 54 derogaria Snellen, 89

INDEX

echinodes sp. n., 46 Ophthalmodes Guenée, 8
epiclithra sp. n., 36 oriadelpha ssp. n., 98
etesiae ssp. n., 94
eumelana ssp. n., 107
panarista sp. n., 51
papillifer Prout, 25

flavivenata sp. n., 22 Paradromulia Warren, 8
funesta Warren, 20 pavlitzkiae Fletcher, 92
phaea ssp. n., 40
plax sp. n., 71
herbuloti Fletcher, 38 prosema Prout, 33
Hypopalpis Guenée, 7 Pseudalcis Warren, 8
insularum ssp. n., III quadrimaculata Janse, 53
lacrymata sp. n., 44 radula sp. n., 104
lamella ssp. n., 94 raphis sp. n., 48
lamottei Herbulot, 73 rostella sp. n., 77
legrasi Herbulot, 80 rothkirchi Strand, 108

leptopasta ssp. n., 106

lima sp. n., 100

longifibulata Fletcher, 43 saltuensis ssp. n., 98
scobina sp. n., 10
Scotorythra Butler, 7

macracantha Herbulot, 87 serena sp. n., 84
melanochorda Fletcher, 24 subcincta Warren, 41
Meyrickia Butler, 5 subspurcata Warren, 91

munda Warren, 71

tamsi sp. n., 28

Neocleova Janse, 5, 6, 7 thyris sp. n., 62

nigrisparsalis Janse, 64 toulgoetae Herbulot, 76
transversaria Pagenstecher, 18
tulbaghata Felder, 67

obsitavia Mabille, 89

oculata sp. n., 31

oligodranes Prout, 86 viettei Herbulot, 30

PAA Ee

Cleora
Fic.
147. acaciavia 3 upperside B.M.Neg.35235
148. 3 a B.M.Neg.35223
149. 2 an B.M.Neg.35225
150. 6 underside B.M.Neg.35222
I51. 2 s B.M.Neg.35224

152. tvansversaria 3 upperside B.M.Neg.34875

153. 3} es B.M.Neg.34868
154. 3 si B.M.Neg.34877
155. 3} i B.M.Neg.34878
156. 3 nf B.M.Neg.34869
157. 3} 56 B.M.Neg.34873
158. 2 5 B.M.Neg.34874
159. 2 73 B.M.Neg.34871
160. Q 3 B.M.Neg.34876
161. 2 8 B.M.Neg.34872
162. Q B.M.Neg.34867
163. Q B.M.Neg.34870
164. 6 underside B.M.Neg.41170
165. Q a B.M.Neg.41169

Bull. Br. Mus. nat. Hist. Entomology, Suppt. 8 PAs fal Dy a

156 157 158 159

Fic.
166.
167.
168.
169.
170.
zal
172.
173.
174.
175.

PAM R, 2

Cleovra

betularia 3 upperside B.M.Neg.35238
3g underside B.M.Neg.35236
3 upperside B.M.Neg.35240
3 underside B.M.Neg.35239
3 upperside B.M.Neg.35241
3 underside B.M.Neg.35237
3 upperside B.M.Neg.35229
3 underside B.M.Neg.35232
3$ upperside B.M.Neg.35231
3g underside B.M.Neg.35227

Bull. Br. Mus. nat. Hist. Entomology, Suppt. 8 PLATE 2

Fic.
176.
OTe
178.
179.
18o.
18t.
182.
183.
184.
185.
186.
187.
188.
189.
19o.
IOl.

PLATE 3

Cleora

flavivenata paratype 3 upperside B.M.Neg.35228
rf 6 underside B.M.Neg.35230
i; 2 upperside B.M.Neg.35234
a 2 underside B.M.Neg.35233
melanochorda holotype 3 upperside B.M.Neg.39886
3¢ upperside B.M.Neg.38646A
Q - B.M.Neg.38647A
papillifer holotype 3 upperside B.M.Neg.38212A
melanochorda holotype 3 underside B.M.Neg.39887
2 underside B.M.Neg.38646B
Q 3 B.M.Neg.38647B
papillifer holotype 3 underside B.M.Neg.38212B
cancey paratype 2 upperside B.M.Neg.38211A
ys 2 underside B.M.Neg.38211B
a 2 upperside B.M.Neg.38210A
a 2 underside B.M.Neg.38210B

Bull. Br. Mus. nat. Hist. Entomology, Suppt. 8 PLATE 3

Fic.
192.
193.
194.
195.
196.
197.
198.
199.
200.
201.
202.
203.
204.
205.
206.
207.
208.
200.
210.

PLATE 4

Cleova

tamsi paratype g upperside B.M.Neg.38648A
¢ underside B.M. Neg.38648B
- 2 upperside B.M.Neg.38649A
2 underside B.M.Neg.38649B
. 2 upperside B.M.Neg.38650
vietter 3 upperside B.M.Neg.38651A
underside B.M.Neg.38651B
upperside B.M.Neg.38652
5 B.M.Neg.38653A
underside B.M.Neg.38653B
upperside B.M.Neg.38968
oculata holotype ¢ upperside B.M.Neg.38213A
Fn 6 underside B.M.Neg.38213B
paratype 2 upperside B.M.Neg.38214A
A 2 underside B.M.Neg.38214B
prosema 3 upperside B.M.Neg.38215
6 underside B.M.Neg.38212
anacantha holotype 3 upperside B.M.Neg.38216A
a4 6 underside B.M.Neg.38216B

+0 +0 +0 Os Oy Oy

Bull. By. Mus. nat. Hist. Entomology, Suppt. 8

FIG.
Ze,
PNP
Bish
214.
215.
210.
AMT
218.
219.
220.
22%
222%
PIR...
224.
225.
226.
PNT
228.
229.
230.

PLATE 5

Cleova

epiclithva holotype g upperside B.M.Neg.38217A
Re ¢ underside B.M.Neg.38217B
caycassoni paratype g upperside B.M.Neg.38218A
5 6 underside B.M.Neg.38218B
herbuloti herbuloti 3 upperside B.M.Neg.38219A
3 $f B.M.Neg.38321
3 ys B.M.Neg.38320
¢ underside B.M.Neg.38219B

herbulott phaea paratype g upperside B.M.Neg.38322A
* 3g underside B.M.Neg.38322B
_, 2 upperside B.M.Neg.38323A
= 2 underside B.M.Neg.383235

acevata paratype ¢ upperside B.M.Neg.38324
, 3 underside B.M.Neg.38325
lacyymata paratype 3 upperside B.M.Neg.38970
i 6 underside B.M.Neg.38971
vaphis holotype g upperside B.M.Neg.38974
y 6 underside B.M.Neg.38975
paratype 2 upperside B.M.Neg.38976
i ® underside B.M.Neg.38977

Bull. Br. Mus. nat. Hist. Entomology, Suppt. 8 PLATE 5

877

Fic,
Baie
222;
Baer
234.
235.
236.
P2eX af
238.
239.
240.
241.
242.
243.
244.
245.
246.
247.
248.
249.
250.

251

252.

PLATE 6

Cleova

subcincta subcincta 3 upperside B.M.Neg.38325A
2 4 B.M.Neg.38326A
3} B.M.Neg.38327A
subcincta longifibulata 3 paratype upperside B.M.Neg.38644A
3 upperside B.M.Neg.38328A
subcincta subcincta 3 underside B.M.Neg.38325B
Q a B.M.Neg.38326B
3 Ns B.M.Neg.38327B
subcincta longifibulata paratype g underside B.M.Neg.38644B
6 underside B.M.Neg.38328B
6 upperside B.M.Neg.38329A
echinodes holotype 3 upperside B.M.Neg.38972
aculeata paratype 3 upperside B.M.Neg.38978
Re Q a B.M.Neg.38980
subcincta longifibulata 2 underside B.M.Neg.38329B
echinodes holotype 3 underside B.M.Neg.38973
aculeata paratype 5 - B.M.Neg.38979
ne eo) “ B.M.Neg.38981
panarista paratype 3 upperside B.M.Neg.38982
ie 3 underside B.M.Neg.38983
5 2 upperside B.M.Neg.38984
7 © underside B.M.Neg.38985

Bull. By. Mus. nat. Hist. Entomology, Suppt. 8 PLATE 6

PLATE 7

Cleova

quadvimaculata 3 upperside B.M.Neg.38994
underside B.M.Neg.38995
upperside B.M.Neg.38998
underside B.M.Neg.38999
upperside B.M.Neg.38996
boetschi 3 upperside B.M.Neg.38986
6 underside B.M.Neg.38987
2 upperside B.M.Neg.38988
2 underside B.M.Neg.38989
bicovnis paratype 3 upperside B.M.Neg.38990
a 6 underside B.M.Neg.38q91
F 2 upperside B.M.Neg.38992
* 2 underside B.M.Neg.38993
dargei 3 upperside B.M.Neg.39000
6 underside B.M.Neg.39001
2 upperside B.M.Neg.39002
2 underside B.M.Neg.39003
dactylata paratype 3 upperside B.M.Neg.39004
A 3g underside B.M.Neg.39005
- 2 upperside B.M.Neg.39006
2 underside B.M.Neg.39007

+0 +0 40 Oa

Bull. Br. Mus. nat. Hist. Entomology, Suppt. 8 PIRATE 7

Fic.
274.
275.
276.
277
278.
279.
280.
281.
282.
283.
284.
285.
286.
287.
288.

PLATE 8

Cleova

thyvis holotype 3 upperside B.M.Neg.39773
is 3 underside B.M.Neg.39774
paratype 2 upperside B.M.Neg.39777
ts 2 underside B.M.Neg.39778
3 3 upperside B.M.Neg.39776
2 r B.M.Neg.39779
“ie ot r» B.M.Neg.39775
nigvisparsalis 3 upperside B.M.Neg.39793
3 underside B.M.Neg.39794
2 upperside B.M.Neg.39795
2 underside B.M.Neg.39796
35 upperside B.M.Neg.39798
2 r B.M.Neg.39797
plax holotype 3 upperside B.M.Neg.39787
i 3$ underside B.M.Neg.39788

Bull, Br. Mus. nat. Hist. Entomology, Suppt. 8 PEATE

Fic.
289.
290.
291.
292.
293.
294.
295.
296.
297.
208.
299.
300.
301.
302.
303.
304.

PLATE 9

Cleova

tulbaghata 3 upperside B.M.Neg.39780
§ underside B.M.Neg.39781
2 upperside B.M.Neg.39785
2 underside B.M.Neg.39786
ab. flavipleta 3 upperside B.M.Neg.39783
ab. fumata 4 upperside B.M.Neg.39782
§ upperside B.M.Neg.39784
holotype 3 upperside B.M.Neg.41281
munda 3 upperside B.M.Neg.39789
$ underside B.M.Neg.39790
2 upperside B.M.Neg.39791
2 underside B.M.Neg.39792
tulbaghata 2 upperside B.M.Neg.41284
munda holotype 2 upperside B.M.Neg.41172
scobina paratype 2 upperside B.M.Neg.41282
2 underside B.M.Neg.41283

Bull. Br. Mus. nat. Hist. Entomology, Suppt. 8 PLATE 9

Fic.
305.
300.
307.
308.
309.
310.
311.
312.
313.
314.
315.
316.
317-
318.
319.
320.
321.
Bees
B23:
324.

PLATE to

Cleova

lamottei 3 upperside B.M.Neg.39881
6 underside B.M.Neg.39882
2 upperside B.M.Neg.39883
2 underside B.M.Neg.39884
toulgoetae 3 upperside B.M.Neg.39862
¢ underside B.M.Neg.39863
2 upperside B.M.Neg.39864
2 underside B.M.Neg.39865
vostella paratype 3 upperside B.M.Neg.39799
> 6 underside B.M.Neg.39800
Fi 2 upperside B.M.Neg.39801
A 2 underside B.M.Neg.39802
oF 2 upperside B.M.Neg.39803
Fr Oe B.M.Neg.39804
3 3g underside B.M.Neg.39805
lamotter 3 upperside B.M.Neg.39885
angustivalvis 3 upperside B.M.Neg.39840
¢§ underside B.M.Neg.39841
2 upperside B.M.Neg.39842
2 underside B.M.Neg.39843

Bull. By. Mus. nat. Hist. Entomology, Suppt. 8 PLATE, to

Fic.
325.
3206.
BS 27h
328.
329.
330.
331.
332.
333:
334-
335:
3306.
337:
338.
339.
340.
341.
342.
343:
344.

PLATE it

Cleova

legvast § upperside B.M.Neg.39835
6 underside B.M.Neg.39836
2 upperside B.M.Neg.39837
2 underside B.M.Neg.39838
6 upperside B.M.Neg.39839
macracantha 3 upperside B.M.Neg.39830
6 underside B.M.Neg.39831
6 upperside B.M.Neg.39834
Q A B.M.Neg.39832
2 underside B.M.Neg.39833
scobina paratype ¢ upperside B.M.Neg. 39860
Pa 6 underside B.M.Neg.39861
sevena paratype g upperside B.M.Neg.39810
a 6 underside B.M.Neg.39811
FP 2 upperside B.M.Neg.39812
is 2 underside B.M.Neg.39813
oligodvanes 3 upperside B.M.Neg.39814
¢ underside B.M.Neg.39815
2 upperside B.M.Neg.39816
9 underside B.M.Neg.39817

Bull. By. Mus. nat. Hist. Entomology, Suppt. 8 PAE se

Fic.

345-
346.
347-
348.
349.
350.
351.
352.
353:
354-
355:
356.
357:
358.

359.
360.

361.
362.
363.
364.

PLATE 12

Cleova

devogaria 3 upperside B.M.Neg.39806
3 underside B.M.Neg.39807
9 upperside B.M.Neg.39808
2 underside B.M.Neg.39809
pavlitzkiae pavlitzkiae holotype 3 upperside B.M.Neg.39844
. 3 underside B.M.Neg.39845
paratype 2 upperside B.M.Neg.39846
* underside B.M.Neg.39847
3 upperside B.M.Neg.39848
fe) i B.M.Neg.39849
pavlitzkiae lamella paratype 3 upperside B.M.Neg.39850
pA 6 underside B.M.Neg.39851
pavlitzkiae oriadelpha paratype 3 upperside B.M.Neg.39852
5 ¢ underside B.M.Neg.39853
pavlitzkiae saltuensis holotype 3 upperside B.M.Neg.39854
5 6 underside B.M.Neg.39855
paratype 2 upperside B.M.Neg.39856
5 2 underside B.M.Neg.39857
lima paratype 3 upperside B.M.Neg.39858
* 3 underside B.M.Neg.39859

Bull. Br. Mus. nat. Hist. Entomology, Suppt. 8

PLATE 12

Fic.
3605.
366.
307.
368.
369.
379.
371.
372:
373:
374:
375:
376.
377:
378.
379:

IPIL NILE, 305}

Cleova

vadula vadula paratype 3 upperside B.M.Neg.39868
- 6 underside B.M.Neg.39869
- 2 upperside B.M.Neg.39871
‘5 2 underside B.M.Neg.39872
Pp 3 upperside B.M.Neg.39870
vadula leptopasta holotype 3 upperside B.M.Neg.39866
nf 3g underside B.M.Neg.39867
vadula avenosa paratype 3 upperside B.M.Neg.39875
Fe 3 5 B.M.Neg.39873
nf 6 underside B.M.Neg.39874
is 2 upperside B.M.Neg.39876
. 2 underside B.M.Neg.39877
vadula eumelana paratype 2 upperside B.M.Neg.39878

a 3 ee B.M.Neg.39879
se 6 underside B.M.Neg.39880

Bull. By. Mus. nat. Hist. Entomology, Suppt. 8 PLATE 13

S9t

99€

Fic.
380.
381.
382.
383.
384.
385.
386.
387.
388.
380.
399.
391.

PEATE or

Cleova

vothkivcht vothkivcht 3 upperside B.M.Neg.39818

6 underside B.M.Neg.38919

2 upperside B.M.Neg.39820

2 underside B.M.Neg.39821

2 upperside B.M.Neg.39822

2 underside B.M.Neg.39823
vothkivcht insularum holotype g upperside B.M.Neg.39824
As 6 underside B.M.Neg.39825
vothkivcht amydvopa paratype 3 upperside B.M.Neg.38926
o underside B.M.Neg.39827
2 upperside B.M.Neg.39828
® underside B.M.Neg.39829

”

”

”

Bull. By. Mus. nat. Hist. Entomology, Suppt. 8 PLATE 14

G8e
18€

C8E

L8E
€8€

4

y acaciaria

© betularia

a Cancer

# flavivenata

ao melanochorda
a papillifer

e transversaria

N 4 a4
\ ~S4
Sete
a re | eae
| °
\ |4 x |
5 hey, ]

. - y
\ = / = =
\ Ger Saas #
) o UF

Map ft.

Distribution of Ethiopian Cleova, species-group acaciaria

s anacantha
y carcassoni
e epiclithra
a oculata

= prosema

Map 2. Distribution of Ethiopian Cleova, species-group oculata

\

4 aculeata

e echinodes

4 lacrymata

o panarista

vy quadrimaculata
o raphis

v boetschi

Map 3. Distribution of Ethiopian Cleova, species-group vaphis

a dactylata

e darge@i

’ munda

o plax

ja thyris

» tulbaghata

o nigrisparsalis

Map 4.

Distribution of Ethiopian Cleora, species-group tulbaghata

# angustivalvis
= derogaria

/vlamottei
i :
o legras|

' a macracantha

a oligodranes

| v~rostella
e scobina

-eserena

0 toulgoetae

Map 5. Distribution of Ethiopian Cleova, species-group rostella

i | \
\

(e pavlitzkiae pavlit zk iae
opavlitzkiae lamella
epavlitzkiae ® lamella
apavlitzkiae etesiae
epavlitzkiae oriadelpha
vy pavlitzkiae saltuensis
'e pavlitzkiae ? saltuensis

Map 6. Distribution of Ethiopian Cleova, species-group vostella, species pavlitzkiae

-¢radula ? radula

_eradula |eptopasta
-eradula ’?leptopasta

\

_afadula arenosa
>a rddula eumelana

Distribution of Ethiopian Cleorva, species-group rostella, species vadula

Map 7.

‘a rothkirchi tothkirchi
ia rothkirchi insularum
“e rothkirchi ? insularum
_erothkirchi amydropa

_erothkirchi ?amydropa
'olima

Map 8. Distribution of Ethiopian Cleova, species-group rostella, species vothkirchi and lima

o Acerata

aherbuloti herbuloti

| aherbuloti phaea

| |e subcincta subcincta

lo subcincta longifibulata
vtamsi

vy viettei

| = bicornis

Map 9g. Distribution of arbitrarily placed species of Ethiopian Cleova

hy
|
Lm!
f
Pte
-
a
=
,
i
J
ie
*
.
Mi
t
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aa

“

A’ LIST OF SUPPLEMENTS
TO <THE ENTOMOLOGICAL SERIES
OF THE . _BULEETIN: | OF
THE BRITISH MUSEUM (NATURAL HISTORY)

. Masner, L. The types of Proctotrupoidea (Hymenoptera) in the British
Museum (Natural History) and in the Hope Department of Entomology, Oxford.
Pp. 143. February, 1965. 5.

. Nrxon, G. E. J. A reclassification of the tribe Microgasterini (Hymenoptera :
Braconidae). Pp. 284: 348 Text-figures. August, 1965. {6.

. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177 :
18 plates, 270 Text-figures. August, 1965. {4 4s.

. SaAnDs, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera,
Termitidae) from the Ethiopian Region. Pp. 172 : 500 Text-figures. September,
1965. £355.

. AHMAD,I. The Leptocorisinae (Heteroptera : Alydidae) of the World. Pp. 156:
475 Text-figures. November, 1965. {2 I5s.

. OxaDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso-
philidae. Pp. 129: 328 Text-figures. May, 1966. £3.

. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 Text-figures. January,
T9Q67, 63°35.

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