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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 



f 



MORPHOLOGY AND TAXONOMY 
OF ADULT MALES OF THE 

FAMILY COCCIDAE 
(HOMOPTERA : COCCOIDEA) 




BY 



J. H. GILIOM EE 

Dept. of Entomology, University of Stellenboscli} South Africa 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Supplement 7 

LONDON : 1967 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. 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 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 20th January, 1967 Price £3 3s. 



MORPHOLOGY AND TAXONOMY 
OF ADULT MALES OF THE 

FAMILY COCCIDAE 
(HOMOPTERA : COCCOIDEA) 

By J. H. GILIOMEE* 



I 'INTENTS 

Acknowledgements 

Introduction 
Review of the Literai i r] 
Material and Technique 
Key to Lettering on Figures 

Abbreviations used in the I 
General Morphology 

General Characteristics 
The Head 

I le id Capsule 
Antennae . 
The Thorax 

1 Yothorax . 
Mesothorax 
Metathorax 
Wings and Halteres 
Legs . 
The Abdomen . 

I'regenital Segments 

Genital Segment and External Genitalia 
Description of the Species 
The EULECANIUM Grou 
The ERIOPELTIS Group 
The INGLISIA Croup 
The COCCUS Group . 
Discussion 

Taxonomic Significance of the Characters 
Classification and Interrelationships of the 

Males 

Groups of Genera 

Genera ..... 
Species ..... 
Relationships of the Coccidae with the other subdivisions 
Coccoidea 
Keys .... 
References . 
Index .... 

* The information contained in this work was submitted in the form of a thesis for 
Doctor of Philosophy in the University of London, June, 1964. 



Coccidae based on the 



of the 



Page 

4 

4 

5 

7 

9 

9 
1 1 
12 
14 
14 
19 
-'i 
21 

23 

28 

30 
3i 
33 
33 
36 
37 
37 
77 
88 
92 

125 
126 

L39 
139 
146 

L5i 

153 
159 
164 
168 

the degree of 



4 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

SYNOPSIS 

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 Would 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. Mitic- 
Muzina, Institute for Plant Protection, Belgrade ; Mr. J. Munting, Plant Protect- 
ion Research Institute, Pretoria ; Dr. J. Rehacek, 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, 1941 ; Bodenheimer, 1953) indicated the close relationship of a small number 



OF THE FAMILY COCCI DAK 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 ; (ii) 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 Xewstead's (1901, 190.5) 
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 (1919a, 1919b, 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) : Aspidiotits hederae (Vail.), Lichtensia viburni 
Signoret, Aclerda berlesii Buffa, Pseudococcus citri (Risso), Eriococcus anricariae 
Mask., Micrococcus silvestrii Leon., Trabutina leonardii 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 Eulecanium 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 (i960) 
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 hesperidnm. 
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, Parthenolecaniam pomeranicum 
(Raw.). 

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 (i960). 
However, in view of Wilkey & McRenzie'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 Runckel d'Herculais. Husseiny & Madsen 
(1962) dealt with the life history of Lecanium kunoensis Ruw. 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 Steingelia. 

MATERIAL AND TECHNIQUE 

The males of 23 species, belonging to 19 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 



Eidecanium 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. bi tuber cidatum (Targioni Tozetti, 
1868). 



Phyllostroma Sulc, 1942. 

P. myrlilli (Kaltenbach, 1874). 
Filippia Targioni Tozetti, 1868. 

F. viburni (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 



Eriopeltis Signoret, 1871. 
E. species. 
E. Ifcstucae (Fonscolombe, 1834). 



Luzidaspis Cockerell, 1902. 
L. luzulae (Dufour, 1864). 



The INGLISIA Group 

Inglisia Maskell, 1879. 

/. thcobromae Newstead, 1917. 



The COCCUS Group 



Coccus Linnaeus, 1758. 

C. hesperidum Linnaeus, 1758. 
Genus B species (nr. Pidvinaria). 
Pulvinaria Targioni Tozetti, 1868. 

P. ?betulae (Linnaeus, 1758). 

P. acericola (Walsh & Riley, 1868). 



Parthcnolecanium Sulc, 1908. 

P. corni (Bouche, 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 INGLISIA group only one species (/. 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. Rehacek 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. 1). 

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 prescutum 



OF THE FAMILY COCCIDAE g 

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 ridges. The maximum 
length and width of the median membranous area of the scutum, the length and width 
across the middle of the scutelhim, 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 


1 >orsal and ventral aspects of body 


1. 


Hind claw, posterior view 


A 1 


Lateral aspect of body 


M 


Part of inner margin of fore coxa, 


B 


Head, dorsal view 




showing setae and coxal bristle(s) 


C 


Head, ventral view 


N 


Articulation of fore wing, showing 


D 


Mesoprephragma, anterior view 




pteralia 


E 


Mesopostphragma, posterior view 


() 


3rd axillary wing sclerite, posterior 


F 


3rd segment of left antenna, 




view 




ventral view 


1' 


Subalare, dorsal view 


G 


10th segment of left antenna. 


Q 


Caudal extension of 8th abdominal 




ventral view- 




segment, dorsal view 


H 


Membranous area <>l st utum 


K 


Apex of penial sheath, ventral 


I 


3rd axillary wing sclerite, dorsal 




view 




view 


S 


Apex of penial sheath, ventral 


J 


Fore claw, posterior view- 




view 


K 


Middle claw, posterior view 


T 


Tentorium and cranial apophysis 




ABBREVIATIONS l 


SED 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 



apar 


anterior postalar ridge 


mc 


median crest 


as 


abdominal sternite 


mdr 


median ridge 


asc 


additional sclerite 


med 


media 


ase 


apical seta 


mo 


mouth opening 


astrijS 


anteprosternal seta 


mpns 


medial pronotal setae 


at 


abdominal tergite 


mr 


marginal ridge 


ata 


anterior tentorial arm 


mts 


metatergal setae 


atp 


anterior tentorial pit 





ocellus 


avs 


abdominal ventral setae 


ocs 


ocular sclerite 


ax 1 


first axillary wing sclerite 


pa 


postalare 


ax g 


second axillary wing sclerite 


pcr 2 


precoxal ridge of mesothorax 


ax 3 


third axillary wing sclerite 


pcr 3 


vestigial precoxal ridge of meta- 


bas 


basalare 




thorax 


bma 


basal membranous area 


pdc 


pedicel 


bra 


basal rod of aedeagus 


pepcv 


proepisternum + cervical 


bs 


sensilla basiconica 




sclerite 


c 


cicatrix 


pla, 


propleural apophysis 


ca 


cranial apophysis 


pla 2 


mesopleural apophysis 


cb 


coxal bristle(s) 


pla 3 


vestigial metapleural apophysis 


ccx 


costal complex of wing veins 


plTj 


propleural ridge 


ce 


caudal extension 


P lr 2 


mesopleural ridge 


cl 


claw 


P lr 3 


metapleural ridge 


ex 


coxa 


pms 


postmesospiracular setae 


dhs 


dorsal head setae 


pmss 


posterior metasternal setae 


dmcr 


dorsomedial part of midcranial 


pn 2 


mesopostnotum 




ridge 


P n s 


metapostnotum 


dos 


dorsal ocular setae 


pna 


postnotal apophysis 


dps 


dorsopleural setae 


pnp 


posterior notal wing process 


dse 


dorsal simple eye 


pnr 


pronotal ridge 


dss 


dorsospiracular setae 


pocr 


postocular ridge 


epm 2 


mesepimeron 


ppar 


posterior postalar ridge 


epm 3 


metepimeron 


pra 


prealare 


eps 2 


mesepisternum 


prn 


lateral pronotal sclerite 


eps 3 


metepisternum 


prnr 


pronotal ridge 


eps 3 s 


postmetaspiracular setae 


procr 


preocular ridge 


f 


furca 


pror 


preoral ridge 


Fiii- X 


segments of flagellum, 3rd to 


prsc 


prescutum 




10th 


ps 


penial sheath 


fm 


femur 


pscr 


prescutal ridge 


fp 


furcal pit 


pscs 


prescutal suture 


fs 


fleshy seta 


Pt 


post-tergite 


g 


gena 


pta 


posterior tentorial arm 


gls 


seta of glandular pouch 


ptp 


posterior tentorial pit 


gp 


glandular pouch 


ptr 2 


peritreme of mesothoracic 


gs 


genal setae 




spiracle 


gts 


setae of genital segment 


Ptr 3 


peritreme of metathoracic 


h 


haltere 




spiracle 


hs 


hair-like seta 


pts 


post-tergital setae 


ior 


interocular ridge 


pwp a 


mesopleural wing process 


lmcr 


lateral branch of midcranial 


Pwp 3 


vestigial metapleural wing pro- 




ridge 




cess 


lpl 


lateropleurite 


rad 


radius 


Ise 


lateral simple eye 


sa 


subalare 



OF THE FAMILY COCCIDAE 



set. scla 


subapical seta 


t 


tendon-like apodeme 




scl 


seutellum 


tar 


tarsus 




self 


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 




sp 2 


mesothoracic spiracle 


tp 


triangular plate 




sp 3 


metathoracic spiracle 


tr 


trochanter 




spl 


sensillum placodeum 


udgt 


ungual digitule 




ss 


suspensorial sclerite 


vhs 


ventral head setae 




Stllj 


prosternum 


vmcr 


ventral part of midcranial 


ridge 


stn.. 


basisternum <>! mesosternum 


vps 


ventropleural setae 




stn :) 


metasternum 


vs 


ventral sclerite 




stn,s 


prosternal setae 


vse 


ventral simple eye 




stn 2 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 
seutellum (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 (1919a, 1919b, 
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), Sulc (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 10th antennal segments, seutellum 
and terminal abdominal segments (including the genital segment) in the species that 
he studied. A small number of workers, namely Pesson (1941), Durr (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 taxi) 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 terminologv 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 \l long) 
and the largest E. pela (2500-3100, average 2864 jj. long) ; L. luzulae had the shortest 
wing-span (2090-2350, average 2213 \l) and E. pela the longest (5330-5700, average 
55°3 V-)- Some species are slender in appearance (e.g. C. hesperidum) while others 
are rather robust (e.g. Ceroplastes 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 EULECANIUM 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. iB, 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 antero ventral 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. 



OF THE FAMILY COCCIDAF 



'3 



?%^S^ 




Fig. i. Generalized drawing of the male of the Coccidae. 



14 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

The abdomen is parallel-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. E. 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 (/. 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 ridge. 



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 posteriori}- into the ocular sclerite, but in some cases (/. 
theobromae, Text-fig. 29, C ; the ERIOPELTIS 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 (/. theobromae, Text-fig. 29, C), short in others 
(Eriopeltis sp., Text-fig. 24, C), or even absent (E. Pfestucae, 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-tig. 
20, C) or the area is both reticulated and sclerotized {Eriopeltis 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 ERIO- 
PELTIS group, Ctenochiton sp., E. pda, 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 (1919a, 1919b, 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 I. 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. E. tiliae, 
Text-fig. 3) or flat (e.g. L. luzulae, Text-fig. 28). The lateral eyes (lse) 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 (I. 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 (E. tiliae, F. viburni, E. pela ; Text-figs. 2, 14, 18) additional lateral eyes. 
The lateral ocellus (o) usually appears as a weakly sclerotized spot on a membranous 
bulge, which is somewhat conical in Eriopeltis 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. corni were studied by 
Pflugf elder (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. Eriopeltis 
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 
ERIOPELl IS group, Genus A ; Text-figs. 24, 26, 27 and 20 ; C). 

The preoral ridge gives support to the cranial apophysis (ca), which is a strong 
scoop-like structure (Text-figs. 24, 39 ; T). Its apex is usually bifurcate, but 
sometimes trifurcate (Eriopeltis sp., Text-fig. 24, C) or truncate (E. pela, E. Pfestucae, 
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 /:. 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 Pseiidaspidoproctits PfuUeri. 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. In a 
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 posterior tentorial pits (ptp) are found posterolateral 
to the mouth opening, immediately median to the posterior ends of the postocular 
ridges ; they are always present. From each posterior pit a thread-like posterior 
tentorial arm (Text-figs. 24, 39 ; T ; 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 

(ii) 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 EULECANIUM and ERIOPELTIS 
groups) or of both fleshy and hair-like ones (the INGLISIA and COCCUS groups). 
The number of hair-like setae, which are present in all the species, varies from 1-7 
(average 5) in P. acericola to 16-19 (average 17) in P. corni ; the number of the 
additional fleshy setae, in the species in which they occur, varies from 4-1 1 (average 
7) in /. 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. 

(iii) 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. prunastri, L. luzulae; Text-figs. 
20, 22, 27 ; C) one pair of median hair-like 3etae 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 COCCI DAE 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 9-16 (average 12) in Eriopeltis sp. to 
54-108 (average 84) in Pulvinaria Pribesiae. 

(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 /. theobromae to 7-13 (average 9) in P. conii, and the number 
of fleshy setae large, varying from 5-1 1 (average 7) in /. theobromae to 17-30 (average 
23) in P. corni. 

No other dermal structures, like disc pores or specialized sensilla are present on 
the head. 

A ntennae 

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 niidcranial 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 9th are intimately fused. The 
relative length of the antennae varies considerably within the family. They are 
very long in the ERIOPELTIS group, being about §-£ as long as the body in 
Eriopeltis 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 /:. tiliae 
(EULECANIUM group) where they are J-f as long as the body. The average 
absolute length varies from 622 [i in C. berliniae to 1922 \i 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 10th 
segments, but in E. pela they occur regularly on the 1st 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. E. tiliae, 

F. viburni, 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 10th 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 10th segment there also occur the long, 



20 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

capitate, subapical setae (set. scla) found in the Pseudococcidae (Sulc 1943 ; 
Giliomee, 1961) and Diaspididae (Bustshik, 1958 ; Ghauri, 1962). Sulc called them 
setae semi-claviformes. Both types probably have a sensory function. Specialized 
sensilla are present on the 2nd, 3rd and 10th 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. Pbetulae (Text-fig. 35, B) and I. 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 (Fujux) is composed of the 3rd to 10th segments. The 3rd segment 
(Text-figs., F) varies considerably in size and shape ; in Eriopeltis spp. (Text- 
figs. 24, 26 ; F) it is long and club-shaped, while it is short and barrel-shaped 
in P. Pbetulae (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 9th 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 10th (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 /. theobromae there are only 2 and in Genus A there are 4-6 
(average 5). On the ventral surface of the 10th 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 COCCIDAK 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 x ). Crampton (1926) called the structure a " neck plate " or 
" laterocervicale " 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 (stnj 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. hesperidum ; Text-figs. 12, 31), interrupted in the middle (e.g. E. tiliae, E. 
Pfestucae, P. corni ; Text-figs. 2, 26, 38), developed anteriorly only (e.g. Eriopeltis 
sp., P. pomeranicum ; 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. Pbetulae the ridge may be complete 
or developed anteriorly only, while in P. bituberculatum and S. prunastri 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 presternum. 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 (/. 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 (lpns), 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. prunastri, 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. theobromae) one or both setae may be absent, 
while medial pronotal setae are absent altogether in some species (e.g. E. 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). 

(iii) 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 (astnjs) consisting of a number of fleshy setae (up to 7) 
which occur immediately ventral to the anterior part of the prosternum 4- cervical 
sclerite (pepcv). They are present in the COCCUS group (Text-figs. 31-43) and 
sometimes in L. luzulae. 

(v) Prosternal setae (stn^), 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 > 3 

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 B 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-i), 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. 

Mesotergam. 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. E. Pfestitcae, 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. bituberadatum, 
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. hesperidum, 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 scutum (set) 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 triangular 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 ; posterolateral^ 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. pomeranicum. 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 (Eriopeltis 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 (self) (F. viburni, Ctenochiton 
sp., S. prunastri, 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 (/. 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 (pn 2 ) 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. tiliae, 
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 postulate (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 2 ). 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 (pla 2 ) is invaginated. In some species (e.g. F. vibumi, 
Ctenochiton sp., E. pcla ; Text-rigs. 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 2 ) 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 subalare (Text-figs. 1, A 1( 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) is a 
narrow but distinct sclerite (in the EULECANIUM, 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 (i960). The 
episternum (epS2) 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 sitbepisternal 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 



26 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

by a band of darker sclerotization. The later opieurite (lpl) is always well developed 
and the anterior margin often bounded by an extension from the anterior part of 
the marginal ridge (e.g. E. tiliae, L. luzulae, I. theobromae ; Text-figs. 2, 27, 29). The 
epimeron (epm 2 ) is represented by a small sclerite posterodorsal to the coxal articula- 
tion. The mesothoracic spiracle (sp 2 ), with its supporting peritreme (ptr 2 ), 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 (i960). 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 2 ). 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 (pcr2) 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 (Eriopeltis spp. ; Text-figs. 24, 26) or 
vestigial (/. theobromae ; Text-fig. 29). The posterior margin of the basisternum 
is invaginated to form a transverse fur col 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 " f urea- 
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 x ) 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. 1 ; 18, N ; ax 2 ) 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 lies directly below it. The 
structure of the third axillary sclerite (Text-figs. 1 ; 18, N ; ax 3 ) 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. Pfestucae) 
they also extend beyond the posterolateral corner of the membranous area to occur 
on the sclerotized parts. In a number of species (INGLISIA 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 B). In the other 
species the scutal setae are either absent (E. 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. Pulvinaria spp.). 



28 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

(ii) 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 tegular setae (tegs) are carried on the anterior part of the tegular bulge and 
consist usually of a small number (up to n) of hair-like setae. Fleshy tegular setae 
were only observed in Ceroplastodes chiton Green and occasionally in /. 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. corni. 

(v) The basisternal setae (stn 2 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. E. tiliae, F. viburni, 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 it may be absent or present within the 
same species. In the intersegmental region between the metathorax and 1st ab- 
dominal segment there is an irregular, transverse, lateral sclerite which corresponds 
to the acrotergite or postnotum (pn 3 ) 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 (/. 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 3 ) 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 EULECANIUM group), it extends towards the base 
of the haltere where it is slightly expanded to form a small metapleural wing process 
(pwp 3 ). 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 apophysis (pla 3 ). 



OF THE FAMILY COCCIDAE 29 

The epistemum (eps 3 ) 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 3 ) is represented by an irregular, posteriorly produced sclerite. In 
most species a vestigial precoxal ridge (pcr 3 ) extends anteriorly for a short distance 
along the ventral margin of the epistemum, but the absence or presence of this ridge 
varies individually. The metathoracic spiracle (sp 3 ), supported by a peritreme (ptr 3 ), 
is situated in the membrane anterior to the epistemum. 

The metasternum (stn 3 ) 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. prunastri, 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 Eriopeltis 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) U P 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. 

(ii) Dorsospiracidar 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 INGLISIA and COCCUS groups. The number of fleshy setae varies 
from 1-8 (average 3-6) in /. theobromae to 10-23 (average 15) in P. acericola ; hair- 
like setae are rarely present and never total more than 3. 



3 o MORPHOLOGY AND TAXONOMY OF ADULT MALES 

(iii) A ntemetaspiracular 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 INGLISIA and COCCUS groups." 

(iv) Postmetaspiracular setae (eps 3 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. corni), but few in the other groups and absent in 
some species of the EULECANIUM group (e.g. E. tiliae). 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 ERIOPELTIS, 
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. Pfestucae 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 ERIO- 
PELTIS, INGLISIA and COCCUS groups (Text-figs. 24-43) where their number 
varies from 5-16 (average 8) in /. 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. 3-1 1 (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 1-9-2-3 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 radius (rad) and media (med) ; these veins were also called radius and " medius 
+ subcosta " by Durr (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 COCCJ DAE 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 ; ccx). 
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. prunastri, the ERIOPELTIS, 
INGLISIA and COCCUS groups ; Text-figs. 12, 22-43) or reduced to halter es (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. spiraeae, 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 comperi and Microparsus 
variabilis (see Baker, 1920) to a considerable degree, presumably through con- 
vergence. The halteres were called " pseudohaltcres " 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 (is) and hair-like (hs) setae, although the fleshy ones are sometimes absent on 
the tarsus (Genus A and E. Pfestucae). 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 E. 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 f 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 digitides (tdgt), i.e. long, capitate 
setae, is present near the dorsal apex of each tarsus and two smaller ungual digitides 
(udgt) occur on each claw. 



32 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

The coxa (ex) 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. tiliae, F. viburni, Pidvinaria 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. abietis ; Text-figs. 2, 3 ; M ; cb) and all the species of the 
COCCUS group (Text-figs. 31, 33, 35, ^y, 38, 41 ; M ; cb), except C. berliniae. 
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. bitubercidatum to 5-8 (average 
6-2) in E. tiliae. 

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, i.e. less than i\ 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. tiliae, 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 B ; Text-figs. 31, 33) to short and broad, i.e. 3I 
times longer than wide (Genus A, S. pranastri ; 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 tibia (tib) is long and slender. The width/length ratio varies from about 
1 : 11 in S. prunastri and L. luzidae (Text-figs. 22, 27) to about 1 : 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. pda, some of the hair-like setae 
near the apex also have a spur-like appearance. In /. 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 9 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 E. Pfestucae. 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 ungual 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 9th or genital segment ; this was 
recognized by Putnam (1879), Silvestri (1919a, 1919b, 1920), Sulc (1932), Pesson 
(1941), Borchsenius (1957), Theron (1958) and Bustshik & Saakjan-Baranova (1962). 

Pregenital 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. viburni, Ctenochiton sp., Eriopeltis spp., /. theobromae and P. Pbetulae ; 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 /. 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 Sulc (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. prnnastri, 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 Parthenolecaniam 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 all segments. 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- 
pieural 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. htzulae, C. hesperidum, Ceroplastes spp.). In most species 
(S. prunastri, 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 INGLISIA 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 (Eriopeltis, Pidvinaria 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 potich (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 fined 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 Diirr (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. hesperidnm, 
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 Vinsonia stellifera (Newstead, 1903) and Ceroplastes 
japonicus (Borchsenius, 1957). The glandular pouch corresponds to what is called 
the "glandular plate " (Pflugf elder, 1939 ; Giliomee, 1961) in the Pseudoccidae. 

Genital 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 penial 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 
cjaculatorius 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 (1919a, 1919b, 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), Dvirr (1954) and Husseiny & Madsen (1962) called the entire 9th 
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 10th and nth 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) y. ; width at mesothorax 570-660 
(average 601) y. Wing expanse 3950-4550 (average 4296) y. 

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) y, width across genae 296-338 (average 315) y.. 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 0-4 (average 2) longer setae, anterior to the eyes. Midcranial ridge 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) y. in diameter and 2-2-2-9 (average 2-5) times 
as much apart ; those of the ventral eyes 32-42 (average 35) y in diameter and 1 -i-i -6 (average 
1-3) 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. 
Preocular ridge extending only a short distance below articular process. Postocular ridge 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. Interocular ridge 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. Preoral ridge present. Tendon-like apodeme long. Cranial 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 10-segmented ; filiform ; 895-1075 (average 988) y long, i.e. shorter than half 
body length (ratio 1 : 2-46-2-92, average 2-62) ; shorter than posterior leg (ratio 1 : 0-73-0-87, 
average o-8o) and longer than penial sheath (ratio 1 : 1-16-1-50, average 1-31). Scape 
57-68 (average 61) y. long and 57-67 (average 62) y. wide, with 3 h.s. Pedicel with distinct, 
polygonal, dorsal reticulation ; 49-57 (average 54) |j. long and 49-57 (average 53) y wide, with 
0-3 (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 y long and 42-49, 
average 45 y wide) ; with 4-12 (average 8-5) h.s. and 4-12 (average 5-8) f.s., the latter of 
medium length, 0-7-1-0 (average 0-9) times as long as width of segment ; with 4-9 usual 
sensilla basiconica. Segments IV-IX cylindrical ; lengths of these segments (in y) 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 y, 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) y long and 30-38 (average 33) y 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) y. long. 

Prothorax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral 
pronotal sclerites large, 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, interrupted median ridge and a triangular sclerite. Anteprosternal 
setae absent ; prostemal setae 0-3 (average o-8) h.s. 

Mesothorax. Mesoprephragma with shallow emargination. Pre scutum about twice as wide 
as long (average 268 and 138 y 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 



OF THE FAMILY COCCIDAE 



39 




a 

— 

a 



o 

-a 



8 



- 



4 o 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



95) (x long and 2-06-3-00 (average 2-36) times as wide (width 203-251, average 223 y.) ; without 
setae. Scutellum 93-114 (average 107) u. long and 217-266 (average 236) [x wide, the ratio 
being 1 : 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 
postalare well developed, the latter densely reticulated distally. Mesopostphragma 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. Subalare small. Episternum not reticulated; 
subepisternal ridge well developed, becoming appreciably broader ventrally, but below mem- 
branous cleft indistinct and only marked by a band of dark sclerotization. Epimeron small. 
Later opleurite partly bounded anteriorly by an extension from marginal ridge. Basisternum 
large, about 383 u. wide and 287 u. 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 ridges. Furca well developed. Mesothoracic spiracle with well developed peritreme ; 
postmesospiracular 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 sclerite with a pronounced ventral projection at its base. 
Additional sclerite well defined. Antemetaspiracular setae absent. 

Metathorax. 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 ; epimeron produced 
posteriorly. Metathoracic spiracle similar to mesothoracic one. Dorsospiracular setae absent. 
Postmetaspiracular setae : 0-2 (average 09) h.s. Metasternal plate weak and irregular. 
Anterior metasternal setae consisting of 0-2 (average 1-4) medial h.s. and occasionally with one 
or two lateral to the posterior part of the basisternum ; posterior metasternal setae usually 2 
(range 1-3) h.s. medially. 

Wings hyaline ; of medium length (1750-2000, average 1879 y.) but comparatively broad 
(width 780-870, average 840 u.), ratio width to length being 1 : 2-13-2-35 (average 2-24) ; 
alar lobe present ; alar setae : 1-3 (average 1-7) h.s. on each wing. Halteres well developed, 
163-201 (average 178) u. long and 42-61 (average 53) [i wide, each with 2-4 (average 26) 
apically hooked setae which are about 70 [i long. 

Legs short and slender, with fore pair shortest and hind pair longest ; ratio length of hind leg 
to body length is 1 : 2-07-2-15 (average 2-10). Length of segments (in jx) : 



Leg 


Coxa 


Trochanter 


Femur 


Tibia 


Tarsus 


Claw 


Total 


I 


91-103 


118-148 


274-346 


353-437 


93" io 3 


29-34 


973-1094 




(95) 


(128) 


(301) 


(39i) 


(98) 


(30) 


(1043) 


II 


99-118 


120-133 


251-308 


433-479 


H4- I 33 


29-34 


1058-1203 




(no) 


(129) 


(285) 


(45o) 


(124) 


(32) 


(1130) 


III 


106— 141 


125-143 


281-315 


456-532 


125-148 


3o-34 


1132-1303 




(124) 


(134) 


(295) 


(499) 


(134) 


(32) 


(1217) 




Fig. 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 h.s. 

Coxae each with 24-35 setae ; fore coxa with 5-8 (average 62) coxal bristles, of which some 
are capitate ; apical seta about half as long as trochanter. Trochanters 34-44 \i 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 31) times as long as width of trochanter. Femora of medium width (53-67 
[x), ratio width to length of hind femur being 1 : 4-5-5-1 (average 4-9) ; each with 37-65 setae. 
Tibiae 29-38 \i wide, ratio width to length of hind tibia being 1 : 141 163 (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. Tarsi 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) (jl long and 450-600 (average 521) \j. wide. 

Segments I— VI I : tergites and sternites present on all segments ; tergites on segments 11 and 
111 represented by a small sclerite on each side on the anterior margin, and on the IV— VII by 
a transverse plate ; sternites represented by .1 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 11 VII. Pleural setae consisting of h.s. 
only, which include d or sopleural setae : 0-3 (average 1 -j), 2-8 (average 3-8), 1-6 (average 35) 
and 2-6 (average 3-8) on segments III— VI respectively, and ventropleural 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 74) h.s. Ventral setae : h.s. only, usually 2 medially on each of II and 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 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. Ante- 
anal setae : 2 long h.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) u. long and 49-59 (average 55) U. wide ; lateral sclerotizations not 
joined anterior to anus ; length of basal rod fj— j{ that of aedeagus, the rod extending anteriorly 
from base of aedeagus for $ -f 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 \i), penial sheath longer and basisternum shorter, 
the ratios being 1 : 1-9-2-4 (average 22) 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. Rehacek in Bratislava, Czechoslovakia on 
14.iv.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 



42 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 




c 
a 



o 
-a 



o 
l-c 

o 

pq 






OF THE FAMILY COCCIDAE 



43 



(average 2136) y. ; width at mesothorax 415-480 (average 445) jx. Wing expanse 3870-4190 
(average 4072) jx. 

Head subcorneal 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) [x, 
width across genae 251-289 (average 274) p.. 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) jx in diameter and 2-3-3-8 (average 2-9) times as much apart ; those of the ventral eyes 
21-30 (average 26) |x 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. Preocular ridge long, ventrally extending 
half-way or more of the distance between the articular process and the midcranial ridge. 
Postocular ridge very weak dorsally, well developed latero-ventrallv, and tapering but well 
defined posteromedian y ; below ocellus the ridge usually splits up, with the anterior branch 
partly surrounding ocellus. Interocnlar 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. Preoral 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 10-segmented, filiform ; 927-1087 (average 1025) [x long, i.e. shorter than half body 
length (ratio 1 : 2-02-2-13, average 2-09), slightly longer than posterior leg (ratio 1 : 1-04- 
1 14, average 1 07) and longer than penial sheath (ratio 1 : 1-55-1-71, average 1-64). Scape 
53-68 (average 60) jx long and 46-55 (average 51) |x wide, with 3-4 (average 3-2) h.s., area of 
sclerotization reduced ventrally. Pedicel with distinct, polygonal, dorsal reticulation ; 55-68 
(average 59) u. long and 42-51 (average 47) jx wide ; with 0-3 (average 21) f.s., 1-3 (average 
2-5) h.s. and a sensillum placodeum. Segment III somewhat club-shaped, 2-5-30 (average 2-7) 
times longer than wide (80-95, average 90 |x long and 30-36, average 34 \x 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 IV— IX 
cylindrical ; lengths of these segments (in jx) 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 |x ; 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) jx 
long and 23-29 (average 26) jx wide ; carrying 8-13 (average n) f.s., 3 capitate subapical setae 
and 5 antennal bristles of which the 3 long ones are about f 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. 




Fig. 5. Xemolecanium abietis Borchs., lateral view. 



44 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

Thorax : 619-752 (average 699) y, long. 

Prothorax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral 
pronotal sclerites 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 prostemal setae absent. 

Mesothorax. Mesoprephragma with shallow emargination. Prescutum less than twice as 
wide as long (average 197 and 113 y. respectively) ; anterior margin curved ; laterally bounded 
by the prescutal ridges 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) y. long and 1-72-2-27 (average 2) times as 
wide (width 156-190, average 171 y.) ; without setae. Scatellum 72-91 (average 80) y. long 
and 148-190 (average 164) y. wide, the ratio being 1 : 20-23 (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 
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 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 y. wide and 230 y. long, i.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 0-2 (average 1 -4) h.s. on the median 
ridge. Furca well developed. Mesothoracic spiracle with well developed peritreme ; post- 
mesospiracular 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. Antemetaspiracular setae 
absent. 

Metathorax. 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. Metathoracic spiracle similar to mesothoracic one. Dorso- 
spiracular setae absent. Postmetaspiracular setae : 0-2 (average 0-7) h.s. Metasternal plate 
weak and irregular. Anterior and posterior 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 |j.) and of medium width (700-780, average 
753 y.), the ratio width to length being 1 : 2-42-2-51 (average 2-46) ; alar lobe present ; alar 
setae usually one (range 0-2) h.s. on each wing. Halteres well developed, 125-156 (average 
142) y. long and 38-46 (average 43) y. wide, each with 1-3 (average 1 -8) apically hooked setae 
which are about 73 y. 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 y.) : 



Leg 


Coxa 


Trochanter 


Femur 


Tibia 


Tarsus 


Claw 


Total 


I 


76-80 


93-118 


220-266 


344-391 


95-io3 


24-29 


857-984 




(79) 


(107) 


(245) 


(367) 


(98) 


(26) 


(922) 


11 


76-87 


95-110 


i94- 2 5i 


308-365 


101-114 


25-28 


798-955 




(83) 


(103) 


(222) 


(345) 


(108) 


(26) 


(886) 


III 


91-110 


106-125 


210-247 


331-395 


114-125 


26-30 


878-1034 




(100) 


("4) 


(235) 


(37i) 


(118) 


(27) 


(964) 



OF THE FAMILY COCCI DAK 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 1 1 19 h.s. ; fore 1 oxa with 5-6 (average 5-6) coxa! bristles, each with a small apical 
knob ; apical seta about half as long as trochanter. Trochanters 26-38 fi 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 6-1 1 h.s., the latter including 2 minute setae near 
basal ridge, one small seta on the outer m irgin and .' 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 
I ' rmora of medium width (42-57 y.), ratio width to length of hind femur being 1 : 4-5 5-2 
(average 4-8) ; each with 21 35 Is. and 12 [8 b S. Tibiae 24-30 [x wide, ratio width to length 
of hind tibia being 1 : [2-3 14-6 (average 135) ; each with 09 85 setae of which 20-34 are q.S. 
and 44-59 f.s., the latter about 1 1 [J times as long as width of tibia ; apical spur about the 
same size on all tibiae. Tarsi 23-30 y, wide, hind tarsus 3-9 50 (average 4-5) tunes longer than 
wide ; cub with i> [8 f.s. and 11 i<> h.s. ; tarsal digitules subequal, longer than claw. 1 
of medium length, about as long as width of tarsus . shghtlv curved, with small denticle near 
tip ; anterior ungual digitule with larger apical knob than posterior one, digitules about as long 
as c law. 

Ibdomen 490-650 (average 579) u long and 360-430 (average J90) u wide 
Segments 1 VII : tergites and sternites present on all segments ; tergites on segments II, III 
and sometimes IV represented by a small sclerite on each side on .interior margin, and on 
l\' VII by a transverse plate ; stermtes represented by a weak transverse plate on the anterior 

and posterior segments and .1 small sclentc on <a« h side on the intermediate segments Caudal 
extension of segment VII small, rounded, not sclerotized Dorsal setae : lis only, segments 

1 III occasionally with one, and segments IV VII usually with one seta on each side. Pleural 
setae absent on segments I and II and on 111 VI represented by h.s only, which usually include 

2 (range 1-3) dorsopleural setae and 1 ventropleural seta on each segmenl 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 lute-anal setae ; 2 long, and 
• >< 1 asionally one small h.s. Posterior margin with 3 h s on eai h side 

Genital segment. Penial sheath long, about ? tot d body length (ratio 1 : 3-3-3-5, average 34), 
581-695 (average 628) u. long and 40 53 (average 4c)) iz wide ; lateral sclerotizations not joined 
anterior to anus ; length of basal rod about 5 that of aedeagus, the rod extending anteriorly 
from base of aedeagus for about ,", of the distance to apex oJ basil membranous area ; apex 
of sheath without membranous extension. The area from base ol 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 \i), penial sheath longer and basisternum 
shorter, the ratios being 1 : 2-02-3 (average 21) and i : 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) y. ; width at mesothorax 380-500 (average 429) u,. Wing expanse 
2950-3200 (average 3063) u.. 



4 6 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



Head subcorneal 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) 
[i, width across genae 171-224 (average 195) \l. 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. Midcranial 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) \l 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) [i in diameter and 1-0-1-5 (average 1-2) times as much apart. Ocellus 
small. Ocular sclerite well sclerotized and polygonallv reticulated throughout. Preocular ridge 
with ventral part reaching or 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. Interocular 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. Preoral ridge weak, sometimes 
interrupted. Tendon-like apodeme long. Cranial apophysis short ; apex bifurcate, not 
reaching level of posterior margin of ventral eyes. Mouth opening irregular. Anterior tentorial 
pits apparently absent. 





Fig. 6. Physokermes piceae (Schr.), dorsal and ventral view. 



()!•• THK FAMILY COCCIDAE 



47 



Antennae io-segmented, filiform ; 604-101 1 (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 1 : 0-91-1 11, 
average 0-99) and longer than penial sheath (ratio 1 : 1-32-1-69, average 147). Scape 46-72 
(average 59) li long and 42-61 (average 51) Lt wide, with 3 h.s. Pedicel with distinct, polygonal 
dorsal reticulation ; 38-57 (average 46) li long and 42-53 (average 46) li wide ; with 2 h.s. 
and a sensillum placodeum. Segment III bulging in middle, 1-82-6 (average 2-1) times longer 
than wide (61-91, average 73 pi long and 30-42, average 35 li 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 11) times as long as 
width of segment ; with 1-6 usual sensilla basiconica. Segments IV- IX cylindrical ; lengths 
of these segments (in li) 80-160 (average 114), 68-137 (average 98), 76-137 (average 101), 
65-114 (average 86), 57-95 (average 74) and 46-84 (average 61) respectively, widths varying 
from 25 to 34 Li, 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 slightlv 
thicker than f.s. Segment X : terminal part not constricted ; 46-68 (average 58) li long and 
27-31 (average 30) li wide ; carrying 7 1 1 (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 Is. ; with 2 sensilla basiconica ventrallv, one near apex 
and the other more proximal. 

Thorax 494-673 (average 565) \i long. 

Prothorax . Pronolal ridge strong, but medially interrupted by weak sclerotization. Lateral 
pronotal sclerites large, without setae. Medial \ctae absent, Post-tergites medium-sized, without 
striations and without setae I'leiiral structures typical of the family. Sternum with trans- 
verse ridge strong, median ridge reduced to a basal stalk, and a triangular sclerite. Ante- 
prosternal setae absent ; occasionally a hairdike prosternal seta present. 

Mesothorax. Mesoprephragma with no cmargination. Presciitum about twice as wide as long 
(average 163 and 88 li 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) li long and 1-48-2-30 (average 1-79) times as wide, (width 122 170, average 
146 [a) ; without setae. Scutellum 59-103 (average 78) li long and 133 104 (average 151) li 
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 ; postnatal apophysis 
and poslalare well developed, the latter densely reticulated distallv. Mesopostphragma with 
shallow emargination. Mesopleuro>i. 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 not reticulated ; sub- 
episternal ridge well developed, but below the membranous cleft indistinct and only marked by 
a band of dark sclerotization. Epimeron small, l.ateroplcm ite bounded anteriorly by an 
extension from marginal ridge. Basisternum large, about 251 li wide and 168 li 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 i-8) h.s. on or 
near median ridge. Furca well developed. Mesothoracic spiracle with well developed peri- 




FlG. 7. Physokermes piceae (Schr.), lateral view. 



48 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

treme ; postmesospiracular 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. Antemetaspiracular setae 
absent. 

Metathorax. Metanotum with posterior margin usually strong and well developed throughout ; 
siispensorial sclerites small, spot-like. Postnotnm consisting of a transverse sclerite on each side. 
Metatergal setae absent. Pleural ridge well developed, though interrupted near middle ; with 
a small wing process. Episternum with anterior margin not ridge-like ; vestigial precoxal 
ridge present ; epimeron produced posteriorly. Metathoracic spiracle similar to mesothoracic 
one. Dorsospiracular setae absent. Postmetaspiracular setae : occasionally one h.s. present. 
Metasternal plate weak and irregular. Anterior and posterior metasternal setae : 1-5 (average 
2-4) and 0-3 (average 1 -8) h.s. respectively, arranged medially. 

Wings hyaline; medium-sized : 1300- 1450 (average 1367) y. long and 470-580 (averago 542) y 
wide, ratio width to length being 1 : 2-44-2-77 (average 2-53) ; alar lobe present ; alar setae 
absent. Halteres well developed, 103-137 (average 117) y long and 27-43 (average 35) y wide ; 
each usually with 2 (range 1-3) apically hooked setae, which are about 62 y. 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 

I 

II 
III 

F.s. 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 f as long as trochanter. 
Trochanters 27-38 y. 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. Femora of medium width (42-49 y), ratio width to length of hind femur being 

1 : 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. Tibiae 23-30 u. 
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 y wide, hind tarsus 2-9-3-9 (average 3-3) 
times longer than wide ; each with 1-6 f.s. and 6-10 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 ; ungual digitules subequal, about as long as claw. 

Abdomen 380-760 (average 499) y. long and 320-440 (average 376) y. 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, 0-2 (average 1 -i) on III and usually 2 on each 
of segments IV-VI, and ventropleural 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. 



Coxa 


Trochanter 


Femur 


Tibia 


Tarsus 


Claw 


Total 


57-7° 


78-110 


165-217 


209-300 


63-80 


23-27 


600-805 


(66) 


(94) 


(187) 


(248) 


(7i) 


(25) 


(691) 


65-91 


78-110 


154-205 


209-289 


74-95 


26-28 


608-815 


(73) 


(88) 


(171) 


(242) 


(82) 


(27) 


(684) 


7 2 -95 


82-110 


163-220 


220-314 


76-110 


27-30 


667-866 


(80) 


(94) 


(i«7) 


(262) 


(9o) 


(28) 


(74i) 



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 ? total borly length (ratio 1 : 3-2-3-6, average 
34), 456-597 (average 530) [i long and 43-53 (average 48) |j. 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 | - ^ 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 \i), penial sheath longer and basisternum shorter, 
ratios being 1 : 1-49-1-61 (average 1-55) and 1 : 0-43-0-54 (average 0-49) respectively. 

Material examined : 10 specimens, collected by J. Rehacek 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) \i ; width at mesothorax 400-420 (average 411) jz. Wing 
expanse 2860-3000 (average 2940) [i. 

Head subcorneal 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) [x, width across genae 194-232 (average 217) \i. 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) [i 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. Preocular ridge of variable length, ventral part usually extending about 
half-way from articular process to midcranial ridge. Postocular 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. Inter- 
ocular ridge absent. Dorsal ocular setae absent ; ventral head setae : 0-4 (average 2-1) h.s., 
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. 



5° 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



Antennae io-segmented, filiform ; 980-1017 (average 995) y long, i.e. longer than half body 
length (ratio 1 : 1-52— 1-61, average 157), longer than posterior leg (ratio 1 : 1-14-1 21, average 
1-17) and longer than penial sheath (ratio 1 : 2-44-2-63, average 2-53). Scape 38-51 (average 
45) y long and 46-49 (average 48) y wide, with 3 h.s. Pedicel with distinct, polygonal dorsal 
reticulation ; 46-57 (average 54) y long and 42-48 (average 45) u. 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 y long and 23-34, average 29 y. 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 y) 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 y ; 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. ; antenna/ 
bristles on segments VIII-IX not markedly different from f.s. Segment X : terminal part 
not constricted ; 61-72 (average 67) y long and 17-23 (average 21) y wide ; carrying 5-1 1 
(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) y long. 

Prothorax. Pronotal 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. 




Fig. 8. Rhodococcus spiraeae (Borchs.), dorsal and ventral view. 



OF THE FAMILY COCCIDAE 



5' 



Mesothorax. Mesoprephragma with shallow emargination. Presciitum 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 prescittal suture ; sometimes with weak polygonal 
reticulation. Scutum. Median membranous area subrectangular ; 49-65 (average 59) [x long 
and 1-81-2-36 (average 2-01) times as wide (width 106-125, average 117 u.) ; without setae. 
Stutellum large, 76-95 (average 84) u. long and 125-144 (average 135) u, wide, the ratio being 
1 : 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. Mesopostphragnia 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. Subalare small. Episternum not reticulated ; sub- 
episternal ridge well developed, but below membranous cleft indistinct and only marked by a 
band of dark sclerotization. Epimeron small. Lateropleurite partly bounded anteriorlv by 
an extension from marginal ridge, liasisternum large, about 237 fx wide and 160 u. 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. Eurca well developed. Meso- 
thoracic spiracle with well developed peritreme ; postmesospiracular setae absent. Tegula 
small, membranous bulge with a small weak sclerite posteriorly and 2-5 (average 3-2) h.s. 
Third axillary wing sclerite with a pronounced ventral projection at its base. Additional 
sclerite well defined. Antemetaspiracular setae absent. 

Metatkorax. Metanotum with thickening of posterior margin desclerotized mediallv ; 
suspensorial scleriles 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 ridge present ; epimeron produced posteriorly. Metathoracic spiracle similar to meso- 
thoracic one. Dorsospiracular setae absent. Postmetaspiracular setae : occasionally one h.s. 
Metasternal plate weak and irregular. Anterior and posterior metasternal setae absent. 

Wings hyaline ; long (1270-1350, average 1310 jx) and of medium width (480-540, average 
512 (x), 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) jx 
long and 27-42 (average 37) |x wide, each with 2 (rarely 3) apically hooked setae, which are 
about 69 (x long. 

Eegs 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 |x) : 



Leg 


Coxa 


Trochanter 


Femur 


Tibia 


Tarsus 


Claw 


rotal 


I 


65-76 


99-110 


205-228 


308-315 


76-95 


27-30 


798-834 




(68) 


(103) 


(218) 


(312) 


(89) 


(29) 


(819) 


n 


72-84 


95-105 


182-200 


308-317 


91-99 


30-34 


798-820 




(81) 


(IOI) 


(192) 


(3") 


(97) 


(3i) 


(813) 


in 


76-87 


99-1 10 


186-209 


312-352 


103-106 


30-34 


834-882 




(83) 


(104) 


(200) 


(33«) 


(105) 


(32) 


(862) 




Fig. 9. Rhodococcus spiraeae (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 
f as long as trochanter. Trochanters 29-38 y. 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. Femora of medium width (42-51) y, 
ratio width to length of hind femur being 1 : 3-8-4-4 (average 4-1) ; each with 6-14 f.s. and 
1 1-25 h.s. Tibiae 23-40 [i wide, ratio width to length of hind tibia being 1 : n -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. Tarsi 21-27 H wide, hind 
tarsus 4-3-4-7 (average 4-6) times longer than wide ; each with 4-8 f.s. and 15-26 h.s. ; tarsal 
digitales subequal, long, about i| times as long as claw. Claws long, hind claw about i£ 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) y long and 300-350 (average 331) (x wide. 

Segments I-VII : tergites on segments II— III represented by a small sclerite on each side on the 
anterior margin, and on VI and VII by a weak plate ; sternites 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 : 0-2 (average i-o), 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) h.s. Ventral setae : h.s. 
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 i\-2 times as long 
as section within pouch. A small IXth tergite sometimes perceptible. Ante-anal setae : 1-2 
(average i-8) long h.s. Posterior margin with 2-3 (average 2-8) h.s. on each side. 

Genital segment. Penial sheath long, about \ total body length (ratio 1 : 3-8-4-1, average 
3-9), 376-414 (average 399) y long and 36-40 (average 38) y 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 f 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 u.), 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 : 9 specimens, collected by G. Matesova on Spiraea hypersifolia 
L. in the ravine Talgar, Zailiiski Alatau, Kazakhstan, USSR, on 14.vi.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) |jl ; width at mesothorax 290-320 
(average 310) y. Wing expanse 2470-2660 (average 2566) y. 

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 



OF THE FAMILY COCCIDAK 



53 



220) (x, width across genae 217-241 (average 233) (x. 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 39) times as much apart ; those of the ventral eyes 22-27 
(average 24) \x in diameter and 1-2-1-8 (average 1 -6) times as much apart. Ocellus small. 
Ocular sclerite well sclerotized and polygonally reticulated throughout. Preocular 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. Interocular ridge 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 10-segmented, filiform ; 705-883 (average 819) jx long, i.e. longer than half body 
length (ratio 1 : 1-63-1-96, average 176), 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) u. long and 40-46 (average 42) u. wide, with 3 h.s. Pedicel with distini t, polygonal, dorsal 
reticulation ; 42-51 (average 47) ;x long and 38-42 (average 40) ;x wide ; with 1-7 (average 
J-]) f.s., 2-5 (average 35) f.s. and a scnsillum placodeum. Segment III club-shaped, 2-7-3-6 
(average 3-2) times longer than wide (76-87, average 81 jx long and 21-30, average 26 (x wide), 




Fig. 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 jx) 87-122 (average 109), 195-125 (average 118), 91-114 (average 101), 80-110 
(average 100), 65-91 (average 76) and 57-76 (average 68) respectively, width varying form 17 to 
23 [i., 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) [i. long and 19-23 (average 21) pi wide ; carrying 4-1 1 (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) \x long. 

Prothorax. Pronotal 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-tergites very small and weakly sclerotized, without 
striations and without setae. Pleural structures typical of family. Sternum with strong 
transverse ridge, median ridge reduced to a basal stalk or sometimes absent, and a triangular 
sclerite. Anteprosternal setae absent ; prosternal setae : occasionally one h.s. present. 

Mesothorax. Mesoprephragma with shallow emargination. Prescutum about twice as wide 
as long (average 145 and 74 \l respectively) ; anterior margin curved ; laterally bounded by 
the prescutal ridges 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 area subrectangular ; 76-91 (average 82) y. long and 1-47-1-75 (average 
1-57) times as wide (width 1 18-137, average 128) ; with 1-4 (average 2-7) h.s. Scutellum 
57-68 (average 63) [j. long and 118-133 (average 127) [i wide, ratio being 1 : 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 postalare well 
developed, the latter densely reticulated distally. Mesopostphragma with 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 
narrow basalare. Subalare small. Episternum not reticulated ; subepisternal ridge well 
developed, but below membranous cleft indistinct and only marked by a band of dark sclero- 
tization. Epimeron small. Later opleurite anteriorly bounded by an extension from marginal 
ridge. Basisternum large, about 199 \i wide and 148 [i long, i.e. 1-7-2-1 (average i-8) 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 4-7 (average 5) h.s. Third axillary wing sclerite 
with a pronounced ventral projection at its base. Additional sclerite well defined. Antemeta- 
spiracular setae absent. 




Fig. 11. Palaeolecanium bituberculatum (Targ.), lateral view. 



Coxa 


Trochanter 


l emur 


Tibia 


Tarsus 


Claw 


Total 


57-6i 


76-84 


171-186 


•255-285 


89-99 


25-27 


695-735 


(58) 


(80) 


(183) 


(274) 


(95) 


(26) 


(715) 


61-65 


68-72 


163-175 


247-277 


87-95 


23-25 


654-701 


(63) 


(70) 


(169) 


(261) 


(9o) 


(24) 


(678) 


57-68 


67-72 


r.63- 173 


257 - s '' 


87-99 


24-27 


669-722 


(65) 


(69) 


('70) 


(273) 


(94) 


(26) 


(697) 



OP 1 HE FAMILY COCCI DA 11 55 

Metathorax. Metanotum with anterior margin desclerotized medially ; suspensorial scleriles 
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 precoxal 
ridge sometimes present ; epimeron produced posteriorlv. Metathoracic spiracle similar to 
mesothoracic one. Dorsospiracular setae absent. Postmetaspiracular setae : occasionally one 
h.s. Metasternal plate weak and irregular. Anterior and posterior metasternal setae absent. 

Wings hyaline ; long (1110-1190, average 1155 u.) and of medium width (450-510, average 
474 |i), 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) jz long and 20-30 (average 25) u. wide, 
each with one apically hooked seta, which is about 50 [■. 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 u.) : 

Leg 

I 

II 
III 

Coxae with 7-14 (average 10) f.s. on the fore and 14-19 on the middle and hind coxa, and eai li 
with 7-15 h.s. ; fore coxa with 1-2 (average 1 4) pointed coxa! bristles, about as long as segment ; 
api( il seta about half as long as trochanter. Trochanters 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 [ore tnx banter, is 1 s 2*2 (average 
2) times as long as width of trochanter. Femora of medium width (34-40 (j), ratio width to 
length of hind femur being 1 : 4-4-5-0 (4-7) ; each with 8-14 f.s. and 12-19 h.s. Tibiae 
19-23 [i wide, ratio width to length of hind tibia being 1 : 12-3-14-6 (average 131) ; each with 
35-51 setae of which ig-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 (■. wide, hind tarsus 4-2— 4-9 (average 
4-4) times longer than wide ; each with 4-8 f.s. and 11-17 h.s. ; 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) u, long and 240-300 (average 263) \jl wide. 

Segments I-VII : tergites on segments 1 1 — 1 1 1 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 ventropleural 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, 1 
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 2-2^ times as long 
as section within pouch. No IX tergite observed. Ante-anal setae : 2 long h.s. Posterior 
margin with 2-4 (average 26) 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) u. long and 46-61 (average 52) u. wide ; lateral 
sclerotizations not joined anterior to anus; length of basal rod %-% 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 \i), 
penial sheath longer and basisternum as long or longer, the ratios being 1 : 1-8-2-0 (average 
1-9) and 1 : 0-99-1-14 (average 1-04) respectively. 

Material examined : 10 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. Rehacek on Crataegus oxyacantha L. in Novy Bydzov, 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 \i 
each, (ii) the darker median sclerotization on the prescutum was less well developed 
and not ridge-like in appearance and (iii) 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 1 460-1 620 
(average 1553) jx ; width at mesothorax 370-410 (average 387) [x. Wing expanse 2870-3010 
(average 2940) \l. 

Head subconical in dorsal view ; in lateral view obliquely dorsoventrally elongated, with 
anterodorsal bulge not pronounced ; length to pronotal ridge 209-247 (average 226) ;x, width 
across genae 255-270 (average 262) [x. 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. Midcranial 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) u. in diameter and 3-4-4-5 (average 4-1) times as much apart ; 
those of the ventral eyes 25-30 (average 28) jx in diameter and 1 -3-1 -7 (average 1 -5) times as 
much apart. Ocellus small. Ocular sclerite well sclerotized and polygonally reticulated 
throughout. Preocular 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. Interocular ridge 
absent. Dorsal ocular setae absent ; ventral head setae : 8-14 (average 10) h.s., situated around 
posterior part of the midcranial ridge. Preoral ridge 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 10-segmented, filiform ; 910-1053 (average 986) ;x long, i.e. longer than half bod}' 
length (ratio 1 : 1-43-1-65, average 1-57), longer than posterior leg (ratio 1 : 1-23-1-33, average 
1-27) and longer than penial sheath (ratio 1 : 3 15-3 -47, average 3-36). Scape long (length 
65-68, average 66 jx) and 42-48 (average 45) (x wide, with 1-3 (average 2) h.s. Pedicel not 
reticulated, but with one or two wavy lines ; 61-68 (average 65) fx long and 42-46 (average 44) 
(x 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 [x long 



OF THE FAMILY COCCIDAK 



57 



and 42-46, average 44 u. wide), with 4-8 (average 6) f.s. of medium width, 1-2-1-5 (average 13) 
times as long as width of segment ; with 1-3 usual sensilla basiconica. Segments IV-IX 
cylindrical ; lengths of these segments (in (i) 1 18-152 (average 138), 122-148 (average 137), 
1 1 8-1 4 1 (average 131), 99-133 (average m), 91-106 (average 97) and 80-99 (average 86) 
respectively, widths varying from 21-27 V- 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) 
\x 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 ernes rather 
similar to the f.s. ; with 2 sensilla basiconica ventrally, one near apex and the other more 
proximal. 

Thorax 460-532 (average 503) ;x long. 

Prothorax. Pronotal ridge strong, but medially interrupted by weak sclerotizatioii. Lateral 
pronolal 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 rathei 
distinct sternal apophyses. Anteprosternal setae absent ; prosternal setae : sometimes one h.s. 
on each side. 

Mrsothorax. Mesoprephragma with shallow emergination. Prescutum about twice as 
wide as long (average 160 and 83 \i respectively) ; anterior margin strongly curved ; laterally 
bounded by the prescutal ridges and posteriorly by the presented suture ; slightly more heavily 
sclerotized medially but not reticulated. Scutum. Median membranous area subrectangular ; 




pr. 




Q Eb~ 





Fig. 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 u.) ; with 1-2 (average 1-7) h.s. Scutellum 57-68 (average 64) \i long and 160-171 (average 
166) [x wide, the ratio being 1 : 1-8-2-2 (average 19) ; not tubular ; without setae. Post- 
notum with anterior margin irregular, not overlapped by the metathoracic fold ; postnotal 
apophysis and postalare well developed, the latter densely reticulated distally. Mesopost- 
phragma small, with moderately deep emargination. Mesoplenron. Mesoplenral ridge strong, 
not interrupted above coxal articulation ; pleural apophysis and pleural wing process well 
developed, latter connected with episternum by a narrow basalare. Subalare small. Epi- 
sternum not reticulated ; subepistemal ridge well developed, but below membranous cleft in- 
distinct and only marked by a band of dark sclerotization. Epimeron small. Later opleurite partlv 
bounded anteriorly by an extension from marginal ridge. Basisternum large, about 210 [z wide 
and 151 [i. 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 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 0-4 
(average 1-9) h.s. Third axillary wing sclerite with a small ventral projection at its base. 
Additional sclerite well defined. Antemetaspiracular setae absent. 

Metathorax. Metanotum with anterior margin desclerotized medially ; sitspensorial 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 spiracle similar to mesothoracic one. Dorsospiracular 
setae absent. Postme/aspiracular setae : 2-6 (average 4) f.s. and 0-1 (average 0-7) h.s. Mela- 
sternum represented by a weak, transverse plate. Anterior and posterior metasternal setae 
absent. 

Wings hyaline ; long (length 1260-1330, average 1295 y.) and broad (width 570-600, average 
585 y.), ratio width to length being 1 : 2 19-2-24 (average 2-22) ; alar 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 : 
segments (in \l) : 

Leg Coxa Trochanter Femur Tibia 

I 65-80 91-99 198-213 285-304 

(74) (95) (207) (295) 
II 68-80 87-93 182-194 281-296 

(75) (9i) (189) (289) 
III 76-91 91-99 179—190 289-300 

(83) (95) (184) (296) 



1-89-2-05 (average 1-99). Length of 



Tarsus 
99-122 

(107) 
99-106 

(102) 
99-110 

(106) 



Claw 
23-27 

(26) 
23-29 

(26) 
21-29 

(26) 



Total 
785-828 

(804) 
762-785 

(77i) 
771-800 

(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 A 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 




Fig. 13. Phyllostroma myrtilli (Kalt.), lateral view. 



OF 111 I. 1 AM II.Y COCCI DAE 59 

minute setae near basal ridge, one small seta on outer margin and a long apical seta which, on 
the fore trochanter, is i -3-21 (average 1 -8) times as long as width of trochanter. Femora of 
medium width (38-46 \jl), ratio width to length of hind femur being 1 : 3-9-4-6 (average 4-2) ; 
each with 10-19 f.s., and with 15-24 (average 19) h.s. on tin- lore and 1 1 16 on the middle and 
hind femur. Tibiae 23-27 ji. 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 Is, the latter somewhat 
longer than width of tibia ; apical spur about the same size on all tibiae. Tarsi 22-27 V- wide, 
hind tarsus 4-0-4-8 (average 4-5) times longer than wide ; eai h with j j K and IO-ig h S 
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) ^ long and 320 380 (average 346) u wide 

Segments I— VI I : termite* and siemites present on .ill segments ; tergites represented bj 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 
I and V -VII, Pleural setae 1 onsisting 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 : lis only, 
usually none on II, one on ea< h side on III and IV, and | on ea< h of segments V VI I 

Segment VIII with transverse tergite and sternite ; caudal extension forming a small, simple 
lobe ; glandulai pouch with .' long, pointed setae, whose protruding part is aboul 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 \ (average 3-5) h.s. on each side 

Genital segment. Penial sheath of medium length, about { total body length (ratio 1 497 
5-57, average 5*27), 285 304 (average 294) u. long and 46-49 (averagt p u wide ; lateral 
Si lerotizations narrowly joined .interior to .inns ; length of basal rod about : ,' th.it of aedeagus, 
the rod extending anteriorly from base of aedeagus to the apex oi the basal membranous area . 
apex of shea tli without membranous extension The area from base of sheath to tip of aed< 
with 11-19 (average 15) small setae ; .1 cluster ol small sensilla oa Hiring ventrally near apex 
of sheath. Aedeagus of medium length (118—137, average 127 |i), penial sheath and basisterimni 
longer, the ratios being 1 : 2-2-2-6 (average 23) and 1 : i-ii 1 20 (average 119) respectiveU 

Material examined : 8 specimens, collected by J. Rehacek on Vaccimttm myrtillus 
L. in Czechoslovakia on 6.vii.io,53 (remounted from Swarm's mountant). 



FILIPPIA 
Filippia viburni (Signorel 

(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) u. ; width at mesothorax 440-490 (average 469) pt. 
Wing expanse 3630-3940 (average 3798) u.. 

Head subcorneal 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) jx. Median crest sclerotized and striated posteriorly, 
but usually without polygonal reticulation ; with 7-12 (average 9-3) hairdike 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 



6o 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 




C 

> 

c 



o 



a 

be 

c/5 



a 



o 



OF THE FAMILY COCCIDAK 



61 



which is about half as big as the other two ; corneae of dorsal eyes 27-30 (average 29) [j. in 
diameter and 3-0-4-3 (average 3-6) times as much apart ; those of the ventral eyes 29-33 
(average 30) u. in diameter and 1-2-1-7 (average 14) times as much apart. Ocellus small. 
Ocular sclerite well sclerotized and polygonally reticulated throughout. Preocular ridge with 
ventral part almost reaching midcranial ridge. Postocular ridge tapering dorsally, well developed 
lateroventrally, and narrow but well defined posteromedially ; below ocellus the ridge splits up, 
with anterior branch partly surrounding ocellus. I nterocular ridge absent. Dorsal ocular 
setae absent ; ventral head setae : 2-10 (average 6-3) h.s., situated medially just behind the 
preocular ridges. Preoral 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 10-segmented, filiform ; 1 120 1370 (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 : 106-118, average 
1 13) and longer than penial sheath (ratio 1 : 3-38-3-94, average 3-74). Scape 59-65 (average 
62) u long and 46-51 (average 48) u. wide, with 34 (average 33) h.s. Pedicel with distinct. 
polygonal, dorsal reticulation ; 57-65 (average 62) u. long and 49-59 (average 54) u. wide ; 
with 2-6 (average 3-6) f.s., 7-1 1 (average 8-4) h.s. and a sensillum placodeum. Segment 111 
bulging in the middle, 1-9-2-6 (average 21) times longer than wide (76-87, average 80 u. long 
and 34 30, average 35 (jl wide) ; with 4-0 (average 7-1) h.s and 4 12 (average 7-9) f. s., the latter 
of medium length, 0-9-1-2 (average 11) times as long .is width of segment ; with 1-3 usual 
sensilla basiconica. Segments IV IX cylindrical ; lengths of these segments (in \L) 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 2<> \i; with 33-45 (average 38), 37-44 (average 41), 30 49 (average 41 1, 28 39 (average 33), 
7-22 (average 16) and 10-19 (average 14) f.s, and 24 (average 3), 3 9 (average 51), 2-7 
(average 4), 3-6 (average 4-2), 1-4 (average 2-9) and 0-5 (average 23) h.s. respectively . 
antennal bristles on segments YIII-IX distini tly larger than f.s. Segment X : terminal part 
not constricted ; 86-90 (average 02) (x long and 23-26 (average 24) \i wide ; carrying 6-12 
(average 9) f.s., 0-3 (average i-i) h.s., 3 capitate sitbapical setae and 5 antenna! bristles of whi< h 
the 3 long ones are up to half as long as the segment an 1 the 2 shorter ones somewh it 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) [i. 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-tergiles narrow, elongated, without stn.itions and without setae. 
Pleural structures typical of the family. Sternum 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. 




Fig. 15. Filippia viburni (Sign.), lateral view, 



62 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

Mesothorax. Mesoprephragma with shallow emargination. Prescutum more than twice as 
wide as long (average 237 and 106 y 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. Scutum. Median membranous area sub- 
rectangular ; 120-133 (average 128) y long and 1-54-1-94 (average i-68) times as wide (width 
201-236, average 213 y) ; with n-19 (average 15) h.s. Scutellum 76-86 (average 82) y long 
and 201-228 (210) y 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 postalare well developed, the latter densely reticulated distally. Meso- 
postphragma with 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. 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 anteriorly 
bounded by an extension from marginal ridge. Basisternum large, about 293 y wide and 231 u. 
long, i.e. 1-66-1-95 (average i-8i) 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 5-9 
(average 7) h.s. Third axillary wing sclerite with a pronounced ventral projection at its base. 
Additional sclerite small, but well defined. Antemetaspiracular setae absent. 

Metathorax. 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 
ridge well developed, though interrupted in middle ; with small wing process. Episternum 
with anterior margin not ridge-like ; vestigial precoxal ridge present ; epimeron produced 
posteriorly. Metathoracic spiracle similar to mesothoracic one. Dorsospiracular setae absent. 
Postmetaspiracular setae : 4-9 (average 5-5) f.s. and 0-5 (average 2-5) h.s. Metasternum 
represented by a weak, transverse plate. Anterior and posterior metasternal setae : 0-3 (average 
o-8) and 0-1 (average o-6) h.s. respectively. 

Wings hyaline ; long (length 1650-1770, average 1713 y) and broad (width 750-860, average 
818 pi), ratio width to length being 1 : 2-04-2-25 (average 2-09) ; alar lobe present ; alar setae 
absent. Halteres well developed, 106-133 (average 123) y long and 23-33 (average 26) y 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 i-8o). Length of segments (in y) : 

Leg 
I 

II 

III 

Coxae each with 21-32 f.s. and 12-23 ns - '• f° re coxa with 1-3 (average 2) pointed coxal 
bristles which are about \ as long as the segment ; apical seta about \ as long as the 
trochanter. Trochanters 34-38 y 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. Femora of medium width (46-49 y), ratio width to length of hind 
femur being 1 : 5-5-6-3 (average 5-8) ; each with 34-45 f.s. and 18-34 ns - Tibiae 
27-30 y 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 



Coxa 


Trochanter 


Femur 


Tibia 


Tarsus 


Claw 


Total 


74-84 


105-118 


258-308 


456-4 8 3 


129-139 


23-27 


1062-1144 


(78) 


("3) 


(284) 


(467) 


(133) 


(25) 


(1101) 


91-103 


105-118 


250-277 


418-448 


129-137 


23-26 


1018-1100 


(96) 


(113) 


(266) 


(439) 


(134) 


(25) 


(1072) 


106-118 


110-118 


266-289 


429-479 


127-141 


25-29 


1068-1163 


(113) 


(116) 


(276) 


(45i) 


(136) 


(26) 


(1119) 



OF THE FAMILY COCCI DAE 63 

tibia ; apical spur about the same size on all tibiae. Tarsi z^-zj \j. wide, bind tarsus 51 5-3 
(average 5-2) times longer than wide ; each with 13— ig 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 cursed, with small denticle near tip ; ungual digitules subequal, about as long as c law 

Abdomen 620-710 (average 669) [i. long and 400-500 (average 446) ;z wide. 

Segments [-VII : tergites on segments II-1II represented by 3 small sclerites on anterior 
margin, one medially and one on each side, and on A 1 1 bv a weak transverse plate in middle of 
segment ; sternites present on all segments, represented bv 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 1 and V VI I and sometimes one or two on IV Pleural setae 
consisting of h.s. only which include dorsopleural setae : up to 3 on 11,2 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 : lis 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 twi< e 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) u. long and 56-63 (average 60) u, wi le . lateral sclerotizations ool 
joined anterior to anus ; length of basal rod 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 oi aedeagus with 28 \2 
(average 33) small setae ; a cluster of small sensilla occurring ventrally near apex of sheath 
Aedeagus long (160-186, average 174 \x), penial sheath and basisternum longer, the ratios being 
1 : 1 -8-2-1 (average 2) and 1 : 123 130 (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 (x. The same is true for 4 
specimens collected by J. Rehacek on Hedera helix L. during May, 1953 in Czecho- 
slovakia ; they measured 1660, 1680, 1760 and 1830 0.. 



CTENOCHITON 

Ctenochiton sp. 

(Text-hgs. 16 and 17) 

A moderately small and slender species, with comparatively long antennae and moderatelv 
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) [i. 
Wing expanse 3052-3265 (average 3178) jz. 

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) 
ji,, width across genae 240-280 (average 256) ;x. Median crest sclerotized and striated posteriorly, 
but not polygonally reticulated ; with 9-15 (average 12) hair-like dorsal head setae. Mid- 
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 



6 4 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



(average 23) y in diameter and 31-5-8 (average 4-6) times as much apart ; those of the ventral 
eyes 20-27 (average 23) y. in diameter and 1 -2-1 -6 (average 1 -5) 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. Postocular 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. Interocular 
ridge absent. Dorsal ocular 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. Preoral 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 10-segmented, filiform ; 1024-1144 (average 1070) y, long, i.e. longer than half body 
length (ratio 1 : 1-51-1 69, average 1-63), longer than posterior leg (ratio 1 : 1-01-1-05, average 
1-04) and longer than penial sheath (ratio 1 : 3-30-3-69, average 3-48). Scape 52-60 (average 
58) y. long and 40-48 (average 46) y wide, with 3 h.s. Pedicel with weak polygonal, dorsal 
reticulation ; 44-52 (average 46) y long and 40-48 (average 45) y 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 y long and 26-32, average 29 
y. wide) ; with 2-4 (average 3-1) h.s. and 6-10 (average 8-3) f.s., the latter of medium length, 
1-1-1-4 (average 1-3) times as long as width of segment ; with 2-4 usual sensilla basiconica. 
Segments IV-IX cylindrical ; lengths of these segments (in y) 152-180 (average 166), 148-164 
(average 155), 148-180 (average 166), 124-164 (average 147), 100-124 (average 111) and 72-94 




Fig. 16. Ctenochiton sp., dorsal and ventral view. 



I )l 111 E I- AMI1.Y COCCIDAE 



65 



(average 80) respectively, all of about the same width, varying from 19 to 25 (x ; 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), 1-4 (average 2-2), 1-3 (average 2-3), 1-3 
(average 2-1), 0-2 (average 06) and 0-2 (average 1) h.s. respectively ; antenna! bristles on 
segments VIII-IX distinctly larger than fleshy setae. Segment X : terminal part not con- 
stricted ; 76-84 (average 80) jx long and 22-26 (average 24) (x wide ; carrying 6-12 (average 10) 
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 1 scnsilla basiconica ventrally, one near apex and the other more 
proximal. 

Thorax 584-664 (average 619) ;x long. 

Prothorax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral 
pronotal sclerites small, without seate. Medial pronotal setae absent ; 3-7 (average 61) circular 
pores present on each side posterior to pronotal sclerite. Post-tergites small, without striations, 
and without setae. Pleural structures typical of family. Sternum with a strong transverse 
ridge, a narrow but uninterrupted median ridge and a triangular sclerite. Anteprosternal setae 
absent ; prosternal setae : occasionally one h s 

Mesolhorax. Mesoprephragma with shallow emargination. Prescutiini about twice as wide as 
long (average 201 and 98 u. respectively) ; anterior margin strongly curved ; laterally bounded 
by the prescntal ridges and posteriorly by the prescutal suture ; weakly, irregularly reticidated ; 
slightly more heavily sclerotized medially. Scutum. Median membranous area subrectangular; 
108-128 (average 116) u. long and 1-28-1-78 (average 1-49) time as wide (width 152-192 average 
I 73 V-) '• with 7-13 (average 10) h.s. Scutellum 61-87 (average 72) |x long and 160 192 (average 
175) u. wide, the ratio being 1 : 2-2-2-8 (average 24) ; 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. Basisternum large, about 253 [x wide and 
207 jx 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 pcritreme ; postmesospiracular 
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 sclerite small, but well defined. Antemetaspiracular setae absent. 

Metathorax. Metanotum with anterior margin usually strong medially ; suspensorial 
sclerites small, spotdike ; 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, 3-1 1 (average 6-3) circular pores occurring near each seta. Pleural ridge well developed, 




Fig. 17. Ctenochiton sp., lateral view. 



Leg 


Coxa 


Trochanter 


Femur 


I 


68-76 


92-106 


198-222 




(7i) 


(100) 


(211) 


II 


72-84 


92-106 


178-207 




(81) 


(99) 


(i93) 


III 


80-91 


95-116 


190-218 




(85) 


(104) 


(204) 



66 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

though interrupted in middle ; with small wing process. Episternum with anterior margin net 
ridge-like ; vestigial precoxal ridge present ; epimeron posteriorly directed. Metathoracic 
spiracle similar to mesothoracic one. Dorsospiracular setae absent. Postmetaspiracnlar setae ? 
2-6 (average 3-5) f.s. and 1-7 (average 4) h.s. Metasternum represented by a weak, transverse 
plate. Anterior metasternal setae : 1-3 (average 1-9) medially situated h.s. ; posterior meta- 
sternal setae : occasionally one h.s. 

Wings hyaline ; long (length 1357-1463, average 1427 ja) and of medium width (530-615, 
average 578 \i), ratio width to length being 1 : 2-36-2-56 (average 2-46) ; alar lobe present ; 
alar setae : one or occasionally two h.s. on each wing. Halteres well developed, 104-120 (average 
111) [i long and 20-28 (average 24) [i. wide, each with one apically hooked seta which is about 
64 n 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 |j.) : 

Tibia Tarsus Claw Total 

359-422 1 10-128 23-26 903-1013 

(39i) (119) (24) (964) 

331-414 II5-I33 24-27 859-1010 

(377) (124) (25) (947) 

361-426 1 12-135 24-27 906-1053 

(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 \ as long as the segment ; apical seta about \ as long as trochanter. 
Trochanters 26-30 [j. 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 1 -4-1 -7 (average 1 5) times as long as width of trochanter. Femora of 
medium width (38-42 \l), the ratio width to length of hind femur being 1 : 4-7-5-4 (average 
5-1) ; each with 22-34 Is - an d 13-26 h.s. Tibiae 19-25 \x 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) \i 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 ; ungual digitules subequal, about as long as claw. 

Abdomen 551-721 (average 613) \l long and 371-477 (average 411) jjl 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 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 ; ventroplenral 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) \l long and 42-49 (average 46) \x 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 (x), penial 
sheath and basisternum longer, the ratios being 1 : 26-30 (average 2-7) and 1 : 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. 



ERICER US 

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) y. ; width at mesothorax 570-740 (average 680) jjl. 
Wing expanse 5330-5700 (average 5563) u. 




FlG. 18. Ericerus 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) u. and width across genae 285-342 
(average 315) u.. Median crest sclerotized and densely polygonally reticulated ; with 5-7 
(average 5 -7) hair-like dorsal head setae anteriorly. Midcranial 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) \x in diameter and 
1-4-2-1 (average 1-7) times as much apart, those of the ventral eyes 42-49 (average 46) \i in 
diameter and 0-9-1 -i (average 1) times as much apart. Ocellus small. Ocular sclerite well 
sclerotized and densely polygonally reticulated throughout. Preocular ridge with ventral part 
reaching or almost reaching midcranial ridge. Postocular ridge weak dorsally, well developed 
lateroventrally, and narrow but well defined posteromedially ; below ocellus the ridge splits up, 
with anterior branch partly surrounding ocellus. Interocular ridge absent. Dorsal ocular 
setae absent ; ventral head setae : 3-5 (average 4-4) h.s., situated anterior to ocular sclerite on or 
near midcranial ridge. Preoral 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) \i long, i.e. longer than half 
body length (ratio 1 : 1-46-1-55, average 1-50), longer than posterior leg (ratio 1 : 1-10-1-17, 
average 1-13) and longer than penial sheath (ratio 1 : 3-50-4-04, average 3-89). Scape 80-99 
(average 93) ^i long and 76-95 (average 86) [x wide, with 3 h.s. and 2-4 (average 2-6) f.s. Pedicel 
with distinct, polygonal, dorsal reticulation ; small, 61-84 (average 73) u. long and 57-68 
(average 64) |j. 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 \x long and 38-48, average 43 u. wide) ; with 6-1 1 (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 IV— IX 
cylindrical ; lengths of these segments (in y.) 213-274 (average 248), 235-281 (average 264), 
232-312 (average 277), 194-274 (average 235), 156-205 (average 186) and 1 18-156 (average 
145) respectively, all of about the same width, varying from 27 to 38 \i ; 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) \i long, and 
29-34 (average 31) u. wide ; carrying 6-1 1 (average 8-2) f.s., 3 capitate subapical setae and 5 
antennal bristles of which the 3 long ones are about f 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. 

Thorax 832-1 113 (average 1015) u. long. 




Fig. 19. Ericerus pela (Chav.), lateral view. 



OF THE FAMILY COGCIDAE 69 

Prothurax . 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. Sternum with trans- 
verse ridge strong, median ridge reduced to a short basal stalk and triangular sclerite well 
sclerotized. Anteprosternal setae absent ; prosternal setae : 0-4 (average 1 -8) h.s. 

Mesothorax. Mesoprephragma with shallow emargination. Prescutum less than twice as 
wide as long (average 287 and 185 u. respectively) ; anterior margin curved ; laterally bounded 
l>v the prescntal ridges and posteriorly by the presculal suture ; with weak, irregular reticula- 
tion ; slightly more heavily sclerotized medially. Scutum. Median membranous area 
trapezoidal ; 108-186 (average 156) \i long and 1 -57- 2 40 (average 1 95) times as wide (width 
247—334, average 298 y.) ; without setae. Scutellum large, 76—114 (average 96) \l long and 
209-293 (average 250) \l 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 ; 
postnatal apophysis and postalare well developed, the latter densely reticulated distally. Meso 
postphragma with deep emargination. Mesopleuron. Mesopleural ridge strong, but interrupted 
above coxal articulation ; pleural apophysis and pleural wing process well developed, the 
I. iti> 1 connected with episternum by a narrow basalare. Subalare small, lipisternum not 
reticulated ; subepisternal ridge well developed, but below membranous cleft indistinct and 
only marked by a band of dark sclerotization. Epimeron small. Later opleurite partly 
bounded anteriorly by an extension from marginal ridge. Basisternum large, about 430 u. 
wide and 356 pi long, i.e. 2-08-2-81 (average 2-31) times Longer than membranous area of scutum ; 
with strong median ridge and bounded by strong marginal and precoxal ridges ; without setae. 
/•urea well developed. Mcsotlioracic spiracle with well developed peritrcme; postmesospinu ultii 
setae absent. Tegula small, membranous bulge with a weak sclerite posteriorly and 2-7 
(average 4-7) h.s. Third axillary wing sclerite with ,1 pronounced ventral projo tion at its I 
Idditional sclerite elongated, well defined. Antemetaspiracular setae absent. 

Metathorax. Metanotum with anterior margin usually strong medially ; suspcnsurial 
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, /'.pi- 
sternum with anterior margin not ridge-like ; precoxal ridge absent ; epimeron produced 
posteriorly. Metathoracic spiracle similar to mesothoracic one. Dorsospiracular setae absenl 
Postmetaspiracular setae : 0-3 (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. Metasternum represented by a weak trans 
verse plate. Anterior and posterior metasternal setae absent. 

Wings hyaline ; long (length 2350-2530, average 2490 u.) and of medium width (890-950, 
average 911 [x), ratio width to length being 1 : 2-64-2-77 (average 2-68) ; alar lobe present ; 
alar setae absent. Halteres well developed, 163-228 (average 202) y. long and 42-61 (average 
52) (jl wide, each with 1-4 (average 23) apically hooked setae which are about 97 fi 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 165). Length of segments (in jjl) : 

Leg 
I 

II 

III 

Coxae 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. Trochanters 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 



Coxa 


Trochanter 


Femur 


Tibia 


Tarsus 


Claw 


Total 


106-137 


114-186 


334-4 2 6 


562-760 


194-266 


38-48 


1379-1804 


("9) 


(171) 


(389) 


" (678) 


(242) 


(45) 


(1643) 


106-141 


114-186 


308-388 


543-749 


179-251 


40-49 


1321-1763 


(124) 


(164) 


(353) 


(669) 


(227) 


(46) 


(1584) 


1 14-156 


160-198 


334-4 J 9 


608-798 


188-304 


40-49 


1 444-1 908 


(136) 


(176) 


(384) 


(734) 


(249) 


(47) 


(1726) 



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 -5-1 -o (average 1 7) times as long as width of trochanter. Femora of medium width (60-87 t^ 
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 \l 
wide, the ratio width to length of hind tibia being 1 : 16-2-18-2 (average 171) ; 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. Tarsi 29-38 y. wide, hind tarsus 6-2-8-9 (average 7-3) times 
longer than wide ; each with 12-19 IS - an d 8-15 h.s. ; 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) \i long and 600-780 (average 681) \x wide. 

Segments I-VII : tergites on segments II— II I represented by a small sclerite on each side, and 
on VII by a weak transverse plate ; sternites 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), 
1-3 (average 1-7) and 3 on segments III-VI respectively, and ventropleural setae : usually 
one on each of segments IV-VI. Segment VII with 4 h.s. 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 i-i^ 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 short, about \ total body length (ratio 1 : 5-26-6-20, average 
5-81), 441-536 (average 492) (x long and 61-68 (average 65) jjl wide ; lateral sclerotizations not 
joined anterior to anus ; length of basal rod 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 u.), penial sheath and basisternum longer, ratios being 
1 : 2-8-3-2 (average 3) and 1 : 1-81-2-44 (average 2-17) respectively. 

Material examined : 10 specimens, collected by N. S. Borchsenius on Ligustrum 
sp. in China on 14.ix.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) y. ; width at mesothorax 455-500 
(average 477) \i. Wing expanse 3140-3220 (average 3172) [x. 

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) \i, width across genae 
277-304 (average 286) [i. 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. Midcranial 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) \x in diameter and 6-8-7-8 (average 7-2) times as much apart ; 



OF THE FAMILY COCCIDAF 



7i 



those of the ventral eyes 19-23 (average 21) y. in diameter and 2-8-3-3 (average 3-1) times as 
much apart. Ocellus small. Ocular sclerite well sclerotized and polygonally reticulated 
throughout. Preocular ridge with ventral part reaching midcranial ridge. Postocular 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. Interocular 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 10-segmented, filiform ; 1 003-1 144 (average 1101) \i long, i.e. longer than half body- 
length (ratio 1 : 1-62-1-74, average i-68), longer than posterior leg (ratio 1 : 1-16-1-19, average 
1-17) and longer than penial sheath (1 : 3-06-3-40, average 318). Scape 61-72 (average 66) \l 
long and 57-63 (average 60) [i wide, with 3-6 (average 4-1) h.s. Pedicel with distinct, poly- 
gonal, dorsal reticulation ; 57-68 (average 64) [i long and 53-63 (average 59) u. 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 jx long and 38-46, 
average 43 ji. 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 05) times as long as width of 
segment III ; with 1—3 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths of 




Fig. 20. Genus A, dorsal and ventral view. 



72 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



these segments (in y.) 137-164 (average 149), 141-175 (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 u, ; 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 77), 4-6 (average 4-6), 3-5 (average 4) and 0-4 
(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) u. long and 
57-68 (average 63) u. wide ; carrying 5-10 (average 7-3) f.s., 0-2 (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) \x long. 

Prothorax. Pronotal ridges strong, but medially interrupted by weak sclerotization. 
Lateral pronotal sclerites small, without setae. Medial pronotal setae absent. Post-tergites 
small, without striations and without setae. Pleural structures typical of family. Sternum 
with strong transverse ridge, a median ridge which is occasionally interrupted in the middle, 
and a well sclerotized triangular sclerite. Anteprosternal setae absent ; prostemal setae : 
occasionally one h.s. on each side. 

Mesothorax. Mesoprephragma with shallow emargination. Prescutum more than twice as 
wide as long (average 228 and 90 a. 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 u.) ; with 9-13 (average 12) h.s., usually with one on each side situated posterolateral!}' 
immediately outside the membranous area. Scutellum large, 1 10-125 (average 118) u. long and 
201-235 (average 220) u. wide, the ratio being 1 : 1-8-2-1 (average 1-9) ; not tubular ; with 
0-2 (average 1 -6) h.s. Postnotum with anterior margin irregular and partly overlapped by 
metathoracic fold ; postnotal apophysis and postalare well developed, the latter densely 
reticulated distally. Mesopostphragma with 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. 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. Latero- 
pleurite partly bounded anteriorly by an extension from marginal ridge. Basisternum large, 
about 262 (x wide and 183 u. 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. Mesothoracic spiracle with well developed 
peritreme ; postmesospiracular setae absent. Tegula small, membranous bulge with a weak 
sclerite posteriorly and 7-1 1 (average 8-8) h.s. Third axillary wing sclerite with a large ventral 
projection at its base. Additional sclerite elongated, well defined. Antemetaspiracular setae 
absent. 




Fig. 21. Genus A, lateral view. 



OF I HE 1 \UII.Y COCCIDAE 73 

Metathorax. Metanotum with anterior margin usually strong medially ; suspensorial 
sclerites small, spot-like ; a small, additional sclerite occasionally present anterior to post- 
niitum. 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 ridge absent. Jipl- 
sternum ventrally interrupted so that a small, separate sclerite is formed ; anterior margin not 
ridge-like. Epimeron produced posteriorly. Metatkoracic spiracle similar to mesothoracic "i><- 
Dorsospiracular setae absent. Postrnetaspiraatlar 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 131*4 |i) and broad (610-655, average 
630 jji), the ratio width to length being 1 : 2*17— 2-26 (average 221) ; alar lobe present ; alar 
setae absent. H alter es well developed, 1 j 5 144, (average 136) u. long and 30-36 (average 34) y. 
w ide, each with one apii ally hooked seta which is about 70 y. long 

/. (•;'.? moderately long and slender, with fore pair shortest and the middle or bind pair longest . 
ratio length of hind leg to body length is 1 : 1-82— 2-01 (average [-89) Length of segments 
(in |i) : 

Leg 

I 



Coxa 


troi banter 


lemur 


Tibia 


tarsus 


Claw 


rotal 


76-87 


s 7 '03 


209-224 


338 < s i 


M IO3 


25-27 


842 020 


(82) 


(99) 


(217) 


(364) 


(IOO) 


(26) 


(888) 


76-91 


93-106 


[91-231 


<7«> 44° 


IO3-I IO 


-5 29 


s,, 5-954 


(87) 


(102) 


(209) 


(395) 


(10 


(27) 


(928) 


7 6-y 1 


01 114 


[92 227 


3.57 1'- 


to8 112 


25-27 


859 969 


(87) 


(106) 


(215) 


(390) 


(no) 


(26) 


(027) 



III 

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 Trochanters 30-40 y. wide ; with 
6 oval sensilla, 5— 10 f.s. anil 6—11 lis., 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 y.), ratio width 
to length of hind femur being 1 : 32-3-8 (average 35) ; each with 32-54 f.s. and 15-31 h.s. 
Tibiae 30-34 y. 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 5 as long as width of tibia ; apical spur about the same size on all tibiae 
Tarsi 25-30 y wide, hind tarsus 3-6-4-1 (average 3-7) times longer than wide ; each with 
21-40 h.s., with 0-5 (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) u. long and 430-510 (average 450) y 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 dorsopleural setae : 0-2 (average i-6), 1-4 (average 2-1), 2-4 (average 3-3) and 
0-4 (average 2-9) on segments III-YI respectively, and ventropleural setae : sometimes one on 
each of segments [V— 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 VII ; 
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. No IXth tergite observed. A nte-anal setae absent. Posterior margin with 3 h.s. 
on each side. 



74 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



Genital segment. Penial sheath of medium length, about \ total body length (ratio i : 4-80- 
5-32, average 5-04), 315-369 (average 349) u. long and 53-61 (average 57) y. wide ; lateral 
sclerotizations narrowly joined anterior to anus ; length of basal rod about f 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 (j.) and broad, penial sheath and 
basisternum longer, ratios being 1 : 2-4-2-6 (average 2-5) and 1 : 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) y. ; width at mesothorax 360-430 (average 408) y.. Wing expanse 
2750-2940 (average 2816) y. 




Fig. 22. Sphaerolecanium prunastri (Fonsc), dorsal and ventral view. 



OF THIi FAMILY COCCIDAE 



75 



Head subconica! in dorsal view ; in lateral view obliquely elongated dorsoventrally, with, 
anterodorsal bulge somewhat pronounced ; length from apex to pronotal ridge 205-247 (average 
222) \i, width across genae 243-266 (average 253) li. Median crest sclerotized and distinctly 
polygonally reticulated ; with 9-12 (average 10) 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. 
M idcranial 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) y. in diameter and 3-4-4-2 (average 3-9) times as much apart ; those of the ventral 
eyes 19-27 (average 26) \l 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. Postocular ridge well developed 
dorsally and lateroventrally, but weak posteromedially ; below ocellus the ridge splits up, 
with anterior branch partly surrounding ocellus. Interocular ridge absent, but sometimes 
indicated by a line of dark sclerotization. I tarsal 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. Preoral 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 10-segmented, filiform ; 671-809 (average 770) pi long, i.e. shorter than half body 
length (ratio 1 : 2-13-2-45, average 2-28), longer than posterior le^ (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) la long 
and 46-49 (average 47) (x wide, with 3-4 (average 3-4) h.s. Pedicel with we.ik. polygonal, 
dorsal reticulation ; 55-68 (average 63) \i long and 32 42 (average 38) u, wide, with 0-3 
(average 1-2) f.s., 4-9 (average 6-8) h.s. and a sensilluni rjlacodeum. Segment III < tub-shaped, 
2 '7-3 '7 (average 3-1) times longer than wide (68-86, average 78 ■>- long and 23 jo, average 
26 (x wide) ; with 4—7 (average 6) f.s. of medium length, 1-1-1-3 (average 12) times as long as 
width of segment ; with i— 3 usual sensill.i basiconica Segments IV [X cylindrical ; lengths oi 
these segments (in p.) 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 (x, with distal segments wider than proximal ones ; with 18-27 (average 21), [3 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. ; antenual bristles on segments VIII l\ not distinctly different 
from f.s., sometimes slightly thicker. Segment X : terminal part not constricted ; 57-72 
(average 65) \x long and 21-27 (average 23) \i wide ; carrying o 15 (average 12) f.s., 3 capitate 
subapical setae and 5 antennal bristles of which the ; long ones are about half as long as the 
segment and the 2 shorter ones not distin< try different from the f.s. ; with 2 sensilla basiconica 
ventrally, one near apex and the other more proximal 

Thorax 513-597 (average 566) la 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- 




^AOO\i 



F'ig. 2^. Sphaerolecanium prunastri (Fonsc), lateral view. 



76 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

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

Mesothorax. Mesoprephragma with shallow emargination. Prescutum more than twice as 
wide as long (average 181 and 82 \j. 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 subrectangular ; 72-91 
(average 83) [x long and 1-92-2-48 (average 2-11) times as wide (width 152-220, average 175 (x) ; 
with 2-4 (average 2-8) h.s. Scutellum 65-80 (average 74) \x long and 141-182 (average 163) [jl 
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 ; postnatal apophysis and postalare 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. 
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. Eater opleurite not bounded by an extension from marginal ridge. Basistemum large, 
about 236 u. wide and 170 a. 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. Mesothoracic spiracle with well developed peritreme ; 
postmesospiracular setae absent. Tegula small, membranous bulge with a small weak sclerite 
posteriorly and 4-10 (average 7-3) h.s. Third axillary wing sclerite with a pronounced ventral 
projection at its base. Additional sclerite well defined. Antemetaspiracular setae absent. 

Metathorax. 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 precoxal ridge present. Episternum reduced to a small 
subtriangular plate ; epimeron produced posteriorly. Metathoracic spiracle similar to meso- 
thoracic one. Dorsospiracular setae absent. Postmetaspiracular setae : 0—15 (average 7-5) f.s. 
and 1-3 (average 1-2) h.s. Metasternum 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 u.) and of medium width (500-530, 
average 516 u.), the ratio width to length being 1 : 2-34-2-42 (average 2-38) ; alar lobe and 
alar setae absent. H alter es 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 u.) : 



Leg 


Coxa 


Trochanter 


Femur 


Tibia 


Tarsus 


Claw 


Total 


I 


59-72 


76-95 


175-209 


239-293 


91-110 


26-28 


667-803 




(67) 


(87) 


(i97) 


(280) 


(103) 


(27) 


(759) 


II 


62-76 


84-103 


160-190 


251-304 


106-118 


27-30 


690-813 




(70) 


(95) 


(i79) 


(281) 


(113) 


(28) 


(766) 


III 


65-82 


84-106 


171-201 


255-317 


101-125 


27-32 


701-851 




(76) 


(100) 


(189) 


(295) 


(116) 


(30) 


(805) 



Coxae 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 § as long as trochanter. Trochanters 30-34 y. 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 I II E I- \M I I.Y C( >C< [ DAE 77 

outer margin and a long apical seta which, on the lore trochanter, ism 2-<> (average 2-4! tunes 
as long as width of trochanter. Femora very wide (46-57 |x), ratio width to length of hind 
femur being 1 : 3*3— 3-8 (average V5) i each with 8-16 f.s. and 17-28 lis. Tibiae 23—30 (x 
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 lis. and 10-26 f.s., the latter about .is long as width of tibia ; apical 
spur about the same size on all tibiae. Tarsi 21—25 ! x wl( l e . hind tarsus 4-8-5-5 (average 5-2) 
times longer than wide ; each with 1-6 f.s. and 15 27 lis. ; tarsal digitules subequal, longer 
than claw. Claws of medium length, longer than width of tarsus ; slightly curved, with small 
denticle near tip ; ungual digitules subequal, about as long as 1 l.iw 

Ibdomen 470-530 (average 498) |x long and 320-420 (average $75) \t wide. 

Segments I VII : tergites represented by a small sclerite on each side on anterior margin ol 
segments II ill and by a weak plate on V] VII ; sternites represented l>v 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 : lis only, usually one on each side on I and each 
of segments V V 1 1 . Pleural setae on segments III VI consisting of 0—1 (average o-i), 0-1 
(average 01), 0-2 (average 0-5) and z (average o-q) f.s , and _• (average 0-7), 0-3 (avi 
1 - s ) , 1 3 (average 21) and 1 4 (average 1 <i) hair-like dorsopleural pectivel) . ventre 

pleural setae absent or incorporated into the dorsopleural group. Segment VII with 2 
(average i- 1) f.s. and 3 5 (average 5-4) h s. Ventral setae : 1 5 (average 2 -3) f.s. on segment II, 
occasionally one on [II, and usually one h.s. on each side on II [V and 1 on each of segments 
V VII. 

Segment V 1 1 1 with transverse tergite and sternite, and with .1 large, cylindrii al, 1 audal exten- 
sion ; glandular pouch with 2 long, pointed setae, whose protruding part is z\] times as long 
as section within pouch. No lXih tergite observed lute aual 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 42 385, average 
3-68), 433-517 (average 473) |x long and 46-51 (average [8) [i wide ; lateral sclerotizations 
narrowly joined anterior to anus ; length of basal rod 1 ,' -' tunes that of aedeagUS and extend- 
ing anteriorlv 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 30 
(average ^z) small setae ; a cluster of small sensilla occurring vcntrally near apex ol sheath 
Aedeagus of medium length (1 14 152, average 140) [x, penial sheath and basisternum longer, the 
ratios being 1 : 3-13-8 (average ••,■4) and 1 : 115 132 (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 X. S. Borchscnius (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) \i ; width at mesothorax 260-290 (average 281) fx. Wing expanse 
2730-2930 (average 2839) [x. 



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) \l. Median crest sclerotized, distinctly polygonally 
reticulated, with 7-1 1 (average 9) hair-like dorsal head setae. Midcranial 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) |j. in diameter and 2-7-3-4 (average 3) times as much apart ; those of the 
ventral eyes 19-25 (average 22) [x 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. Preocular ridge long, the 
ridges of each side ventrally meeting or almost meeting each other medially. Postocular ridge 
well developed throughout. Interocular ridge broadly joining pre- and postocular ridges below 
ocellus. Dorsal ocular setae absent. V entral 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 
(0-3, 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. Preoral ridge absent. Tendon-like apodeme variable, 
occasionally fully developed, but usually reduced or absent. Cranial apophysis of medium 
length ; apex trifurcate, extending to around the margin of the posterior level of the ventral 
eyes. Mouth opening irregular. A nterior tentorial pits present, situated anterolateral to mouth 
opening. 

Antennae 10-segmented, filiform ; 1030-1284 (average 1125) [x long, i.e. longer than half 
body length (ratio 1 : 1-23-1-51, average 1-38), longer than posterior leg (ratio 1 : 1-42-1-52, 




Fig. 24. Eriopeltis sp., dorsal and ventral view. 



<>l THE 1\M 1 LY COCCIDAE 



79 



average i 46) and longer than penial sheath (ratio 1 : 304-3-86, average 330). Scape 44 53 
(average 48) \j. long and 45-50 (average 46) jjl wide, with 3 h.s. Pedicel with distinct, poly- 
gonal, dorsal reticulation ; 49-61 (average 55) \± long and 34-42 (average 37) jx wide ; with 
3-6 (average 42) f.s., 1 5 (average 2-g) h.s. and a sensilhun placodeum. Segment III somewhat 
club-shaped with a rugose surface ; 40-4-9 (average 4-6) times longer than wide (99—125, 
average 114 u. long and 23-27, average 25 u. wide) ; with 5-11 (average 7-7) f.s. of medium 
length, 1-3-1-7 (average 15) times longer than width of segment ; with one or two usual 
sensilla basiconica. Segments I \' IX cylindrical, rugose ; lengths of these segments (in jjl) 
141-198 (average 162), 152-205 (average 171), 141-194 (average 159), 114-171 (average 135), 
95 1 18 (average 106) and 84-95 (average 90) respectively, all of about the same width, varying 
from 19 to 23 (i. ; with 14-23 (average 20), 20-30 (average 24), 19-28 (average 23), 14-23 
(average 20), 14-20 (average 18) and 13 10 (average i<>) I s respei tively, but no h.s. ; antennal 
bristles on segments VIII— IX not distinctly different from the l s Segment X : terminal part 
not constricted ; 80 95 (average 86) u. long and [9 21 (average 19) ;i wide ; carrying 10 14 
(average 1 1 -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 .is long as the f.s. ; 
with 2 sensilla basiconica ventrally, one near apex and the other more proximal. 

Thorax 448-486 (average 469) [i. long. 

Prothorax. Pronotal ridge strong, but medially interrupted l>v 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 rid^e only represented on anterior 
part of well sclerotized triangular sclerite, Anteprostemal setae absenl : prosternal setae : 
1-7 (average 4-2) f.s., occurring medially around anterior part of the sternum 

Mesothorax. Mesoprephragma with shallow emargination. Prescutum 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 polygonallv retu 11 
lated ; subrectangular, 65-76 (average 73) u. long and 147 1-79 (average 1-62) times as wide 
(width 106-129, average 1 17 u.) ; with 4-10 (average 7) h.s. Anterior arms of scutum irregularly 
reticulated. Scutellum 38-46 (average 41) u. long and 114 137 (average 127) ix wide, ratio 
being 1 : 2-8-3-5 (average 31) ; tubular, with a small ventral foramen ; without setae /'<>-./- 
notum with anterior margin irregular, weakly sclerotized and not overlapped by metathorat ii 
fold ; postnotal apophysis and postalare well developed, the latter densely reticulated distallv. 
Mesopostphragma short, with shallow emargination. Mesopleuron. Mesopleural ridge strong, 
but becoming weaker immediately above coxa! articulation ; pleural apophysis and pleural 
wing process well developed, the latter connected with episternum by a narrow hasalare. 
Subalare small. Episternum without polygonal reticulation ; sube pi sternal ridge becoming 
broader ventrally, but below membranous cleft indistinct and only marked by a band of dark 
sclerotization. Epuueron very small. I.aterop/eiiritc partly bounded anteriorly by an exten- 




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



80 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

sion from marginal ridge. Basistemum large, about 176 \i wide and 130 y. 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 basistemum ; marginal 
and precoxal ridges well developed ; 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 small, rounded ventral projection at its base. Additional sclerite 
well sclerotized. Antemetaspiracular setae absent. 

Metathorax. 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 ridge considerably reduced, extending only a short 
distance above coxal articulation. Episternum reduced to a small subtriangular plate, epi- 
meron produced posteriorly. Metathoracic spiracle similar to mesothoracic one. Dorsospiracular 
setae absent. Postmetaspiracular setae : 5—13 (average 9-1) f.s. and 0-2 (average 0-4) h.s. 
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 0-3 (average 
o-6) h.s. ; posterior metasternal setae : 8-15 (average 11) f.s. 

Wings hyaline ; long (1 255-1 340, average 1303 \x) and narrow (width 395-430, average 413 (i.), 
ratio width to length being 1 : 3-02-3-29 (average 3-16) ; alar lobe and alar setae absent. 
H alter es 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 (x) : 

Leg 
I 

II 

III 

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. Trochanters 25-30 u. 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 \i), ratio width to 
length of hind femur being 1 : 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. Tibiae 19-23 y. 
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 ns - an d J 7-33 Is -. the latter about 1 1 times longer than width of tibia ; 
apical spur about the same size on all tibiae. Tarsus 19-23 \i 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 i| 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) |j. long and 240-300 (average 270) \l wide. 

Segments I -VI I : 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 : 0-2 (average 0-4), 0-3 (average i-i), 3-10 (average 6-3), 3-10 
(average 5-9), 2-5 (average 3-2) and 0-4 (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 



Coxa 


Trochanter 


Femur 


Tibia 


Tarsus 


Claw 


Total 


59-68 


76-84 


186-198 


272-319 


110-122 


30-34 


736-806 


(61) 


(79) 


(I9i) 


(292) 


(116) 


(33) 


(772) 


57- 6 7 


72-80 


175-198 


270-315 


103-114 


28-31 


711-789 


(60) 


(75) 


(183) 


(291) 


(109) 


(29) 


(748) 


61-68 


72-84 


177-217 


277-323 


99-114 


30-34 


722-809 


(66) 


(76) 


(183) 


(296) 


(106) 


(32) 


(758) 



OF THE FAMILY COCCIDAL 81 

I— III and on IV— VI represented by h.s. only, which include the dorsoplenral setae : usually- 
one on IV and 2 on segments V-VI, and ventropleural setae : occasionally one on \ r and usually 
one on segment VI. Segment VII with 0-6 (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 n) 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 0-2 (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 IXth tergite 
present. Ante-anal setae : 4-9 (average 5-8) f.s. and an occasional h.s. Posterior margin 
with 3-4 (average 31) h.s. on each side. 

Genital segment. Penial sheath of medium length, about j— j total body length (ratio 
1 : 4-5-4-9, average 4-7), 314-342 (average 332) u. long and 29-36 (average 32) ;j. wide ; lateral 
sclerotizations narrowly joined anterior to anus ; basal rod about 1 .J times as long as aedeagus, 
extending anteriorly from base of aedeagus to between £ and J 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 u. 
long), penial sheath and basisternum longer, the ratios being 1 : 4-14-9 (average 4-5) and 
1 : 1-57-1-04 (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 Yfestucae (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 i486) pt ; width at mesothorax 280-310 (average 293) [i. Wing expanse 
2730-2920 (average 2820) fjt. 

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) 
[x, 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) jx in diameter and 3-3-4-2 (average 3-7) times as much apart ; 
those of the ventral eyes 19-23 (average 21) (i. 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. Pre- 
ocular ridge long, the ridges of each side ventrally meeting or almost meeting each other medially. 
Postocular ridge well developed and distinct throughout. Interocular 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 (0-2, 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. Preoral ridge absent. 



82 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



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

Antennae io-segmented, filiform ; 996-1060 (average 1018) y. long, i.e. longer than half body 
length (ratio 1 : 1 -37-1 -52, average 1-45), longer than posterior leg (ratio 1 : 1 -38-1 45, average 
1-41) and longer than penial sheath (ratio 1 : 3-31-3-49, average 3-41). Scape 42-49 (average 47) 
[x long and 44-48 (average 46) (x wide, with 3 h.s. Pedicel with distinct, polygonal, dorsal 
reticulation ; 49-69 (average 57) pt long and 38-42 (average 39) u. 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, with a 
rugose surface ; 4-3-6-0 (average 5) times longer than wide (99-118, average 113 \l long and 
19-25, average 22 \i 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 IV-IX 
cylindrical, rugose ; lengths of these segments (in jj.) 137-156 (average 148), 133-156 (average 
J 45). I 33 _I 63 (average 146), 103-118 (average in), 84-99 (average 90) and 72-87 (average 82) 
respectively, all of about the same width, varying from 19 to 23 (x and with 13-21 (average 18), 
19-23 (average 21), 20-32 (average 26), 18-27 (average 22), 13-25 (average 18) and n-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) y. long and 
19-22 (average 20) \i 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 f 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) \i long. 




Fig. 26. Eriopeltis Pfestucae (Fonsc), dorsal and ventral view. 



01 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. Sternum with strong transverse 
ridge, usually somewhat narrowed medially ; median ridge sometimes complete but usually 
interrupted ; triangular sclerite well sclerotized. A nteprosternal setae absent ; prosternal setae 
consisting of 0-2 (average 07) f.s. occurring medially around anterior part of sternum. 

Mesothorax. Mesoprephragma without emargination. Prescutum about ii times as wide as 
long (average 125 and 81 (x 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 membranoits area without reticulation ; 
subrectangular, 61-72 (average 64) (i. long and 1-47-2-00 (average 1-71) times as wide (width 
99-122, average 109 u-) ; with 2-10 (average 6) h.s. Anterior arms of scutum not reticulated. 
Scutellum 42-49 (average 45) \i long and 1 14-129 (average 122) u. wide, the ratio being 
1 : 2-5-3-0 (average 27) ; 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. Mesopostphragrna 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 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. Basis ternum large, about 171 [i wide and 128 u. 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 halt 
the length of the basisternum ; marginal and precoxal ridges well developed ; without setae. 
h'urca well developed. Mesothoracic spiracle with well developed peritreme ; postmesospiraculai 
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. Antemetaspiracular 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. 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. Dorsospiracular setae absent. 
Postmetaspiracular setae : 8-1 1 (average 92) f.s. and 0-1 (average 03) 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 pi) and narrow (width 400-460, average 422 u.), 
the ratio being 1 : 2-87-3-15 (average 3-04) ; alar 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 


no- 125 


29-34 


726-775 




(62) 


(80) 


(i9o) 


(277) 


("4)" 


(32) 


(754) 


II 


61-68 


72-80 


171-190 


251-274 


106-122 


27-32 


695-751 




(62) 


(76) 


(180) 


(262) 


(in) 


(29) 


(720) 


III 


61-68 


72-78 


167-186 


251-274 


99-108 


30-32 


686-731 




(64) 


(75) 


(177) 


(265) 


(104) 


(3i) 


(7i4) 



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. Trochanters 26-30 [i wide ; with 6 oval sensilla, 5-12 f.s. and 
7-1 1 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 3-1-5-0 (average 4-1) times as long as 
width of trochanter. Femora of medium width (34-42 u.), 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. Tibiae 21-25 ^ 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 i£ times as long as width of tibia ; apical 
spur about the same size on all tibiae. Tarsi 19-23 |j. wide, hind tarsus 4-3-4-8 (average 4-6) 
times longer than wide ; each with 0-3 f.s. and 21-30 h.s. ; tarsal digitules subequal, slightly 
longer than claw. Claws long, about 1 J 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) y. long and 250-300 (average 272) jx wide. 

Segments I-VII : tergites absent ; sternites 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 : 0-3 (average o-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. 

Genital segment. Penial sheath of medium length, about \ total body length (ratio 1 : 4 53- 
5-16 (average 4-95), 289-308 (average 300) ;x long and 30-36 (average 34) \i wide, lateral sclero- 
tizations narrowly joined anterior to anus ; basal rod 1^—2 times as long as aedeagus, extending 
anteriorly from the base of the latter for ^— f 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 65 fi. long), penial sheath and 
basisternum longer, ratios being 1 : 4-3-5-1 (average 4-7) and 1 : 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 Eriopeltis 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 COCCIDAF 



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) \x ; width at mesothorax 240-270 (average 255) \i. Wing expanse 
2090-2350 (average 2213) \l. 

Head subcorneal 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) 
\x, width across genae 167-182 (average 175) jji. 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) \i in diameter and 40-4-5 (average 42) times as much 
apart ; those of the ventral eyes 15-20 (average 16) jx in diameter and 2-2-2-5 (average 23) 
times as much apart. Ocellus small. Ocular sclerite well sclerotized, distinctly polygonally 
reticulated throughout. Preocular ridge long, the ridges of each side ventrally meeting or 
almost meeting each other medially. Postocular ridge well developed, but tapering dorsally. 
fnlerocular 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 (0-6, 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. 




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



86 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

Preoral ridge absent. Tendon-like apodeme short. Cranial 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 io-segmented, filiform ; 729-901 (average 822) y. long, i.e. longer than half body 
length (ratio 1 : 1-28-1-56, average 139), longer than posterior leg (ratio 1 : 1-36—1-40, average 
1-38) and longer than penial sheath (ratio 1 : 4-36-6-02, average 4-87). Scape 36-46 (average 
41) u. long and 36-40 (average 38) y. wide, with 3 h.s. and occasionally one f.s. Pedicel with 
weak, polygonal, dorsal reticulation ; 42-49 (average 45) y. long and 31-34 (average 33) y. 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 jx long and 23 \i wide) ; 
with 0-2 (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 \i) 1 14-158 (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 u., 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) \i long and 19-23 (average 20) [i 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) \x long. 

Prothorax. 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. Sternum 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. 
A ntepro sternal 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 0-1 (average 0-5) h.s., none of these occurring 
anterior to spiracles. 

Mesothorax. Mesoprepkragma short, with shallow emargination. Prescutum about twice as 
wide as long (average 116 and 60 y. respectively) ; anterior margin curved ; laterally bounded 
by strong prescutal ridges and posteriorly by the prescutal suture ; slightly more heavily sclero- 
tized medially ; polygonally reticulated. Scutum. Median membranous area subrectangular ; 
49-72 (average 60) ji long and 1-42-1-92 (average 1-70) times as wide (width 97-114, average 
102 u.) ; with 3-8 (average 4-7) h.s. Anterior arms of scutum irregularly reticulated. 
Scutellum 29-34 (average 32) |j. long and 95-114 (average 102) [t wide, ratio being 1 : 30-3-8 
(average 3-4) ; tubular, with small ventral foramen ; usually without setae. Postnotum with 




Fig. 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. Subalare small. Episternit»i 
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. Basi- 
sternum large, about 163 fx wide and 91 [x 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. Fnrca 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. 
A ntemetaspiracular setae absent. 

Metathorax. 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. Dorsospiracular setae absent . 
Postmetaspiracular setae 6-14 (average 10) f.s. and 0-2 (average 07) h.s. Metasternal plate 
weak, but more heavily sclerotized around anterior margin. Anterior metasternal setae : 
16-23 (average 19) f.s. and no h.s. ; posterior metasternal setae : 7-19 (average 13) f.s. 

Wings hyaline ; long (910-1050, average 969 |x) and narrow (2<>5-36o, average 336 u, wide), 
ratio width to length being 1 : 280-301 (average 2 90) ; alar lobe and alar setae absent. 
Halteres absent. 

Legs short and moderately slender, all 3 pairs subequal in size, fore legs being only slight Iv 
shorter ; ratio length of hind leg to body length is 1 : 1-89-2-01 (average 198). Length of 
segments (in jx) : 



Leg 


Coxa 


Trochanter 


Lemur 


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) 


11 


49-60 


62-72 


129-148 


186-219 


84-89 


26-28 


540-616 




(54) 


(69) 


(139) 


(206) 


(86) 


(27) 


" (578) 


III 


46-57 


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. Trochanters 21-24 V- 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) jx, ratio width to 
length of hind femur being 1 : 41-50 (average 4-6) ; each with 8-16 f.s. and 9-17 h.s. Tibiae 
17-23 (x wide, ratio width to length of hind tibia being 1 : 9-1-11-6 (average 105) ; each with 
20-39 h.s. and 8-22 f.s., the latter about 1 \ times longer than width of tibia ; apical spur about 
the same size on all tibiae. Tarsi 19-21 u. wide, hind tarsus 4-0-4-3 (average 42) times longer 
than wide ; each with 2-6 f.s. and 17-23 h.s. ; 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) pi long and 200-260 (average 227) u. wide. 

Segments I-VII : tergites represented by a weak sclerite on or near anterior margin of each 
segment ; sternites 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 o-6) and V-VII (up to 3) ; usually one h.s. on I and each of 
segments IV— VII. Pleural setae : absent on segments I, II and usually III, on segments IV-VI 
consisting of dorsopleural setae : usually 2 h.s. on each of segments IV-VI, and ventropleural setae : 
usually one h.s. and occasionally one f.s. on segments V-VI. Segment VII with 4-1 1 (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), 0-6 (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 ; glandular 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 | to | total body length (ratio 1 : 6-1-7 -3, 
average 67), 152-186 (average 173) [x long and 30-38 (average 34) [/. wide ; lateral sclerotiza- 
tions narrowly joined anterior to anus ; basal rod about as long as aedeagus, extending anter- 
iorly from base of aedeagus to between £ and f 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 u. long), penial sheath and basisternum 
longer, the ratios being 1 : 3-2-4-3 (average 3-8) and 1 : 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 INGLISIA 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) u. ; width at mesothorax 370-420 (average 399) y.. 
Wing expanse 2870-3270 (average 3032) \i. 

Head subcorneal in dorsal view ; in lateral view obliquely elongated, with the anterodorsal 
bulge not pronounced ; length from apex to pronotal ridge 209-262 (average 245) u., width 
across genae 251-277 (average 266) y.. Median crest well sclerotized, especially around the 
posterior margin, which is obtuse ; weakly polygonally reticulated ; with 4-1 1 (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 5-1 1 (average 7) fleshy and 0-4 (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) u. in diameter 
and 1 -6-2-1 (average i-8) times as much apart ; those of the ventral eyes 30-36 (average 34) y. 
in diameter and o-6-o-8 (average 0-7) times as much apart. Ocellus small. Ocular sclerite 



OK THE FAMILY COCCIDAE 



89 



weakly sclerotized ; reticulated only dorsally and between and anterior to the ventral eyes. 
Preocular ridge with ventral part not extending far below articular process. Postocidar ridge 
well developed throughout ; below ocellus the ridge splits up, completely enclosing the ocellus. 
1 nterocular 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 7-1 1 (average 
9) h.s., of which 1-4 (average 2) f.s. and 0-3 (average 11) h.s. occur anteriorly, around the 
midcranial ridge, and 1-4 (average 3) f.s. and 0-2 (average 1) h.s. between and behind the 
ventral eyes. Preoral 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 10-segmented, filiform ; 903-1047 (average 986) (z long, i.e. longer than half body 
length (ratio 1 : 1-67-1-82, average 173), longer than posterior leg (ratio 1 : 107-113, average 
i 10) and longer than penial sheath (ratio 1 : 2-53-2-78, average 2-66). Scape 46-53 (average 
47) u, long and 49-53 (average 51) (x wide, with 3 h.s. Pedicel not reticulated ; 38-48 (average 
43) (i long and 34-38 jj. wide ; with 1-4 (average 2) f.s., 4-9 (average 63) h.s. and a sensillum 
placodeum. Segment III somewhat club-shaped, 3-6-4-5 (average 4) times longer than wide 
(103-110, average 107 jjl long and 25-29, average 27 u, wide) ; with 1-4 (average 22) h.s. and 
6-12 (average 10) f.s., the latter of medium width, o-8-i-i (average 09) times as long as width 
of segment ; with 1-3 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths of these 
segments (in \x) 114-141 (average 127), 114-137 (average 127), 125-152 (average 138), 1 16-129 
(average 120), 89-122 (average 105) and 76-95 (average 82) respectively, all of about the same 




Fig. 29. Inglisia theobromae (Xewst.), dorsal and ventral view. 



go 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



width, varying from 17 to 25 (x ; 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 90) [i. long and 
42-46 (average 43) \x 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) \i long. 

Prothorax. Pronotal 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. Sternum with transverse ridge strong, median ridge reduced to a short basal stalk and 
triangular sclerite bearing numerous small projections. A nteprosternal setae absent ; prosternal 
setae : 5-1 1 (average 7-8) f.s. and 0-2 (average 0-9) h.s., the setae not occurring in front of 
spiracles. 

Mesothorax. Mesoprephragma with shallow emargination. Prescutum about twice as wide 
as long (average 171 and 90 jx 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) y. long and 2-10-2-88 (average 2-51) times as wide (width 156-179, average 171 (z) ; 
with 3-15 (average 6-6) f.s. and 7-15 (average 12) h.s. Scutellum 53-80 (average 70) (x long 
and 144-175 (average 163) jx wide, the ratio being 1 : 2-1-2-9 (average 24) ; not tubular. 
Postnotum with anterior margin irregular and usually 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, not interrupted 
above coxal articulation ; pleural 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 bound anteriorly by 
an extension from marginal ridge. Basisternum large, about 253 u. wide and 162 u. 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 spiracle with well developed peritreme ; 
postmesospiracular setae : 20-28 (average 24) f.s. and 0-3 (average 1) h.s., arranged in a band 
behind the spiracles and presternum. 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 well defined. Antemetaspiracular 
setae : 1-6 (average 3-8) f.s. 




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



I arsus 


Clau 


Total 


133 164 


20 -1 5 


821 95a 


(148) 


(22) 


(888) 


150-175 


^3-^7 


787-901 


(169) 


(24) 


(860) 


160-170 


25-29 


8^7-937 


(173) 


(26) 


(896) 



OF THE FAMILY COCCIDAE 91 

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 0-2 (average o-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 posteriorlv 
Metathoracic spiracle similar to mesothoracic one. Dorsospiracular setae : 1-8 (average 3-6) 
f.s. and 0-2 (average 0-3) h.s. Postmetaspiracular setae : 6-1 1 (average 8-2) f.s. and 0-5 
(average 1 -7) h.s. Metasternum represented by a transverse plate. .-1 nterior 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 (1 270-1370, average 1333 y.) and width (540-580, av« 
561 u.), ratio width to length being 1 : 2-33-2-45 (average 2-38) ; alar lobe and alar setae absent. 
H alter es absent. 

Legs moderately long and slender, with middle pair shortest and lore or bind pair longest ; 
ratio length of hind leg to body length is 1 : 1 -87- 1 (>8 (average i<)i). Length of segments 
(in n) : 
Leg (Oxa Trochanter Femur Tibia 

1 57-61 68-84 224-255 308 -369 

(60) (80) (239) (339) 

11 57-67 65-76 201-232 285-338 

(64) (7i) (217) (315) 

111 61-72 76-84 201-232 304 357 

(67) (79) (219) (333) 

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 J as long as trochanter. Trochanters 23-29 u. wide ; with b 
oval sensilla, 7-15 f.s. and 6-9 h.s., the latter including -' minute setae near basal ridge, one small 
seta on outer margin and a long apical seta which, on the fore trochanter, is 2024 (average 
2-3) times as long as width of trochanter. Femora of medium width (36-44 u.), the ratio width 
to length of hind femur being 1 : 4-8-5-4 (average 52) ; each with 18-33 f.s. and 13-25 h.s. 
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 72) 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 ; ungual digitules subequal, 
about as long as claw. 

Abdomen 490-610 (average 558) u. long and 320-420 (average 380) u. 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 [V— 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 : 0-3 (average 0-5), 
0-2 (average o-6) and 0-2 (average o-8) f.s. on segments III and VI and 0-1 (average 0-7) 1, 1, 
1-2 (average 1-2), 0-2 (average 1-3) and 1-3 (average i-8) h.s. on segments I— VI respectively, 
and of ventropleural setae : 0-1 (average 0-4), 0-3 (average 1-5) and 0-5 (average 2-1) f.s. on 
segments IV-VI and 0-1 (average 04), 0-1 (average 0-5), 1-2 (average 12), 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. No I Xth 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) \x long and 34-40 (average 37) [x wide ; lateral 
sclerotizations not joined anterior to anus ; length of basal rod about f that of aedeagus and 
extending anteriorly from base of the aedeagus for f-f 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 \i), penial sheath longer 
and basisternum shorter, ratios being 1 : 1-9-2-1 (average 2) and 1 : 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 /. 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 

COCCUS 

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) \jl ; width at mesothorax 280-330 (average 309) [t. Wing expanse 2200-2650 
(average 2446) u.. 

Head subcorneal in dorsal view ; in lateral view obliquely elongated, with anterodorsal bulge 
pronounced ; length from apex to pronotal ridge 198-239 (average 216) |i., width across genae 
186-205 (average 196) iz. 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 49) hair-like genal 
setae. Eyes : two pairs, subequal ; corneae of dorsal eyes 21-32 (average 27) u. in diameter 
and 10-21 (average 13) times as much apart ; those of the ventral eyes 22-32 (average 29) tx 
in diameter and o-6-i-6 (average 0-9) 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 Inter ocular 
ridge absent, although the area between ocellus and preocular ridge is strongly sclerotized. 
Dorsal ocular setae : 2-8 (average 4-<)) f.s. and o 4 (average 1 -6) h a mi i\i< h side. \'ctilral 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 




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



94 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



12) f.s. between and behind the eyes. Preoral 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 10-segmented, filiform ; 576-732 (average 663) [x long, i.e. shorter than half body 
length (ratio 1 : 2-01-2-63, average 226), shorter than posterior leg (ratio 1 : 0-58-0-74, average 
0-69) and longer than penial sheath (ratio 1 : 2-10-2-43, average 2-27). Scape 38-46 (average 
43) \l long and 27-31 (average 30) jx wide, with 3-4 (average 3-1) h.s. Pedicel with wavy stria- 
tion dorsally ; 46-53 (average 50) ;x long and 27-33 (average 31) [x wide ; with 6-13 (average 
9-7) f.s., 2-7 (average 4-2) h.s. and a sensillum placodeum. Segment III somewhat club-shaped, 
3-0-3-7 (average 3-3) times longer than wide (57-72, average 60 \x long and 19-21, average 20 \i 
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 IV-IX cylindrical ; lengths 
of these segments (in [x) 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 |x, 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 £-J constricted ; 57-78 (average 69) jx long and 49-55 
(average 52) [x 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) \l long. 

Prothorax . Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral 
pronotal sclerites small, without setae. Medial pronotal setae : 2 h.s., widely separated, with 
0-1 (average 04) 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 behind the sclerite 
on each side. Pleural structures typical of the family. Sternum with strong transverse and 
median ridges and a small oval sclerite ; the area on each side of the median ridge showing 
distinct polygonal reticulation. Anteprosternal 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. 

Mesothorax. Mesoprephragma with deep emargination. Prescutum about twice as wide as 
long (average 159 and 83 jx respectively) ; anterior margin strongly curved ; laterally bounded 
by the prescutal ridges 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) (x long and 1-23-1-37 (average 1-30) times as wide (width 135-160, 
average 146 jx) ; with 10-32 (average 24) f.s. and 11-15 (average 13) h.s. Scutellum 34-42 
(average 38) jx long and 141-160 (average 154) [x wide, ratio being 1 : 3-8-4-2 (average 4) ; 
tubular, with a small ventral foramen ; setae absent. Postnotum with anterior margin weakly 




<--sm jji JijM i%<& 




L.I I .•-- /&*& 



.400u 



Fig. 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. Hasalare vestigial, incorporated into pleural 
wing process, not joining the latter with episternum. Subalare small. Epi sternum 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. Later opleurite not bounded anteriorly by an extension from 
marginal ridge. Basisternum large, about 182 ji wide and 147 u. 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. Eurca well developed. Mesothoracic 
spiracle with well developed peritreme ; postmesospiracular 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. Antemetaspiracular 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 (0-2, average 1) h.s. on each side. Pleural ridge 1 onsiderably reduced, 
extending only a short distance above coxal articulation. Vestigial precoxal ridge present. 
Episternum reduced to a small subtriangular plate ; epimeron produced posteriorly. Meta- 
thoracic spiracle similar to mesothoracic one. Dorsospiracultw 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 10) f.s. respectively. 

Wings hyaline; of medium length (980-1200, average 1097 (/), but comparatively broad 
(width 430-520, average 481 a.), ratio width to length being 1 : 2-19-2-31 (average 2-27) ; 
alar 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 : 1-58-1-65 (average 162). Length of segments (in u.) : 

Tibia 
291-369 

(341) 
■293-350 

(322) 
34^-395 

(369) 

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 78) h.s. ; fore coxa with 2-3 (average 23) capitate coxal bristles ; 
apical seta about § as long as trochanter. Trochanters 19-23 u. 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. Eemora very slender, 27-34 V- wide, ratio width to length 
of hind femur being 1 : 61-6-7 (average 6-4) ; with 27-46 f.s. and 8-14 h.s. Tibiae 17-21 (x 
wide, ratio width to length of hind tibia being 1 : 18-0-20-8 (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 (i. 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) \j. long and 230-290 (average 263) \i wide. 



-eg 


Coxa 


Trochanter 


Femur 


1 


38-49 


57-72 


171-200 




(46) 


(66) 


(190) 


11 


69-80 


76-84 


163-190 




(75) 


(81) 


(180) 


II 


76-84 


76-87 


178-203 




(81) 


(84) 


(191)" 



Tarsus 


Claw 


Total 


1 17-152 


17-18 


692-850 


(136) 


(17-5) 


(795) 


127-14N 


17-19 


749-864 


(140) 


(18) 


(814) 


141-160 


19-20 


834-943 


(152) 


(19 5) 


(895) 



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, 6 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 I V-VII. Pleural setae consisting of dorso- 
pleural setae : 4-12 (average 8-i), 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 0-1 (average 0-5), 0-3 (average i-6), 1-3 
(average 2-4), 2-4 (average 2-4) and 0-3 (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 4f— 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. 

Genital segment. Penial sheath of medium length, about \ total body length (ratio 1 : 4-8- 
5-1, average 4-9), 274-308 (average 296) y. long and 30-34 (average 32) y. wide ; lateral sclero- 
tizations narrowly joined anterior to anus ; length of basal rod about i£ times that of aedeagus 
and extending anteriorly from base of aedeagus for about f-f 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 \i 
long), penial sheath and basisternum longer, ratios being 1 : 4-5-5-3 (average 4-8) and 1 : 2-12- 
2-80 (average 2-39) respectively. 

Material examined : 9 specimens, 4 from Laurus nobilis L., collected inLeningrad, 
USSR, on 11.vi.60 and 5 from " citrus ", collected in Leningrad on 3. xi.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) \i ; width at mesothorax 3440-3560 (average 3515) y. Wing expanse 
3440-3560 (average 3515) fz. 

Head subconical in dorsal view ; in lateral view dorsoventrally elongated, with anterodorsal 
bulge pronounced ; length from apex to pronotal ridge 289-308 (average 301) u., width across 
genae 353-385 (average 366) y.. 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 46) hair- 
like genal setae. Eyes : two pairs, subequal ; corneae of dorsal eyes 27-30 (average 28) jx in 
diameter and 1-5-2-3 (average 1-9) times as much apart ; those of the ventral eyes 27-30 
(average 29) u. in diameter and 1 -o-i -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. 
Postocular ridge well developed throughout ; below ocellus the ridge splits up with the anterior 
branch partly surrounding ocellus. Inlerocular ridge absent. Dorsal ocular setae : 1-4 
(average 2-3) f.s. and 1-4 (average 2-3) h.s. on each side, i'enlral 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 54) 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 10-segmented, filiform ; 979-1094 (average 1028) (x 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 
070) and longer than penial sheath (ratio 1 : 2-67-3-13, average 2-94). Scape 57-61 (average 
60) (jl long and 40-45 (average 42) jx wide, with 3 h.s. Pedicel with distinct, polygonal, dorsal 
reticulation ; 65-72 (average 70) \x long and 40-45 (average 52) [x wide ; with 9-13 (average 12) 
f.s. 3-6 (average 4-7) h.s. and a sensillum placodeum. Segment 1 1 1 somewhat club-shaped, 2*6 
3-1 (average 2-8) times longer than wide (68-84, average 78 jx long and 27-30, average 28 [x 




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



9 8 



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 \i) 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 \l, 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. ; antenna! bristles on segments VIII-IX distinctly 
larger than f.s. Segment X : terminal \ constricted ; 72-95 (average 81) y. long and 24-27 
(average 25) ;jl wide (near base) ; carrying 4-8 (average 6-i) f.s., 3 capitate snbapical 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. 

Thorax 730-779 (average 755) \x long. 

Prothorax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral 
pronotal sclerites medium-sized, without setae. Medial pronotal setae occasionally present, 
consisting of 0-2 (average o-6) f.s. and 0-2 (average o-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 structures typical of family. Sternum with strong 
transverse ridge, a strong but interrupted median ridge, and an oval-shaped sclerite. Ante- 
prostemal 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. 

Mesothorax. Mesoprephragma with deep emargination. Prescutum about if times as wide 
as long (average 219 and 121 \i respectively) ; anterior margin strongly curved ; laterally 
bounded by the prescutal ridges and posteriorly by the prescutal suture ; slightly more heavily 
sclerotized medially ; with dense irregular reticulation. Scutum. Median membranous area 
subrectangular ; 152-169 (average 157) pi long and 1-40-1-53 (average 1-45) times as wide 
(width 213-243, average 227 y.) ; with 7-18 (average 11) f.s. and 14-22 (average 18) h.s. 
Scutellum 65-72 jx long arid 220-247 (average 229) \l wide, the ratio being 1 : 3-1-3-8 (average 
3-4) ; tubular, with small ventral foramen ; setae absent. Postnotum with anterior margin 
weakly sclerotized, and usually regular 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. Subalare 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 dark sclerotization. Epimeron small. Eater opleurite not bounded anteriorly by an 
extension from marginal ridge. Basisternum large, about 268 fi. wide and 228 [i long, i.e. 
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 




Fig. 34. Genus B, lateral view. 



OF THE FAMILY COO 1 D \K 99 

on or near median ridge. I'urca well developed. Mesothoracic spiracle with well developed 
peritreme ; postmesospiracular setae numerous, consisting of 50-87 (average 66) f.s., arranged in 
a broad hand behind the spiracles and presternum, often with a few on the episternae. Tegitla 
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. Antemetaspiracular setae : 2-4 (average 3-1) f.s. 

Metathorax. 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 (0-3) f.s. near the lateral h.s 
Pleural ridge considerably reduced, extending only a short distance above coxa] articulation. 
Vestigial precoxal ridge present. Episternum reduced to a small subtriangular plate ; epimeron 
produced posteriorly. Mctathoracic spiracle similar to mesothoracic one. Dorsospiracuiar 
setae : 9-19 (average 13) f.s. and in one case 2 lis Pistmetaspiracular setae : 14-23 (average 
18) f.s. and usually one (0-2) 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 lis ; posterior metasternal setae : 18-41 
(average 27) f.s. 

Wings hyaline ; of medium length (1540-1600, average 1571 jjl) , but comparatively broad 
(720-780, average 763 (j. wide), the ratio width to length being 1 : 203-214 (average 2 06) ; 
alar lobe and alar setae absent. Halteres absent. 

Legs long and slender, with fore pair shortest anil hind pair longest ; ratio length of hind 
leg to body length is 1 : 151 103 (average 1*56). Length of segments (in y.) : 

Leg Coxa Trochanter Femur Tibia Tarsus Claw Total 

I 65-80 76-89 }°4-334 498-536 194-217 25-30 1 180-1286 

(74) (84) (317) (5i6) (203) (28) (1221) 

II 99-122 106-118 277-300 490-543 [86 209 28-30 1203-1301 

(in) (in) (289) (517) (197) ( 2 9) (1253) 

III 101-114 1 14-125 277-304 532-597 200-213 27-31 1277-1382 

(in) (117) (290) (560) (206) (30) (1314) 

Coxae : 16-24 (average 19) f.s. on the fore and 26-34 (>n tne middle and hind coxa, and each 
with 6-12 h.s. ; fore coxa with 2-4 (average 2-6) capitate coxa] bristles ; apical seta about J 
as long as trochanter. Trochanters 29-34 (^ wide ; with 6 oval sensilla ; with 8-10 (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 |j. 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 10-18 on the middle and 
hind femur. Tibiae 23-27 [i wide ; ratio width to length of hind tibia being 1 : 206-22 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 (* 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 ; ungual 
digitules subequal, about as long as claw. 

Abdomen 620-700 (average 676) \x long and 370-500 (average 405) \x wide. 

Segments I-VII : tergites present on anterior margin of segments II— III, represented by 3 
small sclerites — one mediallv and one on each side ; sternites represented by a transverse plate 
on segments II, 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 1st and occasionally one or two on each of segments II— V ; usually one 
h.s. on each side on all segments. Pleural setae : consisting of dorsopleural setae : 5-16 (average 



ioo 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 1-5 (average 
2-6) f.s. on II-VI, and 0-1 (average 0-3), 1-3 (average 1-9), 2-4 (average 3-1), 2-4 (average 3-1) 
and 2-5 (average 3-2) h.s. on segments III-VI respectively, and of ventropleuval setae : occa- 
sionally one or two f.s. on IV, 0-3 (average 15) on V and 0-5 (average 2-4) on the VI, with 
occasionally one h.s. on III and usually one on each of segments IV-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 2-2^ times as long as section within pouch. 
Small IXth tergite present. Ante-anal setae : usually 2 strong h.s., with occasionally one or two 
additional small h.s. and with 1-5 (average 25) 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 1 : 5-6-6-1, average 
5-9), 334-367 (average 352) \x long and 34-42 (average 38) (j. wide ; lateral sclerotizations 
narrowly joined anterior to anus ; length of basal rod i|— 2 times that of aedeagus, extending 
anteriorly from the base of the aedeagus for about J 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 y. long), penial sheath 
and basisternum longer, the ratios being 1 : 4-8-6-1 (average 5-3) and 1 : 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. 

PUL VINARIA 

Pulvinaria fbetulae (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 1st wing vein. When mounted, total body length 1 800-2010 
(average 1885) \x ; width at mesothorax 415-460 (average 434) jj.. Wing expanse 3070-3320 
(average 3170) u.. 

Head subconical in dorsal view ; in lateral view dorsoventrally elongated, with anterodorsal 
bulge pronounced ; length from apex to pronotal ridge 247-288 (average 269) jjl, width across 
genae 247-266 (average 257) [i. 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) \i in diameter and 1-1-1-8 (average 1-5) times as much apart ; 
those of the ventral eyes 30-34 (average 32) \i in diameter and o-6-i-o (average 0-7) times as 
much apart. Ocellus small. Ocular sclerite well sclerotized dorsally, but weak ventrally ; 
polygonally reticulated throughout. Preocular ridge extending only a very short distance 
below articular process. Postocular ridge well developed throughout ; below ocellus the ridge 
splits up, with the anterior branch partly surrounding the ocellus. Interocular ridge absent. 



OF THK FAMILY COCCIDAE 101 

Dorsal ocular setae : 0-5 (average 2-5) f.s. and 0-2 (average 06) h.s. on each 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. 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 10-segmented, filiform ; 840-901 (average 871) u. 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 
090) and longer than penial sheath (ratio 1 : 2-30-2-35, average 2-33). Scape 46-53 (average 
47) u. long and 38-46 (average 43) \x wide, with 3 h.s. Pedicel : dorsal reticulation very weak 
or absent ; 51-61 (average 56) u. long and 42-49 (average 45) [i wide ; with 7-1 1 (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 \i long and 27-30, average 29 a. 
wide), with 5-15 (average 9-3) f.s. of medium length, 1-3-1-5 (average 14) times longer than 
width of segment ; with 2 or 3 usual sensilla basiconica. Segments IV-IX cylindrical ; lengths 
of these segments (in \i) 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 (jl, with distal segments slightly wider than proximal ones ; with -'3-44 (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) u. long and 
25-29 (average 26) [i wide ; caryring 6-1 1 (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 




Fig. 35. Pulvinaria Pbetulae (L.), dorsal and ventral view. 



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. 

Prothorax. 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, not striated, with 4-1 3 (average 7 -8) fleshy post-tergital setae occurring on and behind 
the sclerite on each side. Pleural structures typical of family. Sternum with strong transverse 
ridge, irregular median ridge and a small triangular sclerite. Anteprosternal setae : 0-6 
(average 2-9) f.s. on each side ; prostemal setae : 10-54 (average 37) f.s. and 0-2 (average 1) 
h.s., scattered over the sternal area and spreading into the area anterior to the spiracles. 

Mesothorax. Mesoprephragma with shallow emargination. Prescutum about if times as 
wide as long (average 208 and 119 jx respectively) ; anterior margin strongly curved ; laterally 
bounded by the prescutal ridges and posteriorly by the prescutal suture ; slightly more 
heavily sclerotized medially, reticulation weak or absent. Scutum. Median membranous 
area subrectangular ; 118-125 (average 123) \x long and 1-52-1-67 (average 1-59) times as 
wide (width 179-209, average 196 ;x) ; with 15-28 (average 20) h.s. but no f.s. Scutellum 
65-68 (average 67) [x long and 179-209 (average 196) [x wide, the ratio being 1 : 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 postalare well developed, the latter densely reticulated distally and 
occasionally with a fleshy postalary seta. Mesopostphragma with deep emargination. Meso- 
pleuron. 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. Subalare small. Episternum 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 (x 
wide and 212 jx 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. Mesothoracic spiracle with well developed peritreme ; postmesospiracular 
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 
sclerite with a small, pointed, ventral projection at its base. Additional sclerite small, well 
sclerotized. Antemetaspiracular setae : 3-7 (average 4-8) f.s. 

Metathorax. Metanotum with thickening of posterior margin usually desclerotized medially ; 
suspensorial sclerites 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 
ridge considerably reduced, extending only a short distance above coxal articulation. Epi- 
sternum reduced to a small subtriangular plate ; epimeron produced posteriorly. Metathoracic 
spiracle similar to mesothoracic one. Dorsospiracular setae : 10-19 (average 15) f.s. Post- 
metaspiracular setae : 19-32 (average 25) f.s. and 0-2 (average 1) h.s. Metasternal plate weak 



\ N . 




^^%^#^!^i 




Fig. 36. Pulvinaria Pbetulae (L.), lateral view; also referable for lateral view of P. acericola 

(Walsh & Riley). 



OF THE FAMILY COCCIDAK 103 

and irregular, but somewhat more heavily sclerotized near anterior margin. Anterior meta- 
sternal setae : 50-79 (average 68) f.s. and 0-4 (average i-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 \l), but comparatively broad 
(width 580-700, average 637 ji), 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 \l) : 

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) 

II 72-89 01-99 182-213 357-4 10 U3-I4 8 26-28 870-977 

(83) (96) (198) (383) (139) (27) (926) 

III 76-91 91-112 198-217 37°-437 137-160 27-30 018-1034 

(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 1 
with 6-1 1 h.s. ; fore coxa with 2-4 (average 2-7) capitate coxal bristles ; apical seta about £ as 
long as trochanter. Trochanters 26-31 u. 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 21) times as long as width of 
trochanter. Femora of medium width (39-49 u.), ratio width to length of hind femur being 

1 : 4-5-4-8 (average 4-6) ; with 28-48 f.s. and 7-14 h.s. Tibiae 21-27 I* wide, ratio width to 
length of hind tibia being 1 : 16-5-17-7 (average 170) ; each with 70-115 setae of which 
16-31 are h.s. and 54-89 f.s., the latter about ij to 2 times longer than width of tibia ; a] 
spur about the same size on all tibiae. Tarsi 21-23 H- wide, hind tarsus 6-7 (average 6-5) times 
longer than wide ; each with 23-28 f.s. and 10-18 h.s. ; tarsal digilitles 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, with small a] 
knob. 

Abdomen 570-600 (average 594) u. long and 350-450 (average 410) a wide. 

Segments I —V 1 1 : lergiles 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 ; sterniles 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 : 0-2 (average 0-7), 0-1 (average o-6), 0-1 (average 03), 0-3 (average 
0-9), 0-1 (average 03) and 0-2 (average 0-3) f.s. on segments I— VI respectively ; one h.s. on 
each side on I and each of segments IV— VII, and very rarely a single seta on II 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 0-4 (average 21) f.s. on I-YI and 0-1 (average 01), 
0-3 (average 12), 2-3 (average 26), 2-5 (average 34) and 2-6 (average 3-5) h.s. on segments 
II-VI respectively, and of ventropleural setae : 0-2 (average o-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 0-2 (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 2J-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 29) h.s. and an occasional f.s. on each side. 



io 4 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

Genital segment. Penial sheath of medium length, about \ total body length (ratio i : 4-9- 
5-3, average 5-1), 365-388 (average 375) |x long and 40-46 (average 44) jx wide ; lateral sclero- 
tizations narrowly joined anterior to anus ; length of basal rod about equal to 1 \ times that of 
aedeagus, extending anteriorly from the base of the aedeagus for \ to f 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 |x 
long), penial sheath and basisternum longer, the ratios being 1 : 4-9-5-3 (average 5) and 
1 : 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 Ribes 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 1760-2 160 (average 1963 \i). 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-i) 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) h.s. ; 
antennae 809-999 (average 887) [x long, length in relation to body length 1 : 2-08-2-37 (average 
2-23). 

Thorax. Medial pronotal setae absent ; post-tergital setae 0-8 (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) |x long and 205-239 (average 223) |x wide (ratio 1 : 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) f.s. and 0-4 (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) (x long and 610-670 (average 642) ;x 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) [x long, length in 
relation to body length 1 : 5-15-5-63 (average 5-41) ; aedeagus 72-82 (average 77) jx 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) [i. ; width at mesothorax 420-480 (average 458) ;x. Wing expanse 2840-3140 (average 

2978) [X. 

Head subcorneal in dorsal view ; in lateral view obliquely elongated, with anterodorsal bulge 
pronounced ; length from apex to pronotal ridge 228-293 (average 260) [x, width across genae 
251-266 (average 257) (x. 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 COCCIDAE 



105 



Midcranial ridge dorsally weak and irregular ; ventrally narrow, but well denned, 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) u. in diameter and 1 -7-2-4 (average 1-9) times as much 
apart ; those of the ventral eyes 27-34 (average 30) y. in diameter and 0-9-1-4 (average 11) 
times as much apart. Ocellus small. Ocular sclerite well sclerotized dorsally, but weak ven- 
trally ; polygonally reticulated throughout. Preocular ridges extending only a very short 
distance below articular process. Postocular ridge well developed throughout ; below ocellus 
the ridge splits up with the anterior branch partly surrounding the ocellus. Interocular ridge 
absent. Dorsal ocular setae : 0-6 (average 23) f.s. and 0-2 (average 05) h.s. on each side. 
Ventral head setae : 70-90 (average 80) f.s. and 4-7 (average 48) h.s., scattered over the ocular 
sclerite, always with some (4-7, average 48) f.s. occurring in front of this sclerite and with 
10-15 (average 12) f.s. between and behind the eyes. Preoral ridge present. Tendon-like 
apodeme long. Cranial apophysis long ; apex deeply bifurcate, extending to around the anterior 
margin of the ventral eyes. Mouth opening irregular. Anterior tentorial pits absent. 

Antennae 10-segmented, filiform ; 733-828 (average 794) (i long, i.e. shorter than half body 
length (ratio 1 : 2-08-2-45, average 2 24), shorter than posterior leg (ratio 1 : 0-78-0-81, 
average o-8o) and longer than penial sheath (ratio 1 : 2-14-2-35, average 2-20). Scape 46-53 
(average 49) (jl long and 34-46 (average 41) y wide, with 3 h.s. Pedicel with distinct, polygonal, 
dorsal reticulation ; 53-65 (average 61) u. long and 38-46 (average 43) y. wide ; with 10-15 
(average 12) f.s., 3-5 (average 4) h.s. and a sensillum placodeum. Segment III somewhat 
barrel-shaped, 2-1-2-3 (average 22) times longer than wide (57-72, average 64 \i long and 
23-29, average 26 y 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 IV-IX 

i' 




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



106 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

cylindrical ; lengths of these segments (in jj.) 1 10-133 (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 V-, 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) pi long and 22-29 (average 23) jx wide ; carrying 6-13 (average 10) f.s., 3 capitate 
subapical setae and 5 antennal bristles of which the 3 long ones are about f as long as the segment 
and the 2 shorter ones about \ as long as the f.s. ; with two sensilla basiconica ventrally, one 
near apex and the other more proximal. 

Thorax 635-741 (average 687) pi long. 

Prothorax. 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 0-7, (average 3-9) fleshv 
post-tergital setae occurring on or behind the sclerite on each side. Pleural structures typical of 
family. Sternum 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. 

Mesothorax. Mesoprepkragma with shallow emargination. Presculum about if times as 
wide as long (average 218 y. and 1 19 pt 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 area subrectangular ; 99-139 (average 124) u. long and 153- 
2-04 (average 1-76) times as wide (width 201-228, average 212 pi) ; with 6-20 (average 14) 
h.s. and rarely 2 f.s. Scutellum 65-72 (average 69) pi long and 213-247 (average 231) pi 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 ; poslnotal apophysis and postalare 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. Subalare small. Episternum 
usually showing weak polygonal reticulation dorsally ; subepisternal ridge well developed, but 
below membranous cleft indistinct and only marked by a band of dark sclerotization. 
Epimeron small. Later opleurite not bounded anteriorly by an extension from marginal ridge. 
Basisternum large, about 287 pi wide and 214 pi 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 spiracle with well 
developed peritreme ; postmesospiracular 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, well sclerotized. Antemetaspiracular setae : 1-7 (average 2-9) f.s. 

Metathorax. 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 ; epimeron produced posteriorly. Metathoracic spiracle 
similar to mesothoracic one. D or sospiracular setae : 10-23 (average 16) f.s. Postmetaspiracular 
setae : 12-25 (average 16) f.s. and 0-2 (average o-8) h.s. Metasternal plate weak and irregular, 
but more heavily sclerotized anteriorly. Anterior and posterior metasternal setae consisting of 
58-94 (average 77) and 34-49 (average 38) f.s. respectively. 



1 . 1 IS11S 


Claw 


Total 


"8-133 


[9 25 


864-998 


[26 


(24) 


(928) 


"8-13.5 


23-25 


Noo-991 


(125) 


(24) 


(94" 


"5-144 


25-30 


942-1056 


(135) 


(27) 


(995) 



OB THE FAMILY COCCIDAE 107 

Wings : hyaline ; of medium length (1230-1350, average 1292 jx) but comparatively broad 
width (660-690, average 670 u.), ratio width to length being 1 : 1 86-1 -07 (average 1 93) ; alar 
lobe and alar setae absent. Halteres absent. 

Legs moderately long and slender, with the fore pan usually shortest and the hind pan- 
longest ; the ratio length of hind leg to body length is 1 : 1-76-1 -85 (average 1-79). Length 
of segments (in y.) : 
Leg Coxa Trochanter Femur Tibia 

1 61-72 87-99 205-243 369-422 

(65) (95) ("7) (391) 

II 80-91 99-114 194-220 572-418 

(86) (108) (208) (391) 
III 84-91 106-116 193-228 S'i<>~456 

(87) (112) (211) (422) 

Coxae : 1 1 24 (average 18), 18-29 (average 23) and 11 30 (average 26) f.s. on the fore, middle 
and hind coxa respectively, and each with 7 n lis ; fore coxa with 3 5 (average 4) capitate 
coxa! bristles ; apical seta about \ as long .is trochanter. Trochanters 30-38 \l wide ; with 6 
Oval sensilla ; with 11-16 (average 13) f.s. on the fore and 10 22 on the middle ami 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 i-6— i-8 (average 
1-7) times as long as the width of trochanter Femora "t medium width (41 53 u.), the ratio 
width to length of hind femur being 1 : 3-9-4-6 (average 42) ; each with 28 -47 f.s. and 9 t6 
h.s. Tibiae 24-30 jx wide, the ratio width to length of hind tibia being 1 : 13.5 l6-6 (average 
140) ; each with 90-118 setae of which 14-22 are h.s. and 72-100 f.s., the latter about i\ 2 
times longer than width of tibia ; apical spur about the same size on all tibiae. Tarsi 21 27 <i 
wide, hind tarsus 4-3-5-2 (average 5) times longer than wide ; each with zz-},z 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 .is claw 

Abdomen 450-570 (average 538) (j. long and 400 450 (average 427) \i wide. 

Segments I-VII : tergiles on segmenst 1 1 — I II represented by 3 small sclerites on .interior 
margin — one medially and one on each side, on segment IV by a small sclcnte on each side, and 
on VII by a weak transverse plate ; sternites represented 1>\ a weak transverse plate on segments 
II, III and VII, and sometimes by a small weak sclerite on each side on IV 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 I V-VI I but none on II ami 111 
Pleural setae consisting of dorsopleural setae : 0-8 (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 0-2 (average 0-3), 0-2 (average 1), 1-5 (average 2-8), 0-3 (average 24) 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 I V-VI Segment VI I with 924 (average 15) f.s. and 4-6 (average 
48) 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 1 1— VII respectively ; usually no h.s. on II, one on each side on III-1V 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- 
52, average 5), 342-360 (average 357) u. 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 rod equal to about ij 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 u. 
long), penial sheath and basisternum longer, the ratios being 1 : 3-6-4-8 (average 4-3) and 
1 : 2-26-2-95 (average 2-59) respectively. 

Material examined : 9 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. Boratyriski. 

P. acericola can readily be separated from P. Pbetulae 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 (Bouche) 

(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 
1880-21 10 (average 1980) ;x ; width at mesothorax 390-500 (average 446) [i. Wing expanse 
3030-3400 (average 3283) u.. 

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) (x, 
width across genae 258-289 (average 275) y.. Median crest sclerotized and distinctly poly- 
gonally reticulated ; with 23-37 (average 28) fleshy and 16-19 (average 17) hair-like dorsal head 
setae. Midcranial 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 
9-1) hair-like genal setae. Eyes : two pairs, subequal ; corneae of dorsal eyes 34-42 (average 
38) ji. in diameter and 1 -2-1 -7 (average 1 -5) times as much apart ; those of the ventral eyes 
34-42 (average 38) y. in diameter and 0-5-0-8 (average 0-7) times as much apart. Ocellus small. 
Ocular sclerite well sclerotized and polygonally reticulated throughout. Preocular ridge extend- 
ing only a short distance below articular process. Postocular ridge usually well developed 
throughout, usually with a weak anterior branch below ocellus. Interocular ridge absent. 
Dorsal ocular setae : 1-7 (average 4-1) f.s. and 0-4 (average 2) h.s. on each side. Ventral 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 0-2 (average 0-7) h.s. between and behind the eyes. Preoral 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 10-segmented, filiform ; 825-998 (average 895) jx long, i.e. shorter than half body 
length (ratio 1 : 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 : i*89-2-io, average 1-95). Scape 49-61 (average 
55) \l long and 42-53 (average 48) \i wide, with 3 h.s. Pedicel with distinct, polygonal, dorsal 
reticulation ; 61-68 (average 64) \i long and 42-49 (average 46) u. wide ; with 7-1 1 (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 u. long and 27-34, average 28 jx 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 u.) 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 (x ; 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 J 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) \x long. 

Prothorax. 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-tergiles comparatively large, with irregular, wavy striatums ; post- 
tergital setae absent. Pleural structures typical of the family. Sternum with strong transverse 
ridge, interrupted median ridge and small triangular sclerite. Anteprosternal setae : 1-3 
(average 2-1) f.s. on each side ; prostemal setae 19-30 (average 26) f.s., scattered over the sternal 
area and spreading into the area anterior to the spiracles. 

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




Fig. 38. Parthenolecanium corni (Bouche), dorsal and ventral view. 



no MORPHOLOGY AND TAXONOMY OF ADULT MALES 

sclerotized medially ; polygonally reticulated. Scutum. Median membranous area sub- 
rectangular ; 118-133 (average 128) \l long and 1-53-1 -82 (average i-6i) times as wide (width 
182-236, average 206 \x) ; with 11-26 (average 20) h.s., but no f.s. Scutellum 65-87 (average 
74) \x long and 198-247 (average 220) y. wide, ratio being 1 : 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 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. Subulare 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. Later opleurite not bounded anteriorly by an extension from marginal ridge. 
Basisternum : large, about 270 [z wide and 214 y. 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. Mesothoracic spiracle with well 
developed peritreme ; postmesospiracular setae numerous, consisting of 71-98 (average 89) f.s. 
and 0-2 (average 0-9) h.s., arranged in a broad band behind the spiracles and presternum. 
Tegula small, membranous bulge with a small weak sclerite posteriorly and with 6-8 (average 
7-2) h.s. Third axillary wing sclerite with a small ventral projection at its base. Additional 
sclerite small, well sclerotized. Antemetaspiracular setae : 4-12 (average 6-5) f.s. 

Metathorax. 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 : 0-2 (average o-8) f.s. 
and 0-2 (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 spiracle similar to 
mesothoracic one. Dorsospiracular setae : 5-14 (average 10) f.s. Postmetaspiracular setae : 
21-35 (average 30) f.s. and 0-2 (average 0-9) h.s. Metastemal sclerite represented by a weak 
transverse plate. Anterior and posterior metastemal setae : 76-94 (average 86) and 31-60 
(average 42) f.s. respectively. 

Wings hyaline ; short (length 1330-1510, average 1439 u.) but comparatively broad (width 
620-740, average 686 \x), the ratio width to length being 1 : 2-04-2-15 (average 2-10) ; alar 
lobe and alar setae absent. H alter es 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 i-8i). Length of segments (in jx) : 

Tibia Tarsus Claw Total 

395-452 125-141 24-29 984-1091 

(429) (132) (26) (1042) 

418-471 I37-H3 27-30 1003-1113 

(444) (140) (28) (1062) 

441-479 i37-!52 27-32 1051-1146 

(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 
|— § as long as trochanter. Trochanters 29-34 V- wide ; with 6 oval sensilla, 12-21 f.s. and 8-1 1 
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. Femora of medium width (42-53 (x), ratio width to length of hind femur being 
1 : 4-8-5-7 (average 5-2) ; with 36-54 f.s. and 15-27 h.s. Tibiae 23-30 jx 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 



-eg 


Coxa 


Trochanter 


Femur 


I 


72-80 


99-118 


247-285 




(76) 


(108) 


(271) 


II 


87-95 


106-118 


224-262 




(92) 


(112) 


(244) 


II 


95-107 


106-122 


232-266 




(102) 


("7) 


(251) 



el THE FAMILY COCCI D A F. m 

same size on all tibiae larsi 21-27 I* wide, with 13-28 f.s. and 16-26 h.s. ; tarsal digitules 
subequal, somewhat longer than claw. Claws of medium length, a little longer than width of 
tarsus ; slightlv curved, with small denticle near tip ; ungual digitules subequal, about as long 
as claw . 

Abdomen 520-600 (average 570) jx long and 390-500 (average 425) ij. wide. 

Segments I-Vll : tergites represented by a small sclerite on each side on anterior margin of 
segments 1 1 — 1 1 1 and a weak transverse plate on VI I ; 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 — 1 1 ; 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), 1-4 (average 22), 2-6 (average 3*9), 1-6 (average 3-1) and 0-3 
(average 1-4) f.s., and 0-2 (average 0-9), 0-1 (average 01), o— I (average 03), 1-4 (average i-8), 
0-5 (average 3-3), and 2-5 (average 30) lis. on segments I VI respectively, and of rem 'ro pleural 
setae : 0-3 (average i-i) f.s. on VI and usually one h.s, on each of segments I V VI. Segment 
VII with 10-17 (average 15) f.s. and 512 (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), 5-1 1 (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 I V and 4 on each of \ VII 

Segment VIII with weak termite and transverse sternite . caudal extension forming a small, 
sclerotized, papilla-shaped lobe with .1 small, membranous cicatrix laterally ; glandular pouch 
with 2 long, pointed setae, whose protruding pari is i\ \ tunes as long as section within pouch. 
Small IXth tergite present. Ante-anal setae : 1 strung lis, 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"3^)> 456-486 (average 470) u. long and 45 40 (average 46) u wide at base of aedeagus ; lateral 
sclerotizations not joined anterior to anus ; Length ol basal rod about j{ or equal 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 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 11 long), penial sheath and basisternum 
longer, the ratios being 1 : 3 9-4 3 (average 41) and 1 : r;i) 207 (average 1 09) re- 
spectively. 

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

In addition, ten specimens were examined from Belgrade, Yugoslavia (received 
from N. Mitic-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 \x) : 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 
Variation in P. corni from three localities 



Characters 




Localities 






Warsaw 


Belgrade 


Alma-Ata 


Total length 


1880-21 10 


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) 


(74°) 




Antennal length in relation to body 








length 


1 : 2 06-2 4 1 


1 : 2-12-2-33 


1 : 2 04 




(2-23) 


(2-24) 




Thorax 








Prosternal setae : fleshy 


22-39 


18-40 


13-28 




(3i) 


(28) 


(20) 


Median membranous area of scutum: 








length : 


118-133 


84-118 


103 




(128) 


(101) 




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) 


(7o) 


Scutellum: width 


198-247 


144-205 


182-205 




(220) 


(184) 


(194) 


Basisternum: length 


194-236 


156-201 


213 




(214) 


(182) 





OF THE FAMILY COCCIDAE 
Characters Localities 



Basisternum : width 

Postmesospiracular setae (fleshy) 

Tegular setae (hair-like) 

Antemetaspiracular setae (fleshy) 

Dorsospiracular setae (fleshy) 

I'ostmetaspiracular setae (fleshy) 

Anterior metastemaJ setae: fleshy 

Posterior metasternal setae: flesh] 

Wing length 

Wing width 

Ratio width to length 

Hind leg: length 

length in relation to body length 

Tibial setae 

Abdomen 

Pleural setae on segment VII: flesh} 

Ventral setae on segment II : fleshy 

I'enial sheath: length 

length in relation to body length 

Aedeagus: length 



>3 



Warsaw 


Belgrade 


Alma-Ata 


-233-304 


194-262 


266-277 


(270) 


(237) 


(272) 


71-98 


44-85 


73-8o 


(89) 


(63) 


(77) 


6-8 


4-9 


7-9 


(7-2) 


(57) 


(8-o) 


4-12 


2-5 


5 


(6-5) 


(3-i) 




5-14 


5-15 


7 


(10) 


(10) 




2 1-35 


L5-34 


18 


(3o) 


(19) 




76-94 


40-82 


76 


(86) 


(57) 




31-60 


18-45 


39-50 


(42) 


(29) 


(45) 


i33°-i5i° 


1 100-1380 


1310-1400 


(1439) 


( 1 260) 


(1355) 


620-740 


480-650 


650-710 


(686) 


(598) 


(680) 


1 : 2 04 215 


1 : 1 -10-2-31 


1 : 1 -97-2-02 


(2-IO) 


(2-18) " 


(2 00) 


1051-1146 


832-1098 


1 064- 1 139 


(nog) 


(973) 


(1 102) 


1 : [ 74-1*84 


1 : 172-177 


1 : 1 62 


(i*8i) 


(175) 




i°7-*53 


81-123 


101-127 


10-17 


11-25 


11-14 


(15) 


(20) 


(12) 


30-38 


13-42 


26-31 


(34) 


(25) 


(29) 


456-486 


361-410 


410-429 


(47o) 


(394) 


(420) 


1 : 4-19-4-44 


1 : 3 9I-4 39 


■ : 3-84-4-5I 


(4-38) 


(4- 2 3) 


(4-28) 


106-125 


01-114 


114 


("4)' 


(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) \x ; width at mesothorax 405-450 (average 418) \i. Wing expanse 
2590-2910 (average 2750) \l. 



II 4 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



Head subcorneal in dorsal view ; in lateral view obliquely elongated dorsoventrally, with 
anterodorsal bulge pronounced ; length from apex to pronotal ridge 243-277 (average 261) \i, 
width across genae 247-281 (average 265) jx. 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. Midcranial 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) (x in diameter and 2-7-3-3 (average 3) times as much apart ; those of the ventral eyes 23-30 
(average 26) y. in diameter and 1-1-1-5 (average 1-3) times as much apart. Ocellus small. 
Ocular sclerite well sclerotized and polygonally reticulated throughout. Preocular ridge extend- 
ing only a short distance below articular process. Postocular ridge well developed dorsally and 
lateroventrally, but usually weak posteromedially ; below ocellus the ridge usually has a weak 
anterior branch. Interocular ridge absent. Dorsal ocular setae : 2-8 (average 4-2) f.s. and 
0-4 (average i-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 0-1 (average o-6) h.s. between and 
behind the eyes. Preoral 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. 



;g®P^^s£^ 




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



OF THE FAMILY COCCIDAE 115 

Antennae 10-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 1 : 0-72-0-74, average 
073) and longer than penial sheath (ratio 1 : 1-92-2-09, average 2-01). Scape 49-57 (average 
53) |x long and 40-44 (average 42) |x wide, with 3 h.s. Pedicel with distinct, polygonal, dorsal 
reticulation ; 49-57 (average 53) u. long and 42-46 (average 43) y. wide ; with 3-8 (average 
5-1) f.s., 5-9 (average 69) h.s. and a sensillum placodeum. Segment III : club-shaped, 2-0-2-9 
(average 23) times longer than wide (53-63, average 59 ;x long and 21-27, average 26 (x wide) ; 
with 8-15 (average 13) f.s. of medium length, 1 0-1-4 (average 1-2) times longer than width of 
segment ; with 1—4 usual sensilla basiconica. Segments IV— IX : cylindrical ; lengths of these 
segments (in (x) 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 (x ; 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 iq) f.s. respectively, but no h.s. ; antennal 
bristles on segments VIII-IX distinctly larger than the fleshy setae. Segment X : terminal 
.', 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 1 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) ;x long. 

Prothorax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral 
pronotal sclerites small, with 0-3 (average i-i) fleshy and 0-3 (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 striatums ; posl-tergilal 
setae absent. Pleural structures typical of family. Sternum 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. 

Mesothorax. Mesoprephragma with shallow emargination. Prescutum less than twice as 
wide as long (average 115 and 201 |x respectively) ; anterior margin strongly curved ; laterally- 
bounded by the prescutal ridges and posteriorly by the prescutal suture ; slightly more heavily 
sclerotized medially ; polygonally reticulated. Scutum. Median membranous area subrectan- 
gular ; 103-118 (average m) jx long and 1-50-1-78 (average 1 -6o) times as wide (width 175- 
182, average 178 jx) ; with 14-22 (average 19) h.s., but no f.s. Scutellum 53-61 (average 58) u. 
long and 175-182 (average 178) |x wide, ratio being 1 : 2-9-3-8 (average 3-3) ; tubular, with a 
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 ; postnatal 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 




Fig. 40. Parthenolecanium pomeranicum (Kaw.), lateral view; also referable for lateral 

view of P. corni (Bouchd) . 



n6 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. Subalare 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. Latero- 
pleurite not bounded anteriorly by an extension from marginal ridge. Basisternum large, about 
240 [x. wide and 179 u. long, i.e. 1-48-1-74 (average i-6i) 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 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 
sclerite with a small ventral projection at its base. Additional sclerite small, well sclerotized. 
Antemetaspiracular setae : 2-8 (average 5-3) f.s. 

Metathorax. Metanotum with posterior margin desclerotized medially ; suspensorial sclerites 
absent. Postnotum consisting of a small transverse sclerite on each side. Metatergal setae : 
0-3 (average o-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 precoxal ridge 
usually present. Episternum reduced to a small plate ; epimeron produced posteriorly. 
Metathoracic spiracle similar to mesothoracic one. Dorsospiraculav setae : 3-1 1 (average 7-8) 
f.s. and 0-3 (average 1) h.s. Postmetaspiracular setae : 14-24 (average 18) f.s. and 1-4 (average 
2-2) h.s. Metasternal sclerite represented by a weak, transverse plate. Anterior and posterior 
metasternal setae : 56-74 (average 64) and 20-32 (average 28) f.s. respectively. 

Wings hyaline; short (length 1 130-1290, average 1210 jx) but comparatively broad (width 
520-590, average 557 (j.), ratio width to length being 1 : 2-15-2-20 (average 2 17) ; alar 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 : 1-69-1-84 (average 1-77). Length of segments (in ^) : 



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) 


II 


76-87 


84-91 


201-224 


353-38o 


I33-I4 8 


25-27 


882-948 




(81) 


(87) 


(209) 


(366) 


(140) 


(26) 


(972) 


III 


76-95 


80-99 


209-228 


372-410 


141-163 


26-29 


934-1017 




(87) 


(92) 


(216) 


(39i) 


(L5o) 


(27) 


(963) 



Coxae : 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. Trochanters 28-32 [i 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. 
Femora of medium width (42-49 pi), ratio width to length of hind femur being 1 : 4-5-4-9 
(average 4-6) ; each with 26-42 f.s. and 20—37 ns - Tibiae 23-30 y. 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 arc 
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 a. wide, hind tarsus 6-2-8-6 (average 7) times longer than wide ; 
each with 8-19 f.s. and 27-37 ns - '• 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) [x long and 410-530 (average 460) y. wide. 

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



01 THE FAMILY COCCIDAK 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 
21), 0-4 (average 15), 0-5 (average 17), 1-4 (average 26), 0-3 (average 2) and 0-4 (average 
14) f.s., and 0-1 (average 0-2), 0-2 (average 0-9), 0-3 (average 15), 1-4 (average 28), 1-3 
(average 2-1) and 0-2 (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 
11), 9-14 (average 11), 5-1 1 (average 73), 4—1 1 (average 71) and 1-4 (average 2-7) f.s. on 
segments II— VII respectively ; usually one h.s. on each side on segments [II— TV 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 1 long, 
pointed setae, whose protruding part is about twice as long as section within poach. Small I Xth 
tergite sometimes present. Ante-anal setae : 2 strong h.s., which arc sometimes forked. 
Posterior margin with 2-3 (average 2-8) h.s. on each side 

Genital segment. Penial sheath of medium-length, about \ total body length (ratio 1 : 4 40- 
5-09, average 477), 342-361 (average 353) u. long and 36-42 (average 38) u. wide ; lateral 
sclerotizations apparently not joined anterior to anus ; basal rod about J-§ 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), pem.il sheath and basisternum 
longer, the ratios being 1 : 3-8-4-5 (average 4-2) and 1 : 1-84-2-25 (average 210) 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. corm (Bouche). The two species can be 
separated by the number of fleshy pleural setae on the caudal extension of segment 
VII. In P. pomeraniciim these setae vary from 4-8 (average 5) and in P. corni they 
vary from 11-14 (average 12), 10-17 (average 15) and 11-25 (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 /\ 
pomeraniciim, whereas only one medial pronotal seta was present in one specimen 
of P. corni (from Warsaw). The ventral foramen of the scutellum is large in P. 
pomeraniciim, its length being half or more than half that of the scutellum, while in 
P. corni 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. pomeraniciim, but strong in P. corni. 



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) u. ; width at mesothorax 360-400 (average 372) u.. Wing expanse 2170- 
2310 (average 2277) o.. 



n8 



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) jjl, width 
across genae 201-224 (average 213) y.. 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-i) hair-like ones. Midcranial 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) \i in diameter and 1-8-2-7 (average 2-2) times as much apart ; 
those of the ventral eyes 25-27 (average 26) \i in diameter and o-6-i-o (average 07) times as 
much apart. Ocellus small. Ocular sclerite well sclerotized and polygonally reticulated 
throughout. Preocular ridge extending only a very 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. Interocular ridge absent. 
Dorsal ocular setae : 0-4 (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 44) 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 




/adeh 


400p 


BCFGIO 


ISO|i 


SB 


lOOu 


JKL 


60(j 



Fig. 41. Ceroplastes berliniae 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 10-segmented, filiform ; 541-673 (average 622) \i long, i.e. shorter than half body 
length (ratio 1 : 2-39-2-54, average 2-46), shorter than posterior leg (ratio 1 : 0-68-0-74, 
average 071) and longer than penial sheath (ratio 1 : 1-43-1-64, average 1-55). Scape 38-42 
(average 40) (x long and 34-40 (average 37) jx wide, with 3 h.s. Pedicel with distinct, polygonal, 
dorsal reticulation ; 48-53 (average 50) u. long and 27-30 (average 29) u. wide ; with 6-8 
(average 71) 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 jj. wide) ; with 4-8 (average 6-2) f.s. of medium length, 1-0-1-4 times longer than 
width of segment ; with 1-4 usual sensilla basiconica. Segments IV-IX cylindrical ; length of 
these segments (in \i) 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 [i, 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) \x long and 20-23 (average 21) \l 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) ji. long. 

Prothorax. 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. Sternum 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 43) 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. 

Mesothorax. Mesoprephragma with moderately deep emargination. Prescutum more than 
twice as wide as long (average 197 and 85 [i 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) [i long and 
1-69-1-94 (average 1 -8o) times as wide (width 175-199, average 188 jz) ; with 10-20 (average 
14) f.s. and 4-14 (average 10) h.s. Scutellum 53-65 (average 60) [i long and 190-209 (average 
200) (i. 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 




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



120 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

(average i -7) fleshy postalary setae occurring on or immediately posterior to it. Mesopost- 
phragma with deep emargination. Mesopleuron. Mesoplenral 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 dark 
sclerotization. Epimeron small. Later opleurite partly bounded anteriorly by an extension 
from marginal ridge. Basisternum large, about 223 ;x wide and 171 [i 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 spiracle with well developed peritreme ; postmesospiracular 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 
sclerite with a pronounced ventral projection at its base. Additional sclerite small. Antemeta- 
spiracular setae : 1-5 (average 2-7) 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 : 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 precoxal ridge 
present. Episternum reduced to a small subtriangular plate ; epimeron produced posteriorly. 
Metathoracic spiracle similar to mesothoracic one. Dorsospiracular setae : 3-9 (average 5-5) 
f.s. Postmetaspiracular setae : 12-16 (average 14) f.s. and 0-2 (average o-6) h.s. Metasternal 
plate weak and irregular, but more heavily sclerotized anteriorly. Anterior and posterior 
metasternal setae : 36—48 (average 41) and 18-29 (average 23) f.s. respectively. 

Wings hyaline; short (length 920-980, average 968 \l) and comparatively broad (width 450- 
480, average 465 [±), the ratio width to length being 1 : 2-00-2-18 (average 2-08) ; alar lobe and 
alar 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 \i) : 

Tibia Tarsus Claw Total 

289-325 106-122 21-23 7 2 4-798 

(310) (113) (22) (770) 

304-338 1 12-125 20-23 744- 8l 3 

(319) (119) (22) (780) 

338-365 122-133 22-24 825-876 

(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 (* 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 1 -52) times as long as the width of trochanter. Femora of 
medium width (37-45 u.), with ratio width to length of hind femur 1 : 4-1-4-4 (average 4-2) ; 
with 24-35 f.s. and 8-16 h.s. Tibiae 19-27 u. wide ; ratio width to length of hind tibia 1 : 12-8- 
15-9 (average 14-3) ; each with 81-111 setae of which 21-30 are h.s. and 58-83 f.s., the latter 
about 1 \ times longer than width of tibia ; apical spur about the same size on all tibiae. Tarsi 
17-21 \x wide, hind tarsus 6-4-6-6 (average 6-5) times longer than wide ; each with 17-27 f.s. 
and 6-1 1 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 ; ungual 
digitules subequal, about as long as claw. 

Abdomen 380-450 (average 409) (x long and 330-380 (average 345) (x 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 ; sternites represented by a 



Leg 


Coxa 


Trochanter 


Femur 


I 


49-61 


68-76 


186-205 




(54) 


(73) 


(197) 


II 


65-74 


80-89 


158-175 




(7o) 


(84) 


(166) 


III 


76-84 


89-95 


175-186 




(80) 


(92) 


(180) 



OF THE FAMILY COCCIDAE 121 

weak transverse plate on segments II, III, VI 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 — I II, and usually one on each side on each of 
segments [V— VII. Pleural setae consisting of dorsopleural setae : 0-3 (average 0-7), 0-3 
(average 1-3), 0-3 (average 1-7), 1-3 (average 2), 0-3 (average 1 -6) and 0-4 (average 1) f.s. on 
I-VI, and 0-2 (average 0-9), 2, 0-2 (average 1 -6) and 12 (average 17) h.s. on segments III-VI 
respectively, and of ventropleural setae : occasional lv one (range 0-2) f.s. on II, usually 2 
(range 0-3) on III, and 3 (range 1-5) on each of segments IV V I, and usually one h.s. on each of 
segments [V— 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 [I— 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 IXth tergite present. 
Ante-anal setae : 0-3 (average 1 -a) f.s Posterior margin with 2-6 (average 3-8) f.s. and 2-3 
(average 29) h.s. on each side. 

Genital segment. Penial sheath long, about *-\ total body length (ratio 1 : 3-6 41, average 
3' 8 ). 395-4IO (average 42) |j. long and 38-42 (average 40) [j. 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 jj 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 (1 18-137, average 125 u. long), 
penial sheath and basisternum longer, the ratios being 1 : 3-0-3-4 (average 31) and 1 : 1 -28 
1 52 (average 1 30) 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 1). 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) \x ; width at mesothorax 370-410 (average 386) \i. Wing expanse 2320-2500 
(average 2417) u.. 

Head subcorneal in dorsal view ; in lateral view dorsoventrally elongated, with anterodorsal 
bulge not pronounced ; length from apex to pronotal ridge 205-247 (average 225) \i, width 
across genae 213-251 (average 231) y.. Median crest sclerotized, showing distinct, polygonal 
reticulation which does not enclose irregular striations ; with numerous (16-27, average 22) 
fleshy dorsal head setae and 7-1 1 (average 81) 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) \i in 
diameter and 1-6-2-6 (average 21) times as much apart ; those of the ventral eyes 29-32 
(average 30) u. in diameter and o-6-i-o (average o-8) times as much apart. Ocellus small. 
Ocular sclerite well sclerotized, polygonally reticulated throughout. Preoculav ridge extending 



122 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

a very short distance below articular process. Postocular ridge well developed throughout ; 
ventral to ocellus the ridge usually splits up, with the anterior branch partly surrounding 
ocellus. Interocular 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. Preoral 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 10-segmented, filiform ; 678-770 (average 727) \x long, i.e. shorter than half body 
length (ratio 1 : 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) \l long and 38-42 (average 39) \x wide, with 3 h.s. Pedicel occasionally with poly- 
gonal reticulation dorsally, but usually with only wavy striations ; 46-53 (average 50) ^ long 
and 34-47 (average 38) \x wide ; with 8-17 (average 12) f.s., 4-6 (average 5-4) h.s. and a sen- 
sillum placodeum. Segment III somewhat club-shaped, 2-6-3-2 (average 3) times longer than 
wide (59-72, average 69 u. long and 21-25, average 23 \i 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 a.) 106-137 (average 120), 91-105 
(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 \x, with distal segments slightly wider than 




Fig. 43. Ceroplastes sp., dorsal and ventral view. 



OF THE FAMILY COCCIDAK 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 £ constricted ; 
65-84 (average 78) \x long and 19-23 (average 21) (i wide (near base) ; carrying 2— II (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) u. long. 

Prothorax. Pronotal ridge strong, but medially interrupted by weak sclerotization. Lateral 
pronotal scleriles 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. Sternum 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. 

Mesothorax. Mesoprephragma with moderately deep emargination. Prescutum about twice 
as wide as long (average 196 and 96 |jl 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) \i long and 1-75-2-14 
(average 1 -99) times as wide (width 160-209, average 180 jj.) , with 2-1 1 (average 67) f.s. and 9-12 
(average 11) h.s. Scutellum 46-65 (average 55) u. long and 175 198 (average 188) pt 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 
postalare well developed, the latter densely reticulated distally and with 1-6 (average 33) 
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. Later opleurite partly bounded anteriorly by an extension 
from marginal ridge. Basisternum large, about 232 \x wide and 158 (x 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 spiracle with well developed peritreme ; postmesospiracular 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 : z-j (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 o-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. 
Episternum reduced to a small subtriangular plate ; epimeron produced posteriorly. Meta- 
thoracic spiracle similar to mesothoracic one. Dorsospiracular setae : 9-18 (average 15) f.s. 
Postmetaspiracular 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 ^^) f.s. respectively. 

Wings hyaline ; short and comparatively broad : 970-1060 (average 1018) a. long and 445-500 
(average 483) u. wide, the ratio width to length being 1 : 2-04-2-18 (average 211) ; alar lobe and 
alar setae absent. Halleres absent. 



i2 4 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 i : 1-54-1-69 (average i-6i). Length of segments (in |x) : 



Leg 


Coxa 


Trochanter 


Femur 


Tibia 


Tarsus 


Claw 


Total 


I 


57-°5 


70-78 


220-247 


323-376 


124-131 


24-27 


827-928 




(62) 


(76) 


(23O 


(347) 


(129) 


(25) 


(870) 


11 


68-78 


84-95 


194-209 


346-380 


125-137 


25-28 


849-927 




(75) 


(89) 


(201) 


(363) 


(132) 


(27) 


(885) 


III 


80-87 


91-101 


201-220 


365-418 


133-137 


25-27 


903-989 




(83) 


(98) 


(213) 


(394) 


(136) 


(26) 


(95o) 



Coxae : with 14-16 (average 15), 22-30 (average 26) and 30-41 (average 37) f.s. and 7-9 
(average 8), 7-1 1 (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. Trochanters 23-30 [i wide ; with 6 oval sensilla, 10-17 IS - an d 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. Femora of medium width (38-46 \l), ratio width to length of hind femur being 
1 : 4-6-5-3 (average 4-9) ; each with 31-44 f.s. and 12-23 h.s. Tibia 19-25 jx 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 i£ times longer than width of tibia ; apical spur about the 
same size on all tibiae. Tarsi 19-22 y. wide, hind tarsus 6-o-6-6 (average 6-4) times longer than 
wide ; with 20-32 f.s. and 8-16 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 ; ungual digitules subequal, about as long as claw. 

Abdomen 330-460 (average 399) y. long and 300-360 (average 333) \± 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 VI-VII ; sternites represented by a weak 
transverse plate on segments II, III and VII. Caudal extension of segment VII large, tapering, 
weakly sclerotized. Dorsal setae : 5-14 (average 9) f.s. on I, 1-5 on II and III, and up to 3 on each 
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), 1-5 
(average 2-9), 1-6 (average 3-2), 0-6 (average 3-2), 2-6 (average 4-1) and 1-4 (average 2-2) f.s. 
on I- VI, and 0-1 (average 03), 1-2 (average i-6), 1-2 (average 1-7), 1-3 (average 2-1) and 0-2 
(average 1 -6) h.s. on segments II-VI respectively, and of ventropleural setae : up to 5 f.s. on 
each of segments I II-VI, and usually one (range 0-2) 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 11), 5-12 (average 9) and 6-11 (average 9) f.s. on segments I I-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— 3^ times as long as section within pouch. Small IXth tergite present. 
Ante-anal setae : 2-5 (average 3-8) f.s., 0-4 (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 f total body length (ratio 1 : 3-5-3-8, average 
3-6), 407-445 (average 426) fj. long and 38-44 (average 40) jx wide ; lateral sclerotizations 
narrowly joined anterior to anus ; length of basal rod about twice that of aedeagus, extending 
anteriorly from base of the aedeagus for about j| 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 jx long), penial 
sheath and basisternum longer, ratios being 1 : 4-1-4-9 (average 4-5) and 1 : 1-53-1-78 (average 
1 -66) respectively. 



OV THE FAMILY COCCIDAE 125 

Material examined : 10 specimens, collected by J. Munting on Milletia sp. in 
Umkomaas, South Africa, on 7.H.62 ; no adult females present with the males ; 
nearbv females of Ceroplastes Pmimosae 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 \ II. 

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 know ledge 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., Toumeyclla 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 Eriopeltis, 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, Saisselia, Pul- 
vinaria and Paralecanopsis), Filippiinae (including FUippia 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. corni 
from different localities (Poland, Russia and Yugoslavia) showed large differences, 
e.g. those from Yugoslavia were about f 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 COCCIDAK 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 E. Pfestitcae 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 postocidar 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 ERIOPELTIS group and (iii) is 
reduced posteromedially in some species but not in others. A broad interocular 
ridge is characteristic of the ERIOPELTIS 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- 
caninm, 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 Sulc (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 INGLISIA 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 INGLISIA 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 1st, (iii) the length of the 3rd segment in relation to its width, 
(iv) the relative lengths of the 3rd and 10th 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 INGLISIA 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 prescutum and the size 
of the membranous area of the scutum can be used as supplementary characters to 
separate genera and species. A tubular scutellum is characteristic of the ERIO- 
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 (q.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 ERIOPELTIS group to 
short and broad in the COCCUS group ; in the 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 legs 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 EULECANIUM 
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 EULECANIUM 
group, in the INGLISIA 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 EULECANIUM 
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, (iii) 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 pleural 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 lu:ulae 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 INGLISIA 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, (iii) 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. 



130 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 



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OF THE FAMILY COCCI DAI. 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 i 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 «>t analysis used by 
Ghauri (10.62) 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 whk h separate major Groups of Genera 

Characters Eule- Erie- Ingli- Coccus 

canium peltis sia 

I [bad: 

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. B. Snbequal 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, vcntrally: 

a. Reduced, b. Not so. b ab a b 

6. Number of simple eyes: a. 4. 

b. More than 4. ab a b a 



i 4 o 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 - a* - 

8. Pre-oral ridge: a. Absent, b. Present. ab a b b 

9. Fleshy dorsal head setae: a. Present. 

b. Absent. b b a a 

10. 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. 

13. Genal setae: a. Present, b. Absent. 

Thorax: 

14. Prosternum: a. Spinose. b. Not so. 

15. Scutellum: a. Tubular, b. Not so. 

16. Scutellum: a. 3-4 times as wide as 
long. b. 1-3 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, b. Reduced or absent. a ab b a 

20. Fleshy postmesospiracular setae: 
a. Present, b. Absent. b b a a 

21. Antemetaspiracular setae : 
a. Present, b. 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* 



B 


B 


A 


A 


B 


B 


A 


A 


B 


B 


A* 


B 


AB 


A 


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 i\ times longer 
than wide. b. 2j— 2} times longer than 
wide. c. More than 2} times longer than 

wide. ab c* B A 



26. Total number of setae on front tibia : 
a. More than 60. b. Less than 60. 



ab 



27. Hind tibia: a. With more fleshy than 
hair-like setae, b. With more hair-like 
setae. 



Abdomen: 

28. Pleural sclerotization : a. Present on 
segm. VII. b. Present on segm. IV— VII. 
c. Absent. 

29. Caudal extensions of segm. VI I : 

a. Very prominent, tapering, b. Small, 
broadly rounded or somewhat pointed. 

30. Cicatrix on caudal extension of 
segm. VIII. a. Absent, b. Present. 

31. Fleshy pleural setae on segm. I— III: 
a. Present, b. Absent. 

32. Fleshy ventral setae: a. Absent 
posteriorly beyond segm. III. b. Present 
posteriorly beyond segm. III. 

33. Fleshy setae on 8th sternite: a. Mure 
than 6. b. Not more than 3. 

34. Ventral hair-like setae on segm. Ill, 
usually: a. Two setae medially, b. No 
setae medially. 



Total number of exclusive characters 



'4 2 



MORPHOLOGY AND TAXONOMY OF ADULT MALES 

Table 2 A 



Pairs of Groups 



Number of Characters 
(a) (b) (c) 

shared of which exclusive differentiating 



Eulecanium-Eriopeltis 


14 


6 


6 


Eulecanium-Inglisia 


8 


I 


15 


Eulecanium-Coccus 


6 


1 


16 


Eriopeltis-Inglisia 


9 


1 


21 


Eriopeltis-Coccus 


9 


2 


17 


Inglisia-Coccus 


16 


6 


16 



i. 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- 
caniunt and Ceroplastes. Pairs of genera share from 75 to 90 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 (viii) 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 aurantii (Ckll.) by Borchsenius (1957) and Neo- 
pulvinaria imeretina Hadz. by Hadzibejli (1955), and the drawing of the posterior 
part of the abdomen of Saissetia nigra (Nietner) by Green (1904-1909) it seems 
certain that the genera Akermes, Chloropulvinaria, N eopulvinaria and Saissetia 
should also be included in this group. 



OF THE FAM [LY COCCIDAE i i? 

2. The INGLISIA group comprises only the genus Inglisia, 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 INGLISIA 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 IYth-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 presternum and on the front coxae, (iii) 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 VHIth. 

3. The ERIOPELTIS group : This group consists of the two closely related 
genera Eriopeltis and Luzulaspis, which share about 90 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 EULECAX I I'M 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, (iii) 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 (vii) 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 Vlllth. In 
addition, in Eriopeltis the ventromedial part of the epicranium is well sclerotized, 
and tergites and sternites are present on all the abdominal segments. 



i 4 4 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, Ericerus, 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, (hi) 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, Nemolecanium, 
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, (ii) a basalare connecting the pleural wing process 
with the episternum, (iii) 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 VHIth 
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, Nemolecanium, 
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 90). The three remaining genera, i.e. Ericerns, Genus A and Sphaerole- 
canium are not very close to each other or to any of the other above-mentioned 
subgroups (although Ericems 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 2 A 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 INGLISIA 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 INGLISIA 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 ERIOPELTJS and EULECANIUM 

groups 

1. Fleshy dorsal head setae present. Fleshy dorsal bead 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. Vntemetaspiracular 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 Filippiinae, Borchsenius included the 
three genera Luzulaspis, Eriopeltis and Filippia. From the male characters it is 
clear that whereas Luzulaspis and Eriopeltis are very closely related (sharing about 
90 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 Eriopeltis 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 90 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- 



()!■• 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- Krio- Coccus 

anium peltis 

Head: 

1 . Shape of the head. g <. 



2. Absence or presence of polygonal reticulation 
mi median crest. 

3. Presence of interocular ridg 

4 Nature of postocular ridge, dorsally. 

5. Nature of postocular ridge, posteromedial^ 
(>. Midcranial ridge dorsally represented or not. 

7. Nature of scleroti/.ation and reticulation 
around ventral part of midcranial ridge. 

8. Nature of preocular ridge. 

9. Number of simple eyes. 

10. Distance between dorsal eyes in relation to 
diameter of cornea. 

1 1. Distance between ventral eves in relation to 
diameter of cornea. 

12. Presence of preoral ridge. 



14. Length of cranial apophysis. 

15. Nature of apex of cranial apophysis. 

16. Presence of polygonal reticulation on genae. 

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



13. Presence of ventral sclerite near mouth opening. g 



gs 



s 
GS 



18. Location of ventral head setae. 



i4« morphology and taxonomy of adult males 

Table 3. List of Characters which mainly Separate Genera 

Characters Eulec- Erio- Coccus 

anium peltis 

Antenna: 

19. Length in relation to body length. g - g 

20. Length in relation to length of posterior leg. G 

zi. Length in relation to length of penial sheath. g g 

22. Length of 3rd segment in relation to its width. g g 

23. Relative widths of 1st 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 10th 

segment. g — 

27. 10th segment distally constricted or not. — g 

28. Length of two smallest antennal bristles in 

relation to length of fleshy setae. u - gs 

Thorax: 

29. Number of circular pores behind pronotal 

sclerite. g g 

30. Prosternum reticulated or not. — — G 

31. Number of medial pronotal setae. g — gs 

32. Presence of post-tergital setae. - — g 

33. Presence of flesh v prosternal setae. g — 

34. Nature of emargination of mesoprephragma. g s g 

35. Nature of reticulation on prescutum. g g gs 

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 



of the family coccidak 149 

Table 3. List of Characters which mainly Separate Genera 

Characters Eulec- Erin- Coccus 

anium peltis 

Thorax could. 

40. Presence of fleshv scuta] set ic G 

41. Number of hair-like scutal setae. (; - 

42. Presence of postalary sel 11 g 

43. Number of fleshy postmesospiracular setae. 

44. Presence and type >>l basisternal setae. G 

45. Presence of suspensorial sclerite. G - — 
\t> Nature of metapleural ridge. f". - 
47. Number oi fleshy anterior metasternal setae G 

|S Length of wing in relation to its width g 

49. Presence of alar setae. G — — 

50. Presence of halteres. G - 

51. Number of haltere setae. G - 

52. Length of hind leg in relation to body length. G s 

53. Presence on coxa] bristles on front coxa g S 

54. Total number of setae on front tibia. G 

55. Length of apical seta on front trochanter in 

relation to width of trochanter. g g g 

56. Shape of hind femur (length/width). <. G 

57. Relative number of fleshy and hair-like setae 

on hind tibia. 1 : - 

58. Relative length of fleshy setae on tibiae. g 

50. Relative numbers of fleshy and hair-like 

setae on hind tarsus. G G 

60. Length of hind tarsus in relation to its width. g gs 

61. Relative length of hind claw. g g 



150 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. III. 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 

Ahdoman — conic/. : 

83. Length of basal rod in relation to length of 

aedeagus. G g os 

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 s 

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 Liizulaspis are very 
closely related but Liizulaspis is distinct in lacking the glandular pouch and pores. 

EULECANIUM group. The genera Eulecanium, Nemolecanutm, 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. Ericerus 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 Sphaerolccanium 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. 

2. Presence of lateral arms of mid- 

cranial ridge. X 

3. Shape of apex of cranial apophysis. x 

4. Presence of reticulation on gena. X 

5. Presence of reticulation on pedicel. 

6. No. of fleshy setae between and 
behind ventral eyes. 

7. No. of fleshy setae on 4th antennal 

segment. X 

8. Size of two shortest antennal bristles. x 

9. Presence of reticulation on the 
post-tergital region. X 

10. Nature of emargination of 
mesoprephragma. X 

11. Presence of reticulation on 

mesoprescutum. X - 

12. Presence of reticulation on anterior 

arms of mesoscutum. X - 

13. Presence of reticulation on 

membranous area of scutum. — 

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. X 

18. Presence of median pronotal setae. 
i<i. Presence of coxal bristles. 

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

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

12.. 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 midcranial ridge. This short median ridge is present 
in some members of the EU LECANIUM 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. Interocalar ridge. This ridge, which connects the pre- and postocular ridges 
below the ocellus, is present and alwavs well developed in the ERIOPELTIS group. 



i 5 4 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 exceptionallv long in Eriopeltis 
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, 
1 961) 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. pomeranicnm. 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 Pseudococcus (Giliomee, 1961), are present. 

7. Basalare. This structure is vestigial or absent in P. pomeranicnm (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. pomeranicnm. 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. pomeranicnm 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, Nemolecanium, Enlecanium, 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 COCCIDAK 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 I >iaspidoid 

A. Primitive characters: 

1. Head well sclerotized. X X 

2. Tentorium present. X- 

3. Distinct propleural apophyses present. X 

B. Specialized characters: 

1 . Compound eyes absent. X 

2. Lateral branches of midcranial ridge 
present. 

3. Cranial apophysis long. 

4. Pronotal ridges present. 

5. Post-tergites small. 

6. Prescutal ridges shifted medially. 

7. 1'realare 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 (9) 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). 



X 


X 


X 


X 


X 


X 


X 


X 


X 


X 



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 thev 
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 Steingelia 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 



1. Postoccipital ridge completely absent. 

2. Postocular ridge present. 

3. Cranial apophysis extended. 

4. Membranous area of scutum present. 

5. Scutellum short, transverse. 

6. Median ridge of basisternum present. 

7. Disc pores on body (other than 
glandular pouch) absent. X 

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 Phenacoleachia) and diaspidoid types. 



Cocc- 


Stein- 


Phenaco- 


Pseudo- 


idae 


gelia 


leachia 


coccidae 


X 


X 


- 


- 


X 


- 


X 


X 


X 


X 


- 


- 


X 


X 


- 


- 


X 


X 


- 


- 


X 


X 


_ 


_ 



of the family coccidak 
Table 7 



157 



Characters differentiating male Coccidae and Pseudococcidae compared 
with conditions in margaroid and dlaspidoid types 



Margar- Pseudo- Cocc- Diasp- 

oid coccidae idae idoid 



\ Primitive characters: 

1. Anterior tentorial pits/arms separate, x 

2. Membranous area of scutum present 

3. Median ridge of basisternum present 

4. Metasternal sclerite present. 

5. Ostiole absent. X 



X 


X 


X 


- 


X 


X 


X 


X 


X 


X 



15. Specialized characters: 
1 . Postoccipital ridge absent. 
1. Scutellum short, transverse. 

3. Metasternal apophyses absent. 

4. Penial sheath elongated 

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



X 


— 


X 


X 


X 


X 


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 Icerya (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 (i960) 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 querats (L.) by Borchsenius 
(i960), 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 

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 present ......... 2 

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 ......... 3 

Antenna shorter than posterior leg ; midcranial ridge ventrally complete ; with 4 
simple eyes ; presternum 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 J 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— I II present ; fleshy setae on sternite VIII not more than 3, 
if present ........... COCCUS group 

Antenna 1-07-1-15 (average i-io) times longer than posterior leg ; midcranial ridge 
ventrally reduced ; with 8 simple eyes ; presternum 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 

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 2J 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 J times longer than wide ; abdomen without pleural 
sclerotization ; fleshy ventral setae not occurring posteriorly beyond abdominal 
segment IV EULECAN1UM group 



160 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

Keys to Genera and Species 
Key to the EU LECAN I UM Group of Genera 
i 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 if— 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 ............ 2 

2 Head (in lateral view) rounded ; ventral sclerite near mouth opening present ; 

fleshy setae very short and thick, being less than 060 times as long as width of 3rd 
antennal segment and shorter than width of hind tibia ; 10th 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 ; 10th antennal segment 
with 2 or 3 capitate subapical setae ........ 3 

3 Third antennal segment more than 4 times (range 4-6-6-3, average 5-3) longer than 

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 . . . ERICERUS (E. pela (Chav.)) 

- 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 ........ 4 

4 Median crest not polygonally reticulated ; scutellum tubular ; with more than 6 

hair-like scutal setae ........... 5 

Median crest polygonally reticulated ; scutellum not tubular ; with less than 5 

hair-like scutal setae ........... 6 

5 With 8 simple eyes ; with 1-4 (average 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 i-i) 
alar setae on each side ; hind tarsus 5-6-7-0 (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 § to equal that of aedeagus, basal rod not reaching apex of 
basal membranous area; aedeagus short, the basisternum being 1-74-1-92 

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

With 10 simple eyes ; with no dorsal ocular setae ; 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.)) 



OF THE FAMILY COCCIDAK 161 

Halteres and suspensorial sclerites absent, metapleural ridge reduced ; fleshy 
postmetaspiracular setae present ; abdominal segment 111 with no hair-like 
setae medially ; pleural sclerotizations of penial sheath narrowly joined anterior 
to anus PHYLLOSTROMA (P. myrtilli (Kalt )) 

Halteres and suspensorial sclerites present, metapleural ridge complete ; fleshy 
postmetaspiracular setae absent ; abdominal segment 111 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 : 315-413 ........ 8 

Dorsal part of midcranial ridge absent ; with 8 simple eyes ; apex of cranial 
apophysis truncate ; antenna longer than half bodv length, the ratio oi its length 
in relation to body length being 1 : 1-52-1-61 (average 157) ; antenna also 
2-4-2-6 (average 2-5) times longer than penial sheath ; 3rd antenna! 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 : 172- 
1-83 (average 1-78) ; claws long, hind claw about i\ times as long as width of 
tarsus ; abdominal tergites and sternites absent on segments 1 V and V ; abdom- 
inal segment II usually with none, but occasionally with one median hair-like seta 
ventrally . . . RHODOCOCCUS (R. spiraeae (Borchs.)) 

Vestigial dorsal part oi midcranial ridge present ; with 4, 6 or 10 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 2-02 ; 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, tin- 
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 ............. 9 

With 4 simple eyes ; mesoprephragma without an emargination ; prescutum with 
distinct polygonal reticulation ; basisternum 1-7-2-4 (average 21) 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 usuallv with 2 setae on both segments; basal rod 
short, length |-| that of aedeagus PHYSOKERMES (P. piceae (Schr.)) 

With 6 or 10 simple eyes ; mesoprephragma with shallow- emargination ; prescutum 
with weak or no reticulation ; basisternum 2-5-3-6 times longer than mem- 
branous area of scutum ; alar setae present on at 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 ■£-$ 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 
1 : 1-04-1-14 (average i 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 -ft-ths of the length from aedeagus to apex of basal 
membranous area .... NEMOLECANIUM (N. abietis Borchs.) 



iGi 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 i : 0-73-0-87 (average o-8o) ; 3rd antennal segment 1-8-2-2 
(average 2) times longer than wide ; basisternum without setae ; setae of 
glandular pouch with external part about i\ times as long as the section within 
the pouch ; basal rod extending § or f of the length from aedeagus to apex of 
basal membranous area . ... EULECANIUM (£. tiliae (L.)) 



Key to the Genera and Species of the ERIOPELTIS Group 

With a pair of median, hair-like ventral head setae distinctly longer than the rest ; 
3rd antennal segment 30-37 (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 

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 

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, finger-like, membranous extension 
at the apex CEROPLASTES 2 

Head with anterodorsal bulge pronounced ; with more than 43 fleshy postmeso- 
spiracular setae ; usually 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 



OF THE FAMILY COCCIDAK [63 

Vestigial dorsal part of midcranial ridge present on anterior margin of head ; with 
1 1-2 1 (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 6-1 1 (average 9-2) fleshy 
setae ; aedeagus short, basal rod about 2 times longer and penial sheath 
41-4-9 (average 45) times longer ...... Ceroplastes sp. 

Dorsal part of midcranial ridge absent ; with 5-9 (average 71) fleshy setae between 
and behind ventral simple eyes ; 4th antennal segment with 11- iq (average 15) 
fleshy setae ; coxal bristles absent ; abdominal segment VII witli 1 5 (average 3) 
fleshy setae ; aedeagus medium-sized, about as long as basal rod, and penial 
sheath 3-0-3-4 (average 32) times longer . . . . . C. berliniae Hall 

1'ost-tergital setae absent ; hind tarsus usually with more hair-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 1 1 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 respectivelv ...... 5 

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 (Bouche) 

Vestigial dorsal part of midcranial ridge present ; distal part of terminal antennal 
segment not constricted ; mesoprephragma with a shallow emargination ; fleshv 
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 0-7) ; length of basal rod about equal to ij 
times that of aedeagus PULVINARIA 6 

Dorsal part of midcranial ridge absent ; distal part of terminal antennal segment 
somewhat constricted ; mesoprephragma 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 VII 1 with a geniculate or 
cylindrical caudal extension which carries a large cicatrix posteriorly ; fleshy 
ante-anal setae absent ; length of basal rod more than i^ 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 0-6) hair- 
like setae ventrally ....... Pulvinaria ?betulae (L.) 

The two shortest antennal bristles on the terminal 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 9-1 1 
(average 10) times longer than wide ; length of 2 shortest antennal bristles about 
equal that of the fleshy setae ; presternum not reticulated ; with 2-9 (average 
54) fleshy basisternal setae ; abdominal segment VIII with a geniculate caudal 
extension ; setae of glandular pouch with external part about 2-2I- times as long 



164 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

as the section within the pouch ; abdominal segments H 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 4f-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 ..... COCCUS (C. hesperidum L.) 



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166 MORPHOLOGY AND TAXONOMY OF ADULT MALES 

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1958b. Ibid. Part V. The nut or thorn scale Lecanium coryli (L.) sensu Marchal nee 

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OF THE FAMILY COCCIDAE 167 

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

Walsh, B. D. & Riley, C. V. 1868. A new bark louse on the Osage orange. Am. Ent. 1 : 14. 
Weber, H. 1928. Skelett, Muskulatur und Darm der schwarzen Blattlaus Aphis fabae Scop. 

Zoologica, Stnttg. 28 Heft 76, 120 pp. 

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

Wilkey, R. F. & McKenzie, H. L. 1961. Systematic status of the Psendococcus maritimus- 

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Calif. Dep. Agric. 50 : 245-249. 



1 68 

abietis, Nemolecanium, 41 
acericola, Pulvinaria, 104 

berliniae, Ceroplastes, 117 
Pbetulae, Pulvinaria, 100 
bituberculatum, Palaeolecanium, 53 

Ceroplastes, 117 

Ceroplastodes, 92 

chiton, Ceroplastodes, 92 

Coccus, 92 

corni, Parthenolecanium, 108 

Ctenochiton, 63 

Ericerus, 67 
Eriopeltis, 77 
Eulecanium, 37 

Pfestucae, Eriopeltis, 81 
Eilippia, 59 

Genus A, 70 
Genus B, 96 

hesperidum, Coccus, 92 

Inglisia, 88 

luzulae, Luzulaspis, 85 

Luzulaspis, 85 



INDEX 

myrtilli, Phyllostroma, 56 

Nemolecanium, 41 

Palaeolecanium, 53 
Parthenolecanium, 108 
pela, Ericerus, 67 
Phyllostroma, 56 
Physokermes, 45 
piceae, Physokermes, 45 
pomeranicum, Parthenolecanium, 113 
prunastri, Sphaerolecanium, 74 
Pulvinaria, 100 

Rhodococcus, 49 

Sphaerolecanium, 74 
sp., Ceroplastes, 121 
sp., Ctenochiton, 63 
sp., Eriopeltis, 77 
sp., Genus A, 70 
sp., Genus B, 96 
spiraeae, Rhodococcus, 49 

theobromae, Inglisia, 88 
tiliae, Eulecanium, 37 

viburni, Filippia, 59 




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WORLD SPECIES OF CLEORA 

(LEPIDOPTERA : GEOMETRIDAE) 

By D. S. FLETCHER 



CONTENTS 

Synopsis 

Material Examined 

Acknowledgements 

Treatment 

Cleora Curtis . 

Taxonomic History 

Affinities 

Distribution 

Biology 

Species-Groups 

Key to Species 

Description of Species 
Check List of Species 
Species to be moved from Cleora when genera are available; 
Species moved from Cleora to other genera 
References ....... 

Index ........ 



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SYNOPSIS 

The species of the genus Cleora 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 
10 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 Cleora 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 Oberthur and L. B. Prout, material has been loaned by the 
following : 



4 D. S. FLETCHER 

Museum national d'Histoire naturelle, Paris ; Monsieur CI. Herbulot, Paris ; 
Musee Royal de l'Afrique Centrale, Tervuren, Belgium ; Zoologisches Museum der 
Humboldt-Universitat, Berlin ; Staatliches Museum fur 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 CI. 
Herbulot, Dr. H. B. D. Kettlewell, Prof. O. Lindner, Dr. E. C. G. Pinhey and Dr. 
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 Cleora, 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 : pi. 88. Type-species : Geometra cinctavia Denis & Schiffermiiller, 1775, by 

original designation. 
Cerotricha Guenee, 1857 : 284. Type-species : Cerotvicha licornaria Guenee, by monotypy. 

Syn.n. 



REVISION OF AFRICAN SPECIES OF CLEORA 5 

Barsine Meyrick, 1883 : 530 nee 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 nee Walker, 1854. 
Chogada Moore, 1887 : 415. Type-species : Boarmia alienaria Walker, i860, by original 

designation. [First synonymy by Prout, 1928 : 155.] 
Carecomotis Warren, 1896 : 402. Type-species : Carecomotis perfumaria Warren, by mono- 

typy. Syn.n. 
Neocleora 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- 
taria Denis & Schiffermuller, sublunaria Guenee, repetita Butler and alienaria Walker) a second, 
unsealed 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 repetita 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 w-ith 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 R\ and R% free from radius, 
rarely from a point or shortly stalked (munditibia Prout, dodonaeae Prout) ; -7?3-5 on long stalk ; 
A/2 a little closer to M\ than to M3 ; Cii\ from just before lower angle. Hind wing : termen 
slightly crenulate ; Sc + 7?i approximated to basal half of upper margin of cell ; R5 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 M.% 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 albobrnnneata 
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, albobrunneata) ; 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 D. S. FLETCHER 

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 (Guenee) 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 Cleora, 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 Chora 
were treated by Prout in two revisions, 1929a and 1937. The species previously 
included in Carecomotis were revised by Fletcher in 1953. The Ethiopian species 
of Chora then known were treated by Janse in 1932 under the name Neochora. 

Affinities. Chora is closely related to Ascotis Hiibner (1825 : 313), which is 
represented in southern Europe by A. sehnaria sehnaria (Denis & Schiffermuller), 
by sehnaria reciprocaria (Walker) in continental Africa and by further subspecies in 
India, Ceylon, China and Japan. The neuration is almost identical to that of Chora, 
but R% sometimes anastomoses briefly with R3. 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 Chora ; the characteristic shape of the 
signum (with the anterior edge toothed, often strongly) at each corner, separates it 
from the remaining species of Chora. 

Also closely related to Chora is the genus Alois Curtis (1826 : 113) occurring in 
the Palaearctic and Indo-Australian regions. Though the neuration is identical 
with that of Chora, 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 unsealed. 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 Alois so far examined, there are two weakly spiculate signa on the bursa copulatrix. 

Hypopaipis Guenee (1862 : 29) was erected for H. terebraria Guenee and H. 
per for aria Guenee, since shown by Vinson (1938 : 38) to be synonymous ; terebraria 
was placed by Vinson in Chora. Hypopaipis terebraria and H. antemelaria Mabille 
(1893), island endemics on Reunion 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 sehnaria and they have been transferred to the genus Ascotis. 
Hypopaipis terebraria Guenee is here selected for the first time as type-species of 
Hypopaipis, which becomes a junior synonym of Ascotis. Syn. n. 

Prout (1938 : 154) erected the genus Colochora 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 Colochora, Alois ansorgei Warren, show some 
similarity in habitus to that of Chora, but structurally are closely related to Ascotis. 
In the male the neuration is identical with that of Chora, but the pectinations of the 
antennae arise from the base of each segment and are unsealed ; in the females of 
those species of Colochora available for study, veins R\ 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, Ophthahnodes , 
Pseudalcis and Cusiala occur with and approach closely in habitus many species of 
Cleora, but are structurally distinct. 

Paradromulia Warren (1896 : 300) has veins R\ and R2 of the fore wing long- 
stalked in both sexes, R2 anastomosing briefly with R3. 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 Ri and R% 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 unsealed ; 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 Guenee (1857 : 2 &3) nas 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 unsealed. 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 R% in the fore wing are long- 
stalked, Ri anastomoses briefly with Sc and R2 with R$ to form a long, slender areole. 
The male antennae are bipectinate for the greater part of their length ; the long 
pectinations are unsealed 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 R2 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 Cleora 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 
Reunion, legrasi and macracantha on Madagascar, transversaria and angustivalvis 



REVISION OF AFRICAN SPECIES OF CLEORA g 

on the Comoro Is., and three species in the Gulf of Guinea, tamsi on Principe I., and 
viettei and prosema on Sao Thome I. Subcincta subcincta is an endemic subspecies 
on Sao Thome 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, quadrimacidata, 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 Chora ; 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 1. 

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 raphis group, which includes raphis, 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 raphis group on 
the basis of the male genitalia. Quadrimacidata and lacrymata are associated 



io D. S. FLETCHER 

tentatively with the raphis group, principally on the female genitalia ; though the 
males of these two species lack areas of coarse scobination on the valves, quadri- 
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 raphis. 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 
dargei, 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 rothkirchi are 
weakly sclerotized ; those of macracantha are asymmetrical and aberrant. In 
addition to rostella, the species group includes legrasi, angustivalvis, serena, oligo- 
dranes, macracantha, derogaria, pavlitzkiae, lima, scobina, radula and rothkirchi. 

The species toidgoetae and lamottei are tentatively placed in the rostella group on 
the form of the cornuti on the vesica. 

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

The species tamsi and viettei, endemics on the islands of Principe and Sao Thome 
respectively, though closely related to each other, have been tentatively placed in 
the genus ; herbidoti and acerata, another closely related pair, have also been placed 
only tentatively ; subcincta 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. 



Key to Species based on Male Genitalia 

Valve with area of dense, coarse scobination ventrally just basad of cucullus, 

but not extending to dorsal margin ...... 2 

Valve not so developed ........ 6 

(1) Scobinate area in form of a slender, arcuate band from below cucullus reaching 
ventral margin at one-third ; arm of sacculus curved, spatulate, of even width 
and scobinate at apex, extending in length to mid-cucullus ; short, setose 
digitate process medially at one-third valve (Text-fig. 69) . boetschi (p. 54) 

Scobinate area and sacculus not so formed ..... 3 



REVISION OF AFRICAN SPECIES OF CLEORA n 

3 (2) Scobinate area extending to cucullus ; process from sacculus almost rhomboid, 

posterior dorsal angle produced in short, digitate form (Text-fig. 54) 

echinodes (p. 46) 

- Scobinate area separated from cucullus ; process from sacculus aculeate (Text- 

figs. 57, 61, 64) . . . . . . . . 4 

4 (3) Vesica with three cornuti, but without scobinate band (Text-fig. 65) 

panarista (p. 51) 

- Vesica with two cornuti and scobinate band (Text-figs. 58, 60) ... 5 

5 (4) 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. 48) 

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, 61) . . aculeata (p. 49) 

6 (1) Vesica with single cornutus, shallowly bifurcate at apex ; aculeate arm of 

sacculus parallel with and extending to cucullus (Text-figs. 16, 17) 

melanchorda (p. 24) 
Vesica and sacculus not so formed ....... 7 

7 (6) Arm of sacculus three-fourths as long as dorsal margin of valve, apex incurved 

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. 34) 
Sacculus and aedeagus not so formed ...... 8 

8 (7) Arm of sacculus partially or entirely covered by a setose, membranous fold from 

ventral margin of valve ; aedeagus with one or more cornuti . . 9 

Sacculus not so formed . . . . . . . .17 

9 (8) Arm of sacculus parallel with ventral margin of valve, apex incurved and bear- 

ing a cluster of spines . . . . . . . .10 

Sacculus not so formed . . . . . . . .12 

10 (9) Aedeagus smooth ; vesica with one tapered cornutus sub-equal in length to 

mid-width of aedeagus ; arm of sacculus three-fifths as long as dorsal margin 
of valve (Text-figs. 39, 40) ..... epiclithra (p. 36) 

Apical third of aedeagus scobinate along left half of dorsal surface ; vesica with 

two or more cornuti . . . . . . . .11 

11 (10) Arm of sacculus three-fifths as long as dorsal margin of valve, with a short, 

semicircular process at base ; vesica with three weakly sclerotized cornuti 
(Text-figs. 34, 35) ...... prosema (p. 34) 

Arm of sacculus nine-tenths as long as dorsal margin of valve, without basal 
process ; vesica with two cornuti, broad based and strongly sclerotized 
(Text-figs. 31, 32) . . . . . . . oculata (p. 31) 

12 (9) Arm of sacculus with bifurcate, claw-like apex ; aedeagus broadened apicad ; 

vesica with two tapered cornuti and a longitudinal scobinate band (Text-figs. 

22, 23) ........ cancer (p. 28) 

Arm of sacculus with simple, tapered apex ; aedeagus not broadened apicad ; 

vesica with two or three tapered cornuti . . . . .13 

13 (12) Arm of sacculus not extended beyond two-thirds ventral margin of valve, 

tapered and angled through 90 ° at middle (Text-fig. 4) ; endemic on island 

of Reunion ........ acaciaria (p. 16) 

- Sacculus not so formed, species not occurring on island of Reunion . . 14 

14 (13) Arm of sacculus parallel with and extending to four-fifths ventral margin of 

valve (Text-fig. 19) ..... papillifer (p. 25) 

Arm of sacculus arcuate or sinuous, tip extending to dorsal margin of valve . 15 



12 D. S. FLETCHER 

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

usually 30-35 mm. ; endemic on Comoro Is. . . transversaria (p. 18) 

Arm of sacculus finely tapered ; wing-span of moth usually 40-45 mm. ; not 

found on Comoro Is. . . . . . . . .16 

16 (15) Aedeagus six times as long as broad, one and one-half times as long as dorsal 

margin of valve (Text-fig. 14) .... flavivenata (p. 22) 

Aedeagus almost eight times as long as broad, twice as long asdorsal margin of 

valve (Text-fig. 11) . . . . . . betularia (p. 20) 

17 (8) 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) ...... carcassoni (p. 36) 

- Sacculus not so formed . . . . . . . .18 

18 (17) Arm of sacculus slender and spatulate, dorsal margin partly fused with dorsal 

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) 19 

Sacculus not so formed ........ 20 

19 (18) Serrate-edged process on sacculus one and one-half times as long as greatest 

width of valve (Text-fig. 47) ; endemic on Sao Thome I. 

subcincta subcincta (p. 43) 
Serrate-edged process on sacculus twice as long as greatest width of valve 

(Text-fig. 48) ; continental Africa. . . subcincta longifibulata (p. 43) 

20 (18) 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 (Text-fig. 44) . . 21 

Sacculus not so formed ........ 22 

21 (20) Vesica with two cornuti (Text-fig. 45) .... herbuloti (p. 38) 

- Vesica without cornuti ...... acerata (p. 41) 

22 (20) Sacculus with one or more well sclerotized, tapered processes, each as long as, 

or longer than uncus, the apices curving through 90 ° . . . .23 

- Sacculus not so formed ........ 24 

23 (22) Sacculus with two tapered processes, each smooth to tip ; vesica with one 

cornutus equal in length to width of aedeagus (Text-figs. 72, 73) bicornis (p. 57) 

- 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. 53) 

24 (22) Cucullus almost circular and projecting beyond ventral margin of valve ; 

sacculus with a curved, tapered process arising from near mid-valve, with or 
without additional processes ....... 25 

Cucullus and sacculus not so formed ...... 26 

25 (24) Sacculus with a short truncate process at one-third ventral margin of valve ; 

vesica with one long and one short cornutus (Text-figs. 25, 26) ; endemic 

on Principe I. ....... tamsi (p. 28) 

- Sacculus smooth along ventral margin and without additional process ; vesica 

with one tapered cornutus (Text-figs. 28, 29) ; endemic on Sao Thome I. 

viettei (p. 30) 

26 (24) 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 1-13 ; vesica with two cornuti, one scobinate, one 
tapered and smooth (Text-figs. 51, 52) .... lacrymata (p. 44) 
Genitalia not so formed . . . . . . . .27 



REVISION OF AFRICAN SPECIES OF CLEORA 13 

27 (26) 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 usually smooth, but if scobinate 

only lightly so and always smooth in apical eighth .... 35 

28 (27) Longer cornutus coarsely scobinate, scobination extending basad from apex, 

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

Longer cornutus of even width, slightly arcuate and finely scobinate in apical 
two-thirds ; arm of sacculus spatulate and densely but finely scobinate 
(Text-figs. 141, 142) ...... radula (p. 104) 

29 (28) Apex of larger cornutus dilate to one and one-half times width at middle, 

apical perimeter with 8-10 spines ...... 30 

Apex of larger cornutus of even width or tapered . . . .31 

30 (29) Scobinate medial plate of gnathus broad and shallow, equal in width to cucullus; 

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 margin of valve ; larger cornutus 
straight (Text-figs. 87, 88) ..... tulbaghata (p. 67) 

31 (29) Apex of aedeagus produced and narrowed by one-half, irregular in shape ; 

basal half of shorter cornutus bulbous, apical half slender and curved through 

45 (Text-fig. 79) . . . . . . dactylata (p. 60) 

Apex of aedeagus narrowed slightly and evenly rounded. . . .32 

32 (31) Apex of shorter cornutus curved through 45 ° and usually bearing 2-5 spines 

varying in size (Text-fig. 76) ..... dargei (p. 58) 

Shorter cornutus straight and tapered ...... 33 

33 (32) Sacculus with process developed at two-thirds ventral margin of valve. . 34 

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

34 (33) 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) ..... 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) 

35 (27) 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 

3° (35) O ne or both cornuti spined ........ 42 

- Both cornuti quite smooth ........ 37 

37 (36) Tip of uncus produced, tapered and depressed ; 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 ventral 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 

38 (37) Apical process on sacculus sclerotized and greater in length than diameter of 

uncus .......... -39 

- Apical process on sacculus membranous and shorter than diameter of uncus, or 

wanting .......... 40 



14 



D. S. FLETCHER 



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 
Valve not so developed 



legrasi (p. 80) 

41 



UNCUS 



VENTRAL FOLD 



DIGITATE PROCESS 




CUCULLUS 



CRISTAE HAIRS 



ARM OF SACCULUS 



MEDIAL PROCESS 



VENTRAL MARGIN 



AEDEAGUS 




CORNUTI 



Figs. 1-3. Cleora fabricated genitalia. 1, S ; 2, aedeagus ; 3, $ (opposite page). 



REVISION OF AFRICAN SPECIES OF CLEORA 



15 



41 (40) 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. 109) ; endemic on Comoro Is. . . angustivalvis (p. 82) 

42 (36) 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 to seven-eighths, 
apical eighth smoothly tapered ; shorter cornutus smooth or partially 
scobinate .......... 44 



LAMELLA POSTVAGINALIS 



LAMELLA ANTEVAGINALIS 



RIBBING 



STERIGMA 



COLLICULUM 




BURSA COPULATRIX 



16 D. S. FLETCHER 

43 (42) 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 minute, 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) 

44 (42) 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) ....... lima (p. 100) 

- Sacculus not so formed ........ 45 

45 (44) Apical third of sacculus dilate and scobinate, more heavily on inner surface, 

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) .... toulgoetae (p. 76) 

- Sacculus not so formed ........ 46 

46 (45) Process from apex of sacculus slender, spatulate and scobinate, equal 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. 138, 139) . scobina (p. 101) 

- Sacculus not so formed ........ 47 

47 (46) Apical third of spacculus broadly spatulate, apex produced and narrowly 

rounded, minutely and densely scobinate ; gnathus broader at base than 
cucullus, scobination extending posteriorly along subscaphium (Text-fig. 134) 

pavlitzkiae saltuensis (p. 98) 

- Sacculus not so formed ........ 48 

48 (47) Apical projection from sacculus spatulate and slender, apex not incurved 

(Text-fig. 127) ..... pavlitzkiae lamella (p. 94) 

- Sacculus not so formed, apex incurved ...... 49 

49 (48) Apical projection from sacculus blunt, setose but not scobinate and only 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 

50 (49) Apical projection from sacculus one-fourth as broad as cucullus and incurved 

through 90° (Text-fig. 129) . . . pavlitzkiae etesiae (p. 94) 

- Apical projection from sacculus one-third as broad as cucullus and obtusely 

incurved (Text-fig. 131) . . . . pavlitzkiae oriadelpha (p. 98) 

Cleora acacia ria (Boisduval) 

(Text-figs. 4-6 ; PI. 1, figs. 147-151 ; Map 1) 

Boarmia acaciaria Boisduval, 1833 : 264. 

Boarmia acaciaria Boisduval, 1833a : 116, pi. 16 : 4. 

Boarmia acaciaria Boisduval ; Guenee in Maillard, 1862 : 29. 

Cleora acaciaria (Boisduval) Vinson, 1938 : 38. 

Neocleora acaciaria (Boisduval) Viette, 1954 : 5°9- 

Neocleora acaciaria (Boisduval) ; Herbulot, 1957 : 2 34- 

cj. 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 (PI. 1, fig. 147). Under surface of wings 
tilleul buff densely suffused with fuscous (PI. 1, fig. 150). 

o\ 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 a setose, membranous fold from ventral margin of valve ; short setose, digitate process at base 
of arm ; 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). 

$. 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. o* 38-42 mm. ; $ 38-48 mm. 




Figs. 4-6. C. acaciaria genitalia. 4, 6* ; 5, aedeagus ; 6, $. 



18 D. S. FLETCHER 

Ascotis terebraria (Guenee), the only other species known from the island of 
Reunion 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 acaciaria is endemic on the island 
of Reunion. 

The type, which should have passed to Oberthur 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 i). Island of Reunion. 

Material examined. Reunion : (Dr. Roussel), 4 <$, 3 $ ; ibid. (Maillard), 1 $ ; 
ibid., 21-30.V.1922 (G. F. Leigh), 3 c?> 4 ?• 

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

Boarmia transversaria Pagenstecher, 1907 : 97, pi. 6 : 4. 

<J. 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. o* 29-5-35 mm. ; $ 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 PI. 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. 






Figs. 7-9. C. iransversaria genitalia. 7, $ ; 8, aedeagus ; 9, $. 



20 D. S. 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 i). Comoro Islands ; endemic. 

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

Grande Comoro : 26.ix.19n (G. F. Leigh), 1 <$ ; ibid., ix.1921, 2 $ ; ibid., 
x.1921, 2 $ ; ibid., xi.1921, 1 <$ ; ibid., bred, i.viii.1921, 1 <$ ; Grande Comore 
(L. Humblot), 70 <$, 70 $. Mayotte : 1-5.V.1911 (G. F. Leigh), 1 <J ; ibid., 27-31. 
v.1911, 1 $. 

Cleora betularia (Warren) comb. n. 

(Text-figs. 10-12 ; PI. 2, figs. 166-175 ; Map 1) 

Chogada betularia Warren, 1897 : 93. 
Chogada funesta Warren, 1905 : 398. 
Neocleora betularia (Warren) Janse, 1932 : 271, pi. 8 : 7, text-fig. 100. 

cJ. 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. 

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

Measurements. 6* 45-50 mm. {funesta holotype 35 mm.) ; $ 50 mm. 

An exceedingly variable species, displaying a range of polymorphism in the male 
(PI. 2, figs. 166-175) similar to that found in transversaria ; 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 
(PI. 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 (PI. 2, fig. 174) ; in other 
examples these areas are of the ground colour only very sparsely irrorate with fuscous 
(PI. 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 (PL 2, figs. 167, 169, 
171, 173, 175). 

Closely related to transversaria, 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 




Figs. 10-12. C. betnlaria genitalia. 10, (J ; 11, aedeagus ; 12, $. 



22 D. S. FLETCHER 

The holotype of funesta 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 <$ ", but is without abdomen. 

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

Malawi : Zomba Plateau, pupa ex Pinus patula, emerged 19.xii.1962 (R. C. H. 
Sweeny), 1 <$ ; Little Ruo Plateau, Mt. Mlanje, 6400 ft., 3.viii.i956 [A. W. R. 
McCrae), 1 $. Transvaal : Marieps Mtn., 1-9.1.1926 (G. van Son), 10 <$, 4 $, in 
British Museum (Natural History), 5 <$ in Transvaal Museum. Natal : 1 <J ; 
Karkloof, 17.1.1917 (E. E. Piatt), 1 <$ ; ibid., 13.L1918, 1 <$ ; Durban (G. F. Leigh), 
1 <$ ; Balgowan, 5.11.1951 (K. M. Penniton), 1 <$ in Transvaal Museum ; ibid., 
12.ii.1951, 1 (J ; ibid., 9.xii.i95i, 1 <$ ; Mont-aux-Sources, Natal National Park, 
1 $ in Transvaal Museum. Cape Province : Katberg, 4000 ft., x.1933 (R. E. 
Turner), 1 <$ ; Wilderness, Knysna Forest, 20.x-8.xi. 1952, at M.V. light (H. B. D. 
Kettlewell), 1 $ ; Engcobo, 3.L1954 (D. A. Swanepoel), 1 $ in Transvaal Museum. 

Cleora flavivenata sp. n. 

(Text-figs. 13-15 ; PI. 3, figs. 176-179 ; Map 1) 

o*. 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 (PI. 3, fig. 176). Under surface of wings tilleul buff, 
irregularly suffused and patterned with fuscous (PI. 3, fig. 177). 

o*. 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 (PI. 3, fig. 178). Under surface of wings usually less suffused with 
fuscous proximally (PI. 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 : $ 40-45 mm. ; $ 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 tnlbaghata and munda ; from 
these species flavivenata may be clearly separated by the genitalia of both sexes. 

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




Figs. 13-15. C. flavivenata genitalia. 13, <$ ; 14, aedeagus ; 15, $. 



24 D. S. FLETCHER 

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

Paratypes : 13 <$, 5 $ with same data as holotype ; Cape Province, Camps Bay, 
17. xii. 1955 (A. J. Duke), 1 <$, all in Transvaal Museum ; Cape Town, iii.1912 {Lord 
Gladstone), i<$ ; ibid., v.1912, i<^ ; Cape Province, Harkerville, 9.iii.ig2i (Dr. H. G. 
Breijer), 1 <£ ; Knysna, Wilderness, 20.x-8.xi. 1952, at M.V. light (H. B. D. 
Kettlewell), 6 $, all in British Museum (Natural History) ; Knysna, Sourflats, 
22-24.xi.1954 (L. Vari), 1 <$ ; Knysna, Garden of Eden, 16-20.1.1955 (A. J. T. 
Janse), 5 $ ; Groot River Pass, 21-23.L1955 (A. J. T. Janse), 1 $ ; Port St. Johns, 
10-22.ii.1955 (A. J. T. Janse), 3 £, all in Transvaal Museum. 



Cleora melanochorda (Fletcher) comb. n. 
(Text-figs. 16-18 ; PI. 3, figs. 180-182, 184-186 ; Map 1) 

Neocleora melanochorda Fletcher, 1958 : 137, figs. 42, 76, 216. 

o". 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 Sc$ and between vein 
M\ 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 (PI. 3, fig. 184). 

o*. 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. 

$. 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 q* 50 mm. ; Kenya <$ 46 mm. ; $ 48 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 (PI. 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 <$. Uganda : Ruwenzori, Nyinabitaba, 8650 ft., 
7-13. vii. 1952 (D. S. Fletcher) ,Geometridae genitalia slide 1751. 

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



Cleora papillifer Prout 
(Text-figs. 19-21 ; PL 3, figs. 183, 187 ; Map 1) 

Cleora papillifer Prout, 1934 : 89. 

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





17 



Figs. 16-18. C. melanochorda genitalia. 16, q* ; 17, aedeagus ; 18, $. 



26 



D. S. FLETCHER 



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). 

o*. 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. 6* 24 mm. ; $ 34 mm. 




Figs. 19-21. C. papillifer genitalia. 19, 5* ; 20, aedeagus ; 21, ?. 



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 Iridopsis ; 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 $ : [Congo (Leopoldville)] : Kisantu, 1929 
(R. P. J. Van Wing), in Musee Royal de l'Afrique Centrale. 

Paratype <J : [Congo (Leopoldville)] : Lusambo, Kassai, 12.ix.1919, in British 
Museum (Natural History). 




Figs. 22-24. C. cancer genitalia. 22, $ ; 23, aedeagus ; 24, ?. 



28 D. S. FLETCHER 

Cleora cancer sp. n. 

(Text-figs. 22-24 I PI- 3> fig s - 188-191 ; Map 1) 

o*. 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 Ma. 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). 

cj. 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 (PI. 3, figs. 190, 191). 

$. 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. o* 44-47 mm. ; $ 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 <$. Kenya : Ngong, Nairobi, vii.1954 (Fowler & Coulson). 

Paratypes : [Ethiopia] Abyssinia: Harar, 22.xii.1938 (R. E. Ellison), 1 $ ; 
B.E.A. : Kibwezi, 18.V.1917 (W. Feather), 1 $ ; ibid., ii.1929, 1 $. Burundi : 
Usumbura, 900 m., 8. i. 1962 (Dr. M. Fontaine), 1 <$, in Musee Royal de l'Afrique 
Centrale. [Malawi] Nyasa : Zomba, vi.1923 (H. Barlow), 1 $. Rhodesia : 
Salisbury, 25.vi.1918 (O'Neil), 1 $. Transvaal : Louws Creek, near Barberton, 
26 S., 3i°2o" E., xi.1922 (H. G. Williams), 1 J. 

Cleora tamsi sp. n. 

(Text-figs. 25-27 ; PL 4, figs. 192-196 ; Map 9) 

o*. 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 Mi 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 



o*. 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. 

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

$. 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. <J 35-40 mm. ; $ 40-44 mm. 




Figs. 25-27. C. tamsi genitalia. 25, $ ; 26, aedeagus ; 27, $. 



3 o D. S. FLETCHER 

A variable species developing forms comparable with those of acaciaria from 
Reunion and transversaria from the Comoro Is. (PL 4, fig. 196). The male genitalia 
of this and the following species, viettei from Sao Thome 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. Tarns of this 
department, in token acknowledgement of his frequent, kindly advice. 

Distribution (Map 9). Principe I. ; probably endemic. 

HolotypecJ. W. Africa : Principe I., 19.xii.1932 (W. H. T. Tarns). 

Paratypes : St. Principe : 1500-2000 ft., iv-v.1923 (T. A. Barns), 15 <$, 3 $ ; 
Principe I., 19-22.xii.1932 (W. H. T. Tarns), 9 $, 3 $. 



Cleora viettei (Herbulot) comb. n. 
(Text-figs. 28-30 ; PI. 4, figs. 197-202 ; Map 9) 

Neocleora viettei Herbulot, 1958 : 103, fig. 1. 

o* $. Externally similar in both sexes to the preceding species and displaying a similar 
range of colour and wing pattern. 

o*. 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. 6* 37 _ 43 mm. ; $ 39~43 mm. 

Closely related to the preceding species, tamsi, 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). Sao Thome I. ; probably endemic. 

Material examined. Holotype <$. Sao Thome : Bombaim, Traz-os-Montes 
(450 m.), 6-8. vi. 1956, in Museum national d'Histoire naturelle, Paris. 

St. Thome, x-xi.1899 (Mocquerys), 8 $, 10 $ ; Sao Thome, 10-24.1.1926, edge of 
virgin forest (T. A. Barns), 20 q, ii $. 



REVISION OF AFRICAN SPECIES OF CLEORA 



3i 




Figs. 28-30. C. viettei genitalia. 28, <$ ; 29, aedeagus ; 30, $. 



Cleora oculata sp. n. 

(Text-figs. 31-33 ; PI. 4, figs. 203-206 ; Map 2) 

o\ 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). 

o*. 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. 

$. Similar to male externally, but white area of upperside of both wings more densely 
irrorate with cinnamon buff (PL 4, figs. 205, 206). 

$. 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. o* 38-44 mm. ; $ 40 mm. 




Figs. 31-33. C. oculata genitalia. 31, a* ; 32, aedeagus (dorsal view) ; 33, <j>. 



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 £. [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 $ ; Lolodorf, 1894-1895 (L. Conradt), 1 $ ; Epulan [Efulen], 
i.v.1926 (G. Schwab), 1 £. Angola : Quicolungo, 120 km. N. of Lucala, iv.1936, 
800 m. (R. Brann), 1 <$. [Congo (Leopoldville)] : Leopoldville, 27.vii.1954, 1 <$ ; 
Uele, Paulis, 8.^.1956, 2$ ; ibid., 5.VL1956, 1 <$ ; Sankuru, Djeka, 17-18. xii. 1952, 




Figs. 34-36. C. prosema genitalia. 34, <J ; 35, aedeagus (dorsal view) ; 36, $. 



34 D. S. FLETCHER 

2 $ ; Dimbelenge, 24.x. 1950, 1 <$ ; Lusambo, 2g.viii.194g, 1 $ ; ibid., 23.vii- 
28.ix.1g50, 3 (J, 3 ? ; Katako-Kombe, g.xi.ig5i, 1 $, all collected by Dr. M. Fontaine 
and deposited in Musee Royal de l'Afrique Centrale ; W. Kivu, Upper Lowa 
Valley, Nr. Masisi, 5000-6000 ft., forest and long grass, ii.ig24, wet season (T. A. 
Barns), 3 <£. Uganda : Entebbe, 25.viii.1g61, light trap (K. W. Brown), 4^. 

Cleora prosema Prout 

(Text-figs. 34-36 ; PL 4, figs. 207, 208 ; Map 2) 

Cleora prosema Prout, 1927 : 194, pi. 20 : 11. 
Neocleora prosema (Prout) Herbulot, 1958 : 103. 

o*. 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 (PI. 4, fig. 207). 
Underside of wings white, patterned as illustrated with fuscous (PL 4, fig. 208). 

o*. 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. 

$. Similar to male externally. 

$. 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. q* 35-40 mm. ; $ 39 mm. 

Differs externally from the closely related ocnlata 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 Thome I. ; endemic. 

Material examined. Holotype <$. Sao Thome, 10-24. 1.1926, edge of virgin 
forest (T. A. Barns), Geometridae genitalia slide No. 2125. 

Sao Thome : data as type, 2 $, 1 $ ; Sao Thome, 24. i-25.il. 1926 (T. A. Barns), 
2$ ; ibid., ix-x.ig26, i£ ; Sao Thome I., 30.x-24.xi.1g32 (W. H. T. Tarns), 10^. 

Cleora anacantha sp. n. 

(Text-figs. 37-38; PI. 4, figs. 2og-2io; Map 2) 

o*. 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 (PI. 4, fig. 209). Underside of wings tilleul 
buff, weakly suffused with drab (PI. 4, fig. 210). 

o*- 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 





38 





Figs. 37-40. Cleora $ genitalia. 37-38, anacantha. 37, $ ; 38, aedeagus (ventral view) 

39-40 epiclithra. 39, $ ; 40, aedeagus. 



36 D. S. FLETCHER 

and densely spined ; membranous fold slight ; apical third of aedeagus scobinate along left half 
of dorsal surface ; no cornutus. 

$. Not known. 

Measurement. o* 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 (Leopold ville). 

Holotype <$. [Congo (Leopold ville)] : Sankuru, Lusambo, 9.viii.i95o. 

Paratypes : [Congo (Leopoldville)] : Lusambo, 15. vi. 1949, 1 $ ; Uele, Paulis, 
i.v.1956, 1 (J ; all collected by Dr. M. Fontaine, all in Musee Royal de l'Afrique 
Centrale. 

Cleora epiclithra sp. n. 

(Text-figs. 39, 40 ; PI. 5, figs. 211, 212 ; Map 2) 

o*. 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 (PI. 5, fig. 
211). Under surface of wings tilleul buff, very weakly patterned with fuscous, as illustrated 
(PI. 5, fig. 212). 

o*. 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. 

$. Not known. 

Measurement. o* 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 ^. [Congo (Leopoldville)] : Kibali-Ituri, Nioka, 31.viii.1954 (/. Hecq), 
in Musee Royal de l'Afrique Centrale. 

Cleora carcassoni sp. n. 

(Text-figs. 41, 42 ; PI. 5, figs. 213, 214 ; Map 2) 

o*. 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 



o*. 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. 

$. Not known. 

Measurement. o* 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 <$. Kenya : Kakamega, ix.1961 (N. Mitton), genitalia slide Geo- 
metridae No. 5292. 

Paratype : Uganda : Ankole, Kalinzu Forest, xi.1961 (R. H. Carcasson), 1 <$ in 
Coryndon Museum. 




42 




Figs. 41-43. Chora genitalia. 41-42, carcassoni. 41, $ ; 42, aedeagus. 43, acerata, $. 



38 



D. S. FLETCHER 



Cleora herbuloti (Fletcher) comb. n. 

(Text-figs. 44-46 ; PL 5, figs. 215-222 ; Map 9) 

Neocleora herbuloti Fletcher, 1958 : 136, figs. 44, 85, 215. 
Neocleora herbuloti Fletcher ; Anon., 1963 : 4. 

c? $. External characters of colour and pattern vary geographically and are discussed under 
respective subspecies. 

o*. 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. 

$. 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. 




45 ^^ 46 

Figs. 44-46. C. herbuloti genitalia. 44, q* ; 45, aedeagus ; 46, $. 



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 9). 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) 

cJ. 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 wdng 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). 

$. Differs from male in the even and denser, darker irroration of the wings with bister and 
fuscous. 

Measurements. 6* 36-45 mm. ; $ 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 (PL 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. 



4 o 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 <$. Uganda : Kigezi, 5500 ft., (G. D. H. 
Carpenter), genitalia slide Geometridae No. 735. 

Fernando Po : Moka, 28.i-3.ii. 1933 (W. H. T. Tarns), 1 $. Nigeria : Near 
Ibadan, 1$. Mt. Cameroon : Musake, 6350 ft., 8.i. 1932 (M.Steele), ig. Angola: 
Quicolungo, 120 km. N. of Lucala, 800 m., iv.1936 (R. Brawn), 1 <$. Congo 
(Leopoldville) : W. Kivu, Upper Lowa River, nr. Masisi, 5000-6000 ft., ii.1924 
(T. A. Barns), 1 <$, all in British Museum (Natural History) ; Kivu, Rwankwi, 
15.viii.1947 (J. V. Leroy), 1 <J ; ibid., 13.xi.1947, 1 <J ; ibid., iv.1948, 1^,1$; 
Mulungu (Tshibinda), 31.L1956 (/. Hecq), 1 $ ; Mongbwalu (Kilo), 1937 (Me. 
Harford- J or dens), 1 <$ ; Sankuru, Djeka, 17.xii.1952 (Dr. Fontaine), 1 <$ ; Katako- 
Kombe, 2J.1952 (Dr. Fontaine), 1 <$ ; Kibali-Ituri, Kilo (Mines), 1955 (R. Andry), 

1 (J, all in Musee Royal de l'Afrique Centrale. Burundi : Astrida, c. 400 m., 
6.vii.i96i (Dr. Fontaine), 1 <$ in Musee Royal de l'Afrique Centrale. Uganda : 
Ankole, Kalinzu Forest, xi.1961 (R. H. Carcasson), 1 <$ ; Mabira Forest, Jinja, 
x.1962 (R. H. Carcasson), 2 $, all in Coryndon Museum ; Bundibugyo, 3440 ft., 
22.viii-3.ix. 1952 (D. S. Fletcher), 1 <J ; Entebbe, 25.viii.1961 (W. K. Brown), 3 $. 
Kenya : Mt. Kenya, vii.1930 (E. Barns), i<$ ; Nanyuki-Meru, 6.vii.i930 (E. Bams), 

2 (J. Tanzania : Amani, iii-iv.1936 (Cooper), 2 <$. Malawi : Mt. Mlanje, 
25.iv.1913 (Neave), 1 <$. Rhodesia : Mt. Selinda, 8-9.iii.1954 (H. Cookson), 1 $ 
and ibid., 9-17.iv.1956 (Van Son & Vari), 1 $, both in Transvaal Museum. 
Transvaal : Kowyn's Pass, Pilgrim's Rest Distr., 22.1.1962 (Vari & Leleup), 
1 <$ in Transvaal Museum. Cape Province : Transkei, Katberg, 1949 (H. B. D. 
Kettlewell), 1 <$. 

Cleora herbuloti phaea ssp. n. 

(PL 5, figs. 219-222 ; Map 9) 

$. Ground colour of wings vinaceous buff, white only at termen between veins M 3 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 (PL 5, fig. 220). 

$. Similar to male, but pattern largely obscured by dense fuscous black irroration (PI. 5, 
figs. 221, 222). 

Measurements. $ 38-41 mm. ; $ 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 g). Transvaal, Louis Trichardt. 

Holotype $• South Africa : Entabeni, nr. Louis Trichardt, ex pupa 3.viii.ig62, 
Div. of Entomology, S. Africa ; genitalia slide Geometridae No. 5242. 

Paratypes : Entabeni, nr. Louis Trichardt, 30.iii.1962, 1 $ ; data as holotype 
1 <J, 3 $• 

Cleora ace rata sp. n. 

(Text-fig. 43 ; PI. 5, figs. 223, 224 ; Map 9) 

cJ. 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. Mind wing white anteriorly, vinaceous buff 
posteriorly, densely irrorate and patterned with fuscous black ; medial and postmedial fasciae 
sometimes fused posterior of distal spot (PI. 5, fig. 223). Underside white ; fuscous black 
pattern of upper side marked in iron grey, but ill-defined (PI. 5, fig. 224). 

o*. 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. 

?. 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. o* 39-44 mm. ; $ 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 <$. Mt. Cameroon : Mann's Quelle, 7400 ft., 29J.1932 (M. Steele), 
genitalia slide Geometridae No. 5239. 

Paratypes : Mt. Cameroon : Musake, 6350 ft., 8.L1932 (M. Steele), 2 <$, 1 $ : 
ibid., 13.1.1932, 1 $ ; data as holotype, 2 ^ 1 $ ; ibid., 2.U.1932, 1 $. 

Cleora subcincta (Warren) 

(Text-figs. 47-50 ; PI. 6, figs. 231-241, 245 ; Map 9) 

Chogada subcincta Warren, 1901 : 16. 
Cleora subcincta (Warren) Prout, 1927 : 194. 
Neocleora subcincta (Warren) Herbulot, 1958 : 103. 

<J $. An obscurely marked species varying geographically in size and in colour of underside 
of wings ; these characters are discussed under respective subspecies. 



4 2 



D. S. FLETCHER 



o". 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. 




Figs. 47-50. Cleora genitalia. 47, subcincta subcincta, <$. 48-50, subcincta longifibulata. 

48, <J, left valve ; 49, aedeagus ; 50, $. 



REVISION OF AFRICAN SPECIES OF CLEORA 43 

$. 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 subcincta 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 Thome 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) 

o*. Underside of wings white, suffused with fusions, excepl a1 apex oi lore wing ; disca] 
spots and discal third of each wing more densely suffused than remainder (PI. 6, figs. 236-238). 

o*. Genitalia (Text-fig. 47). Serrate-edged process on sacculus one and one half times as 
long as greatest width of valve. 

$. 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. <J $ 33~34 mm. 

Distribution (Map 9). Sao Thome" ; endemic. 

Material examined. LECTOTYPE $. St. Thome, x-xi.1899 (Mocquerys), 
labelled by the author : Chogada subcincta Warr., type $, by present designation. 

Paralectotypes : St. Thome, x-xi.1899 {Mocquerys), 4 $. 

St. Thome, xii.1899-i.1901 {Mocquerys), 2 <y, 6 $ ; Sao Thome, 10. i-24.fi. 1926, 
edge of virgin forest (T. A. Barns), 5 <$, 5 $. 

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. 

o* $. 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. <J 32-38 mm. ; $ 34 mm. 

Distribution (Map 9). Angola ; Cameroun ; Congo (Leopoldville) ; Uganda ; 
Kenya ; Tanzania. 

Material examined. Holotype <J. Uganda : Bundibugyo, 3440 ft., 22.viii- 
3.LX.1952 {D. S. Fletcher), genitalia slide Geometridae No. 1755. 



44 D. S. FLETCHER 

Cameroun : Johann-Albrechts Hohe, 1896 (L. Conradt), 2 <$ ; ibid., 1898, 1 <$. 
Congo (Leopoldville) : Upper Uelle distr., v, 1 <$ ; E. Upper Ituri Valley, 30 miles 
S. of Irumu, 3000-3500 ft., vii.1924, dense forest (T. A. Barns), 1 g, all in British 
Museum (Natural History) ; Boma a Banana, 1933 (Dr. Van Hoof), 1 <$ ; Uele, 
Paulis, 14.ii.1956 (Dr. M.Fontaine), 1 $ ; ibid., 6.viii.i956, 1 <$ ; Mongbwalu (Kilo), 
1937 (Me. Harford- J or dens), 1^,1$; Kivu, Rwankwi, iii,iv,ix (/. V. Leroy), 4^ ; 
Kivu, Nyamunyunye (Mulungu), 20.xii.1955 (/. Hecq), 1 $ ; all in Musee Royal de 
l'Afrique Centrale. Uganda : Bwamba, v. 1958 (R. Carcasson), 1 $ and Bwamba 
Terr., ix.1961 (N. Mitton), 1 J, both in Coryndon Museum ; data as holotype, 1 $ 
and Kigezi, Mafuga forest, 25 miles N.W. Kabale, Rutenga, 7000 ft., 15.vii.1951 
(/. A. Burgess), 1 <$, both in British Museum (Natural History) ; Mabira forest, 
Jinja, x.1962 (R. H. Carcasson), 2 $ in Coryndon Museum. Kenya : Kakamega, 
v.1957 (Mrs. Board), 1 $ in Coryndon Museum. Tanzania : E. Usambara Mts., 
Amani, x.1953 (E. Pinhey), 3 <$. 



Cleora lacrymata sp. n. 

(Text-figs. 51-53 ; PI. 5, figs. 225, 226 ; Map 3) 

o*. 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 (PI. 5, fig. 226). 

o*. 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. 

$. 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. $ 40-43 mm. ; $ 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 <$. Ivory Coast : Bingerville, 1-5.viii.1915 (G. Melon), genitalia 
slide Geometridae No. 5255. 

Paratypes : Ivory Coast : Bingerville, 25.v-3.vi.1g15 (G. Melon), 1 <$ ; ibid., 
11. vi, 1 <J ; ibid., i-i4.vii, 2 $ ; ibid., 1— 5.viii, 1 9- ; ibid., 1-3. ix, 1 $. [Ghana] 
Gold Coast : Ashanti, Goaso (G. N. Gibbs), 1 <$. Nigeria : Mamfe, ii.1958, 1 <$ 
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 $ ; Fazenda Congulu, Amboim district, 






52 



Figs. 51-53. C. lacrymata genitalia. 51, $ ; 52, aedeagus ; 53, $. 



46 D. S. FLETCHER 

7-800 m., 12-16.iv.1934 (Dr. K. Jordan), 1 <$. [Central African Republic] 
Oubangui : Bangassou, vi-viii.1958 (P. Labour), 1 J in coll. C. Herbulot, Paris. 
[Congo (Leopoldville)] : Uele, Paulis, 12. iv. 1956 (Dr. M. Fontaine), 1 g in Musee 
Royal de l'Afrique Centrale. Kenya : Mt. Marsabit, 4500 ft., ii.1946 (T. H. E. 
Jackson), 1 $. 

Cleora echinodes sp. n. 
(Text-figs. 54-56 ; PL 6, figs. 242, 246 ; Map 3) 

(J. 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 (PI. 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 Cui 
(PI. 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. 

$. 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. $ 39-42 mm. ; $ 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 <$. Uganda: Entebbe, iii-v. 1895 (Jackson). 

Paratypes : Fernando Po : 650 ft., end of wet season (Rev. W. Cooper), 1 $. 
[Cameroun] : Afriq. Occid., Station Kamerun, Johann-Albrechts Hohe, 1896 
(P. Conradt), 1 <£. [Congo (Leopoldville)] : Upper Uelle distr., Dungu, v., 2 $ ; 
Yakusu (K. Smith), 1 <$, all in British Museum (Natural History) ; Uele, Paulis, 
18. hi. 1956 (Dr. M.Fontaine), 1$ ; Lusambo, 16. vi. 1949 (Dr. M.Fontaine), itf ; ibid., 
11.vii.1950, 1 cJ ; Sankaru, Katako-Kombe, 11. ix. 1952, 1 J ; Kafakumba, xii.1932 



REVISION OF AFRICAN SPECIES OF CLEORA 



47 



(F. G. Overlaet), i <$, all in Musee Royal de l'Afrique Centrale ; W. Kivu, south Lowa 
Distr., Lowowo Valley, 4000 ft., mountain forest, iii.1924, wet season (T. A. Bams), 
1 <$ ; Escarpment west of Semliki Valley, 20 mis. S.W. of Boga, 3500-4000 ft., 
vii.1924, borders of tropical forest and long grass country (T. A. Barns), 2 <$. 




Figs. 54-56. C. echinodes genitalia. 54, <$ ; 55, aedeagus ; 56, 



D. S. FLETCHER 



Uganda : Bwamba, iv-v. 1940 (T. H. E.Jackson), 2 $ ; ibid., x. 1942, 1^ ; Budongo 
v-vi.1939 (T. H. E. Jackson), 1 £ ; Budongo (G. W. Jeffery), 1 <$ ; Kampala, 
25.vii.1925, 1 (J. [Malawi] Nyasaland : Mt. Mlanje, 15.ii.1913 (S. A. Neave), 
1 J ; ibid., 17.iii.1913, 1 $. 

Cleora raphis sp. n. 

(Text-figs. 57-59 ; PL 5, figs. 227-230 ; Map 3) 

(J. 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 




Figs. 57-59. C. raphis genitalia. 57, <£ ; 58, aedeagus ; 59, $. 



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 (PI. 5, fig. 227). Under- 
side : apex of fore wing and termen of each wing, between veins Mz and Cu\, white ; remainder 
of wings suffused with fuscous, lightly round discal spots, densely elsewhere (PI. 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. 

o*. 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. 

$. (PI. 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. <? 42-44 mm. ; $ 40 mm. 

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 $. [Congo (Leopoldville)] : Kassai district (Taymans), genitalia slide 
Geometridae No. 2308 in British Museum (Natural History). 

Paratypes : [Congo (Leopoldville)] : Tumbulungu, 8.ix.ig30 (G. F. de Witte), 
1 $ ; Coquilhatville, 9.ix.i955 {Dr. M. Fontaine), 1 J ; Fl. Congo, Bolombo 
[3°59"S. 2i°24"E.], vii.1938 (J. Ghesquiere), 1 <$ ; Uele, Bambesa, 20.ix.1933 (/. 
Leroy), 1 $ ; Sankaru, Lusambo, 6.viii.i950 {Dr. M.Fontaine), i$ ; ibid., 8.viii.i95o, 
1 <J ; Bena-Dibele, xi.1921 (L. Verlaine), 1 <$, all in Musee Royal de l'Afrique 
Centrale. 

Cleora aculeata sp. n. 

(Text-figs. 60-62 ; PL 6, figs. 243, 244, 247, 248 ; Map 3) 

o*. 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 Cu\, 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 (PL 6, fig. 247). 



5° 



D. S. FLETCHER 



(J. 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. 

$. (PI. 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. 6* 3& mm. ; $ 40 mm. 




Figs. 60-62. C. aculeata genitalia. 60, aedeagus ; 61, o* ; 62, $. 



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 <$. [Nigeria] : Warri, vi.1897 (Dr. Roth), genitalia slide Geometridae 
No. 2217. 

Paratypes : data as holotype, 1 $ ; Agberi, Niger, io.v.1901 (Ansorge), 1 $. 



Cleora panarista sp. n. 

(Text-figs. 63-65 ; PI. 6, figs. 249-252 ; Map 3) 

<J. Vestiture white, irrorate with smoke grey and black ; patagia banded with black ; 
abdominal crest on first segment immaculate, second and third segments black dorsaily. 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\ (PI. 6, fig. 249). Underside white, patterned with fuscous black (PI. 6, 
fig. 250). 

o*. 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. 

$. (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. <J 40-45 mm. ; $ 49 mm. 

A species strikingly distinct in colour and pattern. Related closely to both 
raphis 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 <J. S. Rhodesia : Vumba Mts., ii.1956, genitalia slide Geometridae 
No. 5224. 



52 



D. S. FLETCHER 




Figs. 63-65. C. panarista genitalia. 63, $ ; 64, <? ; 65, aedeagus. 



REVISION OF AFRICAN SPECIES OF CLEORA 



53 



Paratypes : S. Rhodesia : Vumba Mts., ii.1956, 1 <$ ; ibid., Umtali, i— ii.1960, 
1 $. [Zambia] N. Rhodesia : Mwinilunga, v.1961, 1 $ in National Museum of 
Rhodesia. [Congo (Leopold ville)] : Elisabethville, 29.iii.1935 (Ch. Seydel), 1 <$ ; 
ibid., xii.1935, 1 $ in Musee Royal de l'Afrique Centrale. [Malawi] Nyasaland : 
Mt. Mlanje, 7.iii.igi3 (5. A. Neave), 1 $ ; ibid., 14.iv.1916, 1 <J. 



Cleora quadrimaculata (Janse) comb. n. 
(Text-figs. 66-68 ; PI. 7, figs. 253-257 ; Map 3) 

Neocleora quadrimaculata Janse, 1932 : 269, pi. 8:13; fig. 100. 

<J. 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 



Figs. 66-68. C. quadrimaculata genitalia. 66, $ ; 67, aedeagus ; 68, $. 




54 D. S. FLETCHER 

transverse fasciae slender and bister (PI. 7, fig. 253). Underside of wings tilleul buff, weakly 
suffused with pinkish buff and patterned with fuscous (PI. 7, fig. 254). 

o*. 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). 

$. Genitalia (Text-fig. 68). Lamella antevaginalis funnel-shaped, similar to that of pana- 
rista ; 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. $ 3 I_ 37 mm. ; $ 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 transversaria ; in the male the form of the sacculus is comparable with 
that of raphis, 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 <$. S. Africa : Pondoland, Port St. Johns, 1-7.X.1931 (Janse), in 
Transvaal Museum. Not seen. 

Material examined. S. Africa : Pondoland, Port St. Johns, 10-22.1.1955 
{A. J. T. Janse), irf, 1 $, in Transvaal Museum ; Port St. Johns, 12-30.vi.1923 
(R. E. Turner), 1 <$ ; ibid., v.1924, 1 $ ; Natal, Umkomaas, 25.L1913 (L. Hargreaves), 
1 $. Madagascar : Diego Suarez, 5.11.1917 (G. Melon), 1 $ ; ibid., 10-30.iv.1917, 
9 cJ, 1 $ ; ibid., 15-20.vi.1917, 5 $, 3 $ ; ibid., 16-29. vii.1917, x c?» 2 $ '■> Sakaramy, 
16.ii.1917 (G. Melon), 1 $ ; Tananarive, 1 $ ; Tananarive (Stichel), 2 <$ ; 
Analalava, 1 $. 



Cleora boetschi (Herbulot) comb. n. 
(Text-figs. 69-71 ; PL 7, figs. 258-261 ; Map 3) 

Neocleora boetschi Herbulot, 1961 : 495, fig. 3. 

o*. 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 (PI. 7, fig. 258). Underside of wings white, 
densely suffused and patterned with fuscous (PI. 7, fig. 259). 



REVISION OF AFRICAN SPECIES OF CLEORA 



55 




Figs. 69-71. C. boetschi genitalia. 69, $ ; 70, aedeagus ; 71, $. 



56 D. S. FLETCHER 

o*. 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. 

$. Wings white, lightly irrorate and patterned with bister (PI. 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 (PI. 7, fig. 261). 

$. 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. ; $ 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 raphis. 

Distribution (Map 3). Guinea ; Ghana ; Nigeria ; Cameroun ; Congo (Leopold- 
ville) ; Uganda. 

Material examined. Holotype $. Cameroun : N'long [Ngoulemakong], 18. i. 
1953 (R. 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 <$. 
Ghana : Wassaw dist. 45 miles inland from Sekondi, 1 <$. S. Nigeria : Ilesha 
(L. E. H. Humfrey), 2 <$. Cameroun : Batouri, hi. 1958, 1 £ in National Museum of 
S. Rhodesia ; Bitje, Ja River, x., wet season (G. L. Bates), 2 <$ and Johann-Albrechts 
Hohe, 1898 (L. Conradt), 1 $ in British Museum (Natural History). Congo 
(Leopoldville) : Coquilhatville, 9JX.1955 {Dr. M. Fontaine), 1 <$ ; Equateur, 
Bokuma, 20.xi.1911 (Rev. P. Hulstaert), 1 $ ; Eala, 14.lv. 1936 (/. Ghesquiere), 1 £ ; 
Uele, Paulis, 2-4.iv.1956 (Dr. M. Fontaine), 2 $ ; ibid., 13. xi. 1956, 1 $ ; ibid., 
5.VU.1958, 1 $ ; Uele-Itimbiri, La Kulu, 15. iv. 1930 (/. Van den Branden), 1 $ ; 
Sankuru, Katako-Kombe, 25.iv.1952 (Dr. M.Fontaine), ig ; ibid., 18.vi.1952, 1 $ ; 
ibid., 6.1X.1952, 1 <$ ; Lusambo, 17.vi-5.vii.1949 (Dr. M. Fontaine), 1 $ ; ibid., 
2.viii-3.ix.i950, 3 <$, 3 $ ; Kasai, Luluabourg, 18. vi. 1953 (Dr. M. Fontaine), 1 <$, all 
in Musee Royal de l'Afrique Centrale ; W. Kivu, Upper Lowa Valley, Nr. Masisi, 
5000-6000 ft., ii.1924, wet season (T. A. Barns), 1 <$ in British Museum (Natural 
History) . 

Uganda : Masaka, Katera, Sango Bay, x.1960 (R. H. Carcasson), 1 $, in 
Coryndon Museum. 



REVISION OF AFRICAN SPECIES OF CLEORA 57 

Cleora bicornis sp. n. 

(Text-figs. 72-74 ; PI. 7, figs. 262-265 ; Map 9) 

(J. 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). 

o*. 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. 




Figs. 72-74. C. bicornis genitalia. 72, $ ; 73, aedeagus ; 74, $. 



58 D. S. FLETCHER 

$. Similar to male, but underside of wings with broader and darker terminal bands, which 
are fuscous (PI. 7, figs. 264, 265). 

$. 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. $ $ 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 <$. W. Africa : Lagos, genitalia slide Geometridae No. 2281. 

Paratypes : Ivory Coast : Bingerville, 1-5.viii.1915 (G. Melon), 1 $ ; ibid., 
5-7.viii.1915, 1 $. Nigeria : data as type, 12 £, 3 $ ; Ilesha, (L. E. H. Humfrey), 
1 $. [Congo (Leopoldville)] : Stanleyville, 9.VL1948 (Dr. Faniel), 1 $ in Musee 
Royal de l'Afrique Centrale. 



Cleora dargei (Herbulot) comb. n. 
(Text-figs. 75-77 ; PI. 7, figs. 266-269 > Map 4) 
Neocleora 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 (PI. 7, fig. 266). Under- 
side white, suffused and patterned with bister (PI. 7, fig. 267). 

o*. 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. 269). 

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. $ 34~44 mm. ; 9 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. 




Figs. 75-77. C. dargei genitalia. 



76, aedeagus ; 77, $. 



60 D. S. FLETCHER 

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 <$. Cameroun : N'long [Ngoulemakong], i8.i. 
1953 (R. P. Boetsch), genitalia slide No. C.H.4027 in coll. Herbulot, Paris. 

Sierra Leone, i <$, 1 $ ; ibid. (A. B. Frere), 2 $, 2 $ ; ibid. (Major Bainbridge), 
1 $. Ivory Coast, i <$ ; Bating river, S of Touba, 1200 ft., 4.vii.i926 (C. L. 
Collenette), 1 <$ ; Bingerville, xi.1913 (Gaston Melon), 2 <J, 2 $ ; ibid., vi, viii-x.1915, 
9 <$, 11 $. Ghana : N. Territories, Kete-Krachi (A. W. Cardinall), 3 $ ; Kumasi, 
Odumase Swamp, 1913 (Smeed), 1 <$ ; Kumasi, ii-iii.1923 (N. E. Bell), 1 <$ ; Kumasi, 
ii-iii.1949 (W. Peters), 2 $, 1 $ ; Bibianaha [Bibiani], 700 ft., 10.vi.1912 (H. G. F. 
Spun ell), 1 $ ; Ashanti, Goaso (G. N. Gibbs), 1 <J. Nigeria : Ilesha (L. E. H. 
Humfrey), 2$ ; Lagos, 1 <J, 6 $ ; Warri, 1 $. Cameroun : Lolodorf, 1894-1895 
(L. Conradt), 1 $ ; Johann-Albrechts Hohe, 1898 (L. Conradt), 1 $. Gabon : 
Tchibanga (P. Rougeot), 1 <J. Congo (Brazzaville) ; Brazzaville, viii.1948 (P. 
Rougeot), 1 <J. Congo (Leopoldville) : Bopoto (Kenred Smith), 3 <$ ; Yakusu (K. 
Smith), 1 J, all in British Museum (Natural History) ; Leopoldville, 11. xi. 1953 
(Dr. M. Fontaine), 1 $ ; Leopoldville, Binza, 11.viii.1953 (Dr. M. Fontaine), 1 $ ; 
Ubangi, Gemena, i.1936 (/. Ghesquiere), 1 $ ; Equateur, Flandria, 24.ii-20.iv.1932 
(R. P. Hulstaert), 5 g ; Equateur, Bokota, 1927 (R. P. Hidstaert), 1 <$ ; Bokela, 
17.iv.1940 (R. P. Hulstaert), 1 <$ ; Eala, 20.X.1917 (R. Mayne), 1 $ ; Eala, xi.1936 
(/. Ghesquiere) , 1 $ ; Ifuta, 21.X.1921 (Verlaine), 1 $ ; Luebo, x.1930 (/. P. Colin), 
1 (J ; Stanleyville, vi.1929 (/. Colin), 1 <$ ; all in Musee Royal de l'Afrique Centrale ; 
Upper Uele Dist., Dungu, v, 1 $, in British Museum (Natural History) ; Uele, 
Rungu, 30. iv. i960 (Dr. M. Fontaine), 1 $ ; Uele, Paulis, i,iv,ix.i956 (Dr. M. 
Fontaine), 1 $, 2 $ ; Sankuru, Lusambo, 25-29. vi. 1949 (Dr. M. Fontaine), 2 $ ; 
ibid., 14-20.X.1949, 2 cJ ; ibid., iv.vii-ix.1950, 5 $, 4 $, all in Musee Royal de l'Afrique 
Centrale ; Kasai, Lusambo, ix-x.1919, 4 <$ in British Museum (Natural History) ; 
Sankuru, Djeka, 17.xii.1952 (Dr. M. Fontaine), 1 $ ; Sankuru, Katako-Kombe, 
i,ii,vii,ix-xii. 1951-53 (Dr. M. Fontaine), 9 <$, 4 $ ; Kivu, Rwankwi, 9.viii.i947 
(/. F. Leroy), 1 ^ ; ibid., 6-7. x. 1947, 1 <$ ; Rutschuru, i.1928 (CTz. Seydel), 1 (J, all 
in Musee Royal de l'Afrique Centrale ; Lindi River, Bafwasende, 2000 ft., vii.1921 
(T. A. Barns), 1 <$, in British Museum (Natural History). Uganda : Bundibugyo, 
3440 ft., 22.viii-3.ix.1952 (D. S. Fletcher), 1 $ ; Mpanga Forest, [Kampala], 2i.iv. 
1961, ex Entandrophragma angolense (W. K. Brown), 1 $ ; Mpanga Forest, 21.x. i960, 
ex Cupressus sp. (W. K. Brown), 1 $ ; Kaweri, 21.xii.1963, ex Pinus patula (W. K. 
Brown), 1 <$. 

Cleora dactylata sp. n. 

(Text-figs. 78-80 ; PI. 7, figs. 270-273 ; Map 4) 

cJ $. Externally closely similar in both sexes to dargei 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 



cJ. 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. 

$. Genitalia (Text-fig. 80). Lamella antevaginalis shallow and broad ; lamella post- 
vaginalis with a triangular, sclerotizcd 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. q* 31-42 mm. ; $ 43-46 mm. 

Closely related to dargei with which it occurs, examples of each species having 
been taken by Dr. Fontaine at Katako-Kombe on consecutive days. Differs 




Figs. 78-80. C. dactylata genitalia. 78, <J ; 79, aedeagus ; 80, $. 



62 D. S. FLETCHER 

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 <$. [Congo (Leopoldville)] ; W. Kivu, Nr. Masisi, Upper Lowa Valley, 
5000-6000 ft., forest and long grass, ii.1924, wet season (T. A. Barns), genitalia 
slide Geometridae No. 2297. 

Paratypes. [Congo (Leopoldville)] : data as holotype, 1 $ ; Nr. Walikili, 3000- 
4000 ft., forest, ii.1924, wet season (T. A. Barns), 2 $ all in British Museum (Natural 
History) ; Sankuru, Katako-Kombe, 27.xii.1951 (Dr. M. Fontaine), 1 <$ ; ibid., 
23-24.vii.1952, 2 <$ ; ibid., 10. ix. 1952, 1 <$ ; ibid., 15. xi. 1952, 1 $ ; ibid., n.i.1953, 
1 $ ; Lusambo, 24.vii.1949 (Dr. M. Fontaine), 1 <£ ; Luluabourg, 20.viii.1955 (Ch. 
Seydel), 1 $ ; Uele, Paulis, 18.ii.1956 (Dr. M. Fontaine), 1 <$ ; ibid., 12.iv.1956, 2 cj, 
1 $ ; ibid., 3. v. 1956, 1 <$ ; Boyenga, 1930 (Delpierre), 1 J ; Fomlioko, 20.ii.1921 
(Verlaine), 1 <$ ; Boende, i.iv.1940 (R. P. Hulstaert), 1 J, all in Musee Royal de 
l'Afrique Centrale. Uganda : Bwamba, v. 1956 (R. Carcasson), 1^,1$. 

Cleora thyris sp. n. 

(Text-figs. 81-83 ; PI- 8, figs. 274-280 ; Map 4) 

<J. 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 (PI. 8, fig. 275). 

o*. 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. 

$. 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. $ 38-50 mm. ; $ 40-50 mm. 

The species is variable in colour and pattern, displaying a range of poly- 
morphism similar to that of tulbaghata 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 thyris ; 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^. B.E.A. [Kenya] : Kyambu, 18.vii.1g19 [W. Feather). 




Figs. 81-83. C. thyris genitalia. 81, aedeagus ; 82, <$ ; 83, $. 



64 D. S. FLETCHER 

Paratypes ; Ethiopia : Negelli, v.1962 (S. Chojnacki), 2 <$. Kenya : N.F.D., 
Marsabit, 4500 ft., ii.1956 (/. G. Williams), 1 $, 3 $ ; ibid., vi.1955, 1 g in Coryndon 
Museum ; Mt. Marsabit, 4500 ft., ii.1946 (T. H. E.Jackson), 1J 1 ; 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 $ ; Nyeri, vii.1950 (.4. L. H. Townsend), 1 $ ; Nyeri 
(H. B. Kettlewell), 2 $ ; data as holotype, 12 <3\ 12 $ ; Nairobi, April-June (van 
Someren), 2 (J, ibid., 27.vi.1918, 1 <$, all in British Museum (Natural History) ; 
Nairobi, Karura Forest, iii.1949 (E. Pinhey), 1 $ ; Nairobi, iv-vi. (van Someren), 2 $ ; 
ibid., 27.vi.1918, 1 $ ; ibid., x.1923, 1 $ ; Nairobi, Thika [Road], v.1950 (E. Pinhey), 
1 <$, all in Coryndon Museum ; Meru District, Mt. Kenya, ix.1930 (Mrs. H. Young), 
1 $ ; Mt. Meru, Kenya distr., 5600 ft., xii.1920 (W. N. van Someren), 1 $, all in 
British Museum (Natural History). [Congo (Leopoldville)] : Kibali-Ituri, Mt. 
Rowa, 5.XU.1952 (/. Hecq), 1 <$ in Musee Royal de l'Afrique Centrale. [Tanzania] 
Tanganyika : W. Kilimanjaro, Ngare-Nairobi, 5000 ft., ii-iii.1937 (B. Cooper), 
1 $ ; Amani, 28. xi. 1934 (Mrs. Editha Dalton), 1 <$, both in British Museum (Natural 
History) ; Makoa, 7-27.L1959 (Lindner), 2 $, 1 $ ; ibid., 6-25.ii.1959, 2 (J, 2 $ in 
Zoological Museum, Stuttgart. [Malawi] Nyasaland : Mlanje, 30.iv.1913, 1 $, 
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, pi. 8 : 5 ; fig. 100. 

(J. 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 (PL 8, fig. 282). 

o*. 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. 

$. Similar in pattern to male ; upperside of wings less heavily, but more evenly irrorate 
with fuscous black (PL 8, figs. 283, 284). 

$. 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. $ 4 0_ 44 mm. ; $ 41-52 mm. 

The range of variation in colour and pattern in nigrisparsalis is similar to that 
found in both thyris and tulbaghata ; the similar forms in each species are virtually 
indistinguishable except by genitalia. Typical forms of nigrisparsalis (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. 




Figs. 84-86. C. nigrisparsalis genitalia. 84, $ ; 85, aedeagus ; 86, $. 



66 D. S. FLETCHER 

Distribution (Map 4). Natal ; Transvaal ; Mozambique ; Malawi ; Zambia ; 
Rhodesia ; Tanzania ; Kenya ; Burundi ; Congo (Leopoldville) ; Angola. 

Holotype <$. S. Rhodesia, Umtali, in Transvaal Museum, Pretoria ; not 
examined. 

Material examined. Natal : Pietermaritzburg, Malta, i-ii. 1927-8 {G. van Son), 
8 (J, 2 $ in British Museum (Natural History), 6 <$ in Transvaal Museum ; Natal 
National Park, Mont-aux-Sources, 30.1.1954, 1 J ; Muden, 15 hi. 1955 {H. Cookson), 
1 $ in Transvaal Museum. Transvaal : White River, ii.1909 (A. T. Cooke), 1 J ; 
Barberton {Harrison), 1 <$ ; Barberton {A. A. Williams), i.1933, 1 $ ; Marieps Mtn., 
6-14.1 (G. van Son), 3 $, 1 $, in British Museum (Natural History), 2 $ in Transvaal 
Museum ; Pretoria, i.1913 {Gladstone), 1 <$ ; Tshakoma, Zoutpansberg, xi.1931 
(G. van Son), 1 $ ; Magoebaskloof, iv.1933 [Rev. D. P. Murray), 1 $ ; Erasmus 
Reserve, Pilgrim's Rest District, 30. iv. i960 {F. Neubecker), 3 <$ ; Kowyn's Pass, 
Pilgrim's Rest District, 22.ii.1962 {Vari & Leleup), 1 $, all in Transvaal Museum. 
Mozambique : Dondo, ix.1954 {H. Cookson), 1 $. Malawi : Zomba, 3000 ft., 
iv.1913 {E. Ballard), 1 <$ ; Zomba, i.1921 {H. Barlow), 2 <$, 1 $ ; ibid., v. 1920, 1 $, 
all in British Museum (Natural History). Rhodesia : Nr. Bulawayo, Khami, 
hi. 1956, 2 (J ; Marandellas, ii,x,xi.i96o, 3 <$, 1 $ ; Vumba Mts., Umtali, iii,xi,xii, 
4 $ ; Umtah District, i-iii, 2 $, all in National Museum of Rhodesia ; Umtali, 
i-iii,vi, 1 $, in Transvaal Museum, 2 ^ in British Museum (Natural History). 
Salisbury, iii,iv,v,vii, 5 $ in National Museum of Rhodesia, 2 <$ in British Museum 
(Natural History) ; Inyanga, iv.1961, 1 $ in National Museum of Rhodesia ; 
Penalonga {0. A. Kidwell), 1 $ ; Mt. Selinda, 17-31.L1959 {G. van Son), 1 $ ; Mt. 
Selinda, 8-9. iii. 1954 {H. Cookson), 1 $, all in Transvaal Museum ; Victoria Falls, 
io.iv.1927 {R. H. R. Stevenson), 1 <$ ; Shamva, ii.1921 {O'Neil), 1 $ ; Mountain Inn, 
Melsetter, xi.1950 {H. B. Kettle-well) , 1^,1$. Zambia : Kashitu, 14.V.1915 {H. C. 
Dollman), 1 g ; N'dola, 4.1.1923, 1 <$, all in British Museum (Natural History) ; 
Abercorn, ii-iii.1954 {D. Vesey-Fitzgerald), 2 <J, both in Coryndon Museum. 
Tanzania : Makoa, 7-27.1.1959 {Lindner), 1 $ in Zoological Museum, Stuttgart ; 
Mamboia {Dr. Baxter), 1 $ in British Museum (Natural History). Kenya : Nairobi, 
v.1927 {D. M. Hopkins), 1 $ ; ibid., i.1953 {E. Pinhey), 1 $ in Coryndon Museum ; 
ibid., 8JX.1927 {Mrs. D. M. Hopkins), 1 <$, 1 $ ; ibid., x.1920 {W. N. van Someren), 
1 $ ; ibid., ii.1928, 1 $ ; ibid., 8JV.1912 (C. Montague Smyth) , i$ ; ibid., bred 1937 
{R. H. Simmonds), 1 $ ; ibid., 5500 ft., iii.1939 {Maclnnes), 1 $ ; Nairobi Plains, 
Kikuyu, 5.V.1900 {R. Crawshay), 1 $ ; Makueni, Machakos [District], vi.1958, 1 $ 
in Coryndon Museum ; Ruiru, vi.1963 {P. E. Wheatly), 3 $, 3 $ ; Mt. Kenya, 
vii.1930 {E. Barns), 5 $ ; Ndarugu, 18-19.vi.1917 {W. Feather), 2 $ ; Suna, S. 
Kavirondo, i.1932 {W. Feather), 4 <$, 1 $, all in British Museum (Natural History). 
Burundi : Kitega, iii,vi-x.i962 {Dr. M. Fontaine), 35 £, 17 $. Congo (Leopold- 
ville) : Kibali-Ituri, Nioka, vi,vii,x,xi (/. Hecq), 2^,4$ all in Musee Royal de 
l'Afrique Centrale ; Elisabeth ville, i-iv, x-xii {Ch. Seydel), 9 <$ in Musee Royal de 
l'Afrique Centrale, 7 $, 2 $ in British Museum (Natural History). Angola : Bihe, 
xii. 1934-1.1935 {E. Braun), 2 $. 



REVISION OF AFRICAN SPECIES OF CLEORA 67 

Cleora tulbaghata (Felder) comb. n. 

(Text-figs. 87-89 ; PI. 9, figs. 289-296, 301 ; Map 4) 

Boarmia tulbaghata Felder, 1875, pi. 125 : 5. 
Chogada acaciaria sensu Warren, 1904 : 474. 
Chogada acaciaria ab. flavipleta Warren, 1904 : 474. 
Chogada acaciaria ab. fumata Warren, 1904 : 474. 
Neocleora munda sensu Janse, 1932 : 267, text-fig. 100. 

o*. 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. 

?. 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. <J 33 _ 49 mm - '• ? 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 (PI. 9, fig. 289) ; the underside of the wings is white, patterned with fuscous 
(PI. 9, fig. 290). In this form the female is similarly patterned, but the irroration, 
which is sparse, and the transverse fasciae are bister (PI. 9, fig. 291). 

The holotype (PI. 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. 9. 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 (PI. 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 (PI. 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 (PI. 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, pi. 5), who 
found it resting on the fire-blackened trunks of a species of Acacia. In a 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 cy clops 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 



68 



D. S. FLETCHER 




Figs. 87-89. C. tulbaghata genitalia. 87, aedeagus ; 88, <J ; 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 10 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 $ (without abdomen). Knysna, C. W., ex coll. 
Felder. 

Kenya : Suna, S. Kavirondo, iv.1930 (W. Feather), 1 <J ; ibid., i.1931, 1 <J ; 
Kibwezi, n.i.1917 (W. Feather), 1 $. Zambia : (Gimson), 2$, 1 $. Mozambique : 
Lorenzo Marques (Distant Coll.), 1 $ ; Lorenzo Marques, 3^.1949 (H. B. Kettlewell), 
1 <$ ; Magude, x.1918 (C. /. Swierstra), 1 $ ; Delagoa Bay (Monteiro), 2$. 
Transvaal : Barberton (Harrison), 2 $ ; Three Sisters, 28.ii.19n {A.J. T.Janse), 
1 $. Zululand : (G. F. Leigh), 1 <$, 1 $ in British Museum (Natural History) ; 
Eshowe, i.1956 (D. A. Swanepoel), 1 $ in Transvaal Museum. Natal : Durban, 
Umhlanga, 13-20.viii.1950 (H. B. D. Kettlewell), 2 <$ ; Umhlanga Rocks, 6.H.1955 
(C. G. C. Dickson), 1 $ in Transvaal Museum ; Durban (G. F. Leigh, Col. Bowker, 
Stella Bush, C. G. C. Dickson), 26^, 18 $ in British Museum (Natural History), 3 J\ 

1 $in Transvaal Museum ; Port Natal (Guenzius), 3 (J, 2 $ ; Pinetown (G. F. Leigh), 

2 (J, 1 $ ; Krantzkloof, 18.1.1918 (L. Hargreaves), 1 $ ; Umkomaas, 25.xii.1910 (L. 
Hargreaves), 1 <$ ; ibid., 8.11.1913, 1 <$ ; Port Shepstone, 12. i. 1924 (Lt. Col. G. R. 
Oakes), 1 <$. Cape Province : E. Griqualand (Kendall), 1 $, 1 $ ; E. Cape, 
Kamanassie, ix.1949 (H. Bridges), 1 <J in Transvaal Museum ; Pondoland, Port St. 
Johns, i-v.1924 (R. E. Turner), 5 $, 7 $ ; ibid., 23-26. viii. 1950 (H. B. D. Kettlewell), 
2 (J, 1 $ ; ibid., 10-22.L1955 (A. J. Janse), 3 <$, 1 $ in Transvaal Museum ; W. 
Pondoland, Nggeleni, 17.iii.1904 (H. H. Si&inny), 1 $ ; East London, 19.iii.1913 
(G. F. Leigh), 2<$, 1 $ ; Grahamstown, 1 $ ; Port Elizabeth, ii,iii,iv (C. G. C. Dickson) 
1 3 1 , 2 $ in Transvaal Museum ; Groot River Pass, 22-23. i. 1955 (A. J- T. Janse), 
1 <$ in Transvaal Museum ; Knysna, C.W. (ex coll. Felder), 3 $, 1 $ ; Wilderness, 
Knysna Forest, 20.x-8.xi.1952, at M.V. light (H. B. D. Kettlewell), 23 $, 12 $ ; 
ibid., 6-20.ii.1934, 1 (J, 4 $ ; Knysna, 10-20.1.1958 (A. J. T. Janse), 1 <$, 2 $ and 
24. vi. 1950 (C. G. C. Dickson), 1 $ in Transvaal Museum ; Belvedere, 10.iii-3.iv.1921 
(Dr. H. G. Breijer), 1 <$, 2 $ ; Harkerville, 9.iii.i92i (Dr. H. G. Breijer), 1 $ ; Mossel 
Bay, 15.iii-20.iv.1932 (R. E. Turner), 1 $ ; Cape Town, ix.1877, 1 $ ; ibid., i.1871, 
1 $ ; ibid., v.1912 (Lord Gladstone), 1 $ ; ibid., 14-21. i.1961 (C. G. C. Dickson), 1$, 
1 $ in Transvaal Museum ; Cape Town, Camps Bay, i,ii,viii,ix,xi,xii (A. J. Dtike), 
5 (J, 1 $ in Transvaal Museum ; Cape Town, Seapoint, 13.x. 1953 (A.J. Duke), 1 $ in 
Transvaal Museum ; Mowbray, ii.1950 (A. J. Duke), 1 <$ in Coryndon Museum ; 
Cape Town, Kirstenbosch, 5-29. xii. 1954 (A. T. J. Janse), 2 <$, 1 $ and Clovelly, 
ii,iv,xii (D. A. Swanepoel), 3 <$, 1 $ in Transvaal Museum ; Simonstown, (H. G. 



7° 



D. S. FLETCHER 




Figs. 90-92. C.plax genitalia. 90,^ ; 91, aedeagus ; 92, $. 



REVISION OF AFRICAN SPECIES OF CLEORA 71 

Hunt), 3 $ ; Fish Hoek, x-xi.1949 (H. B. D. Kettlewell) , 2$ ; ibid., 1949-1950, 6<$, 
1 $ ; ibid., 28.iii.1950, 23 $, 5 $ ; ibid., 15. iv. 1950, 1 (J ; ibid., i-iii.1950, at light, 

22 (J, 7 $ ; Fish Hoek Valley, 10.xi.1952-3.iii.1953, M.V. light (H. B. D. Kettlewell), 

2 3 c?» 9 9 ', Fish Hoek-Kommetje, 1951 (H. B. D. Kettlewell), 1 <$. 

Cleora plax sp. n. 

(Text-figs. 90-92 ; PL 8, figs. 287, 288 ; Map 4) 

o". 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 (PI. 8, fig. 287). Underside of 
wings white, suffused with fuscous (PI. 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. 

$. 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. 6* 40-41 mm. ; °. 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 <y. Uganda : Masaka, Katera, Sango Bay, x.1960 (R. H. Carcasson), 
genitalia slide Geometridae No. 5289. 

Paratypes : [Congo (Leopoldville)] : Escarpment, west of Semliki Valley, 20 
mis. 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, i(Jin British 
Museum (Natural History) ; Mongbwalu (Kilo), 1937, (Me. Harford- J or dens), 1 $ ; 
Kilo, i.1940 (RR. FF. Maristes), 2 <$ ; Mambunga via Boma (R. P. Bittremieux) , 
1 $, all in Musee Royal de l'Afrique Centrale. 

Cleora munda (Warren) comb. n. 
(Text-figs. 93-95 ; PL 9, figs. 297-300, 302 ; Map 4) 
Chogada munda Warren, 1899 : 52. 

o*. Genitalia (Text-figs. 93, 94). Medial plate of gnathus narrowly rounded, as in tulbaghata ; 



7 2 



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. 

$. 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. 6* 3 D ~45 mm. ; $ 42—50 mm. 




Figs. 93-95. C. munda genitalia. 93, 5* ; 94. aedeagus ; 95, $. 



REVISION OF AFRICAN SPECIES OF CLEORA 73 

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 $. Zululand : Edukumbaan Hills, v.1895, 
genitalia slide Geometridae No. 2145. 

Kenya : Kitale, v. 1953 (C. R. Howard), 1 $ in Coryndon Museum ; Nairobi 
{van Someren), 4 <J ; Kenya coast, Shimba Hills, xii.1961 (R. H. Carcasson), 1 $ in 
Coryndon Museum. Tanzania : Usambara, Amani, iv.1952 (C. Howard), 1 <J in 
Coryndon Museum ; E. Usambara, Amani, ii.1953 (E. Pinhey), 1 $ in Coryndon 
Museum ; Amani, iii-iv.1936 (B. Cooper), 2 $, 2 $ ; Amani, 28. xi. 1934 (Mrs. Edith 
Dalton), 2 $. Malawi : Mlanje, 10.iv.1913, 1 <$ ; Mlanje Boma, 2400 ft., 27.iv.1910 
(S. A. Neave), 1 $ ; Chintiche (T. H. Lloyd), 1 $. Rhodesia : Mt. Selinda, 17-31. 
i.1959 (G. van Son), 1 <$ ; ibid., 9-17.lv. 1956 (G. van Son & L. Vari), 1 <$, 1 $ in 
Transvaal Museum ; Umtali District, 16. i. 1933 (P. A. Sheppard), 1 $ ; ibid., 23.iii. 
1934, 1 $, both in National Museum of S. Rhodesia. Transvaal : Zoutpansburg, 
Tshakoma, xi.1931 (G. van Son), 1 $ ; Magoebaskloof Rest Camp, 4000 ft., 21.ii.1960 
(R. F. Lawrence), 1 <$ ; Cyprus Farm, near Ofcolaco, 24-28.ix.1961 (L. Vari), 2 $ ; 
Mariepskop, 16. iv. 1955 (H. Cookson), 1 <$, all in Transvaal Museum ; Marieps Mtn., 
5. xii. 1925-6. i. 1926 (G. van Son), 5 <3\ 2 $ ; Pilgrim's Rest Distr., Erasmus Reserve, 
30.iv.1960 (F. Neubecker), 4 <$, 1 $ in Transvaal Museum ; Pilgrim's Rest Distr., 
Kowyn's Pass, 22.ii.1962 (L. Vari & Leleup), 1 $ in Transvaal Museum ; Louws 
Creek, xi.1922 (H. G. Williams), 1 <$. Natal : Hluhluwe, 16.iii.1952 (Dr. G. van 
Son), 1 (J in Transvaal Museum ; Pinetown, 6.iii.i9io (G. F. Leigh), 1 <$ ; Durban, 
ii,v,vi,x (C. G. C. Dickson), 7 <$, 1 $ in Transvaal Museum ; Pietermaritzburg, 
Malta, i-ii.1928 (G. van Son), 4 <J, 3 $ in British Museum (Natural History), 2 $ in 
Transvaal Museum. Cape Province : Port St. Johns, 10-22.fi. 1955 (A. J. T. 
Janse), 2 $ in Transvaal Museum. 

Cleora lamottei (Herbulot) comb. n. 

(Text-figs. 96-98 ; PL 10, figs. 305-308, 320 ; Map 5) 

Neocleova lamottei Herbulot, 1954 : 322, pi. 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. 10, fig. 305). Underside of wings white, densely suffused with bister 
(PL 10, fig. 306). 

o*. 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 




Figs. 96-98. C. lamottei genitalia. 96, aedeagus ; 97, o* ; 98, $. 



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 (PL io, fig. 307). Differs in underside in reduction of bister suffusion in proxi- 
mal half of each wing (PI. 10, fig. 308). 

$. 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. 6* 34~39 mm - '• ? 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 (PL 10, 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 tulbaghata 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 (Leopold ville) ; Principe I. ; Sao Thome" I. 

Material examined. Holotype g. Guinea : Monte Nimba, 30.viii.1951 (M. 
Lamotte & R. Roy) in Museum national d'Histoire naturelle, Paris. 

St. Principe : 1500-2000 ft., iv-v.1928 (T. A. Barns), 10 <$, 5 $. Sierra Leone : 
1 $. Ivory Coast : Bingerville, v,vi,viii-xii (G. Melon), 38 <$, 18 $ ; Agboville, 
1-8.vi.1915 (G. Melon), 1 $. Ghana : A'koon, 4.1.1919 (G. Harrison), i<$ ; Ashanti 
{Mrs. D. Houston), 2 $ ; Ashanti, Goaso (G. H. Gibbs), 1 $ ; Bibianaha [Bibiani] 
70 miles N.W. of Dimkwa, 700 ft., ii,v (H. G. F. Spurrell), 2 <$, 2 $ ; Coomassie 
[Kumasi] (H. Whiteside), 3 <$, 2 $ ; Ashanti, Juaso (G. H. Gibbs), 1 <$ ; Abossi {Dr. 
J. J. Wilson), 1 $ ; Tarkwa, 3 <$, 2 $ ; Wassaw distr., 45 miles inland from Sekondi, 

1 $ ; Sekondi, xii.1921 (N. E. Bell), 1 $. Nigeria : Lagos, 2 <$ ; Ilesha {Capt. 
L. E. H. Humfrey), 1 $, 1 $ ; Warri, iv-viii.1897 {Dr. Roth), 4 <$, 2 $ ; Degama 
[Degema] {Dr. Ansorge), 1 <$ ; Ikom, 18. v. 1930 {E. Haig), 1 $ ; Old Calabar, 2 <$, 

2 $. Cameroun : Johann-Albrechts Hohe, 1898 (L. Conradt), 7 (J, 2 $ ; Lolodorf, 
1894-1895 (L. Conradt), 6 £ ; Bitje, Ja River, x, wet season {G. L. Bates), 1 $ ; 
Epulan [Efulen], 2.1V.1926 {G. Schwab), 1 <$. Gabon : Ogowe, 1890-1894 (L. 
Gazengel), 2 $ ; Lastourville {P. C. Rougeot), 3 $ ; Tchibanga, iv,xi {P. C. Rougeot), 
6 (J. Congo (Leopoldville) : Kitobola, 191 1 {Rovere), 2 <$ ; Equateur, Bokote, 
19.1.1926 {R. P. Hidstaert), 1 $ and Tshuapa, Flandria, 18. x. 1947 (Rev. P. Hulstaert), 
1 $, in Musee Royal de l'Afrique Centrale. 



76 



D. S. FLETCHER 



Cleora toulgoetae (Herbulot) comb. n. 
(Text-figs. 99-101 ; PI. 10, figs. 309-312 ; Map 5) 

Neocleora toulgoetae Herbulot, 1961 : 493, fig. 1. 

6*. 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 (PI. 10, fig. 310). 

o*. 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 




Figs. 99-101. C. toulgoetae genitalia. 99, aedeagus ; 100,5* '< IOI » ?• 



REVISION OF AFRICAN SPECIES OF CLEORA 77 

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. 

$. 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 (PI. 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. $ $ 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.fnmata. Structurally the form of the female 
genitalia suggest an affinity with the preceding species, lamottei. 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 $. 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 $. Ivory Coast : 2$, 2 $ ; Bingerville, 
1915 (G. Melon), 1 S, 1 $ ; Agboville, 1-8.vi.1915 (G. Melon), 1 $. Ghana : 
Bibianaha [Bibiani], 70 miles N.W. of Dimkwa, 700 ft., 19.x. 1910 (H. G. F. Spnrrell), 
1 $ ; ibid., 28.1.1912, 1 $ ; Coomassie [Kumasi], ii-iii.1923 (N. E. Bell), 1 <$, 1 $ ; 
Enchi (Capt. B. D. Peake), 1 $ ; Manso, N.E. of Cape Coast, iii-iv.1922 (N. E. Bell), 
1 $. Nigeria : Lagos 1 $ ; Ilesha (L. E. H. Humfrey), 1 $ ; Old Calabar (F. C. 
Pudney), 1 $. Fernando Po : (W. Cooper), 1 J. Cameroun : xi. (G. Schwab), 
1 $ ; Johann-Albrechts Hohe, 1898 (L. Conradt), 1 $ ; Bitji, Ja River, 2000 ft., 
wet season, iv-v.1909 (G. L. Bates), 1 <$ ; ibid., ix-xi.1911, 1 <$. Gabon : Lastour- 
ville (P. C. Rougeot), 1 $. Angola : Quicolungo, 120 km. W. of Lucala, 800 m., 
iv.1936 (R. Braun), 1 $. Congo (Leopoldville) : Eala, ix.1936 (/. Ghesqniere) , 1 $ 
in Musee Royal de l'Afrique Centrale. Dungu, Upper Uele distr., iv, 1 $ ; Middle 
Lowa Valley, Nr. Walikali, 3000-4000 ft., forest, ii.1924, wet season (T. A. Barns), 
1 <$. Burundi : Usumbura, 900 m., 8.xii.ig6i {Dr. M. Fontaine), 1 <$. 



Cleora rostella sp. n. 

(Text-figs. 102-104; PI. 10, figs. 313-319; Map 5) 

o*. 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 



7 8 



D. S. FLETCHER 




Figs. 102-104. C- vostella genitalia. 102, $ ; 103, <$ ; 104, aedeagus. 



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 Mi 
and A/3 ; cubital veins and other veins distad of postmedial fasciae interruptedly warm buff ; 
a lateral fuscous streak between veins A/ 2 and M 3 on fore wing, extending from postmedial 
fascia to termen (PI. 10, fig. 313). Underside white patterned with fuscous (PI. 10, fig. 314). 

o*. 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 aedcagus incised ; vesica with two stout, tapered cornuti fused at base, 
one one-half and one three-eighths as long as aedeagus. 

$. 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, 
fi gs- 3*5- 3i6). 

$. Genitalia (Text-fig. 102). Lamella postvaginalis strongly sclerotized medially ; posterior 
margin of colliculum sharply defined ; anterior sixth of bursa copulatrix dilate and membranous, 
remainder cylindrical, ribbed and sclerotized. 

Measurements, o* 36-42 mm. ; $ 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 tidbaghata ; 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 tidbaghata 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. [Malawi] Nyasaland : Limbe, ix-x.1926 (H. Barlow), genitalia 
slide Geometridae No. 2336. 

Paratypes : S. Rhodesia : Chirundu, on Zambesi, 22.ii.1950 (N. Mitton), 2 <$ in 
Transvaal Museum : Salisbury, 17.iv.1916, 1 $ in National Museum of Rhodesia. 
[Zambia] N. Rhodesia : N'kana (L. Ellison), 1 <$ ; Chisorwe [i4°52"S. 29°o5"E.], 
Luano Valley, 11.ii.1928 (M. Burr), 1 <$ ; Mpeta, Loangwa R., aff. of Zambesi, 
xi-xii.1895, beginning of rainy season (Coryndon), 3 <$. [Malawi] Nyasaland : 
Zomba, 3000 ft., iv.1913 (E. Ballard), ig ; Zomba, vi.1923 (H. Barlow), 1 $ ; Mlanje 
ii.1925 (H. Barlow), 1 $ ; Magunda Estate, Luchenza (F. Nisbet), 1 $. [Tanzania] 
Tanganyika : Old Shinyanga, 29.L 1954 (E. Burtt), 1 $. Uganda : Kampala, 
15.fi. 1932 (H. Hargreaves), 1$. Sudan : Tambura, Southern Bahr-el-Ghazal, 4 <J. 



80 D. S. FLETCHER 

[Congo (Brazzaville)] Congo Republic : Fort Crampel, 2 <J. Burundi : 
Usumbura, 900 m., 6-7.vii.1961 (Dr. M. Fontaine), 2$, 1 $ ; ibid., 13.viii.1961, 1 $ ; 
ibid., i.xi.1961, 1 $ ; ibid., 11.xii.1961, 1 $ ; ibid., 31.xii.1961, 1 <$ ; ibid., 7-10.L 
1962, 1 (J, 1 $ ; Kitega, 5. v. 1962 (Dr. M. Fontaine), 1 $, all in Musee Royal de 
l'Afrique Centrale. [Congo (Leopoldville)] Belgian Congo : Region de M'Pala 
(R. P. Guilleme), 1 $ ; E. Lake Kivu, ix.1919 (T. A. Barns), 1 $ ; 150-200 miles 
W. of Kambove, 3500-4500 ft., 29.X.1907 (S. A. Neave), 1 <$ ; Katanga Distr., 
Kafakumba, i.1927, 1 <$ ; S.E. Katanga, 4000 ft., 24.xi.1907 (S. A. Neave), 1 <$, all in 
British Museum (Natural History) ; Elisabeth ville, 26. ix. 1932 (Ch. Seydel), 1 $ ; 
ibid., 27.X.1937, 1 (J ; ibid., 16. xi. 1940, 1 $ ; ibid., x.1948, i$, 2 $ ; ibid., 20. xi. 1949, 
1 $ ; ibid., 9.1V.1950, 1 $ ; ibid., 15.1.1951, 1 $, all in Musee Royal de l'Afrique 
Centrale; Elisabeth ville, 23.X.1951 (Ch. Seydel), 1 $ ; ibid., 24-29. xi.1951, 1 <$, 1 $ ; 
ibid., 23.xii.1953, 1 $ ; ibid., 14.xii.1954, 1 $ ; ibid., 7.L1956, 1 <J ; ibid., 25L1956, 
1 $ ; ibid, 2.XL1956, 1 $ ; ibid., 22.xii.1956, 1 <$ ; ibid., i.i.1957, 1 $ ; ibid., 
28.1.1957,1 $, all in British Museum (Natural History) ; Elisabethville, r.Lubumbashi, 
11-24.vi.1914 (Overlaet), 1 <$, 1 $ ; Lubumbashi, xi.1926 (Ch. Seydel), 1 ^ ; Katanga, 
Muteba, xii.1922 (C/z. Seydel), 1 ^ ; Katanga, Luashi, 1933 (Freyne), 2$; ibid., 1935, 
1 cj ; Katanga, Luashi, 3.vii.i924 (Ch. Seydel), 2$ ; Lulua, Kapanga, 13-14. xi. 1932 
(i 7 . G Overlaet), 4^ ; ibid., i.1933, 1 S '< ibid., viii.1933, 1 £ ; Lulua, Sandoa, iv.1932 
(F. G. Overlaet), 1 <$, all in Musee Royal de l'Afrique Centrale ; Kasai, Lusambo, 3-10. 
ix.1919, 1 (J, 1 $ ; Upper Kasai Distr. (P. Landbeck), 1 $, all in British Museum 
(Natural History) ; Kasai, Luluabourg, 12. vi. 1952 (Dr. M. Fontaine), 1 $ ; ibid., 
6.VL1953, 1 $ ; Luluabourg, 24.X.1921 (Verlaine), 1 $ ; Lusambo, 8.L1926 (Ch. 
Seydel), 2$ ; Sankuru, Katako-Kombe, 15. xi.1951 (Dr. M.Fontaine), 1^ ; Sankuru, 
Dimbelenge, 19.V.1953 (Dr. M. Fontaine), 1 <$ ; Kilo 300 de Kindu (Dr. Russo), 
1 <$ ; Kilo 345 de Kindu (Dr. Russo), 1 $ ; Leopoldville, 12.viii.1953 (P. Jobels), 1 $ ; 
ibid., 6.xii.i955, 1 <$ ; Leopoldville, 29.xi.1953 (Dr. M. Fontaine), 1 $, all in Musee 
Royal de l'Afrique Centrale ; Upper Uele distr., Dungu, ix, 1 $ in British Museum 
(Natural History). Angola : Gamba, Bihe, xii.1934 (R. Braun), 7 <$ ; ibid., 
i- I 935> 3 c? ! Talala, Benguella, 1 xii.1905 (Dr. Ansorge), 1 <$. 

Cleora legrasi (Herbulot) comb. n. 
(Text-figs. 105-107 ; PI. 11, figs. 325-329 ! Map 5) 

Neocleora legrasi Herbulot, 1955 : 40, fig. 2. 

<J. 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 : c? 34~44 mm. ; $ 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 




Figs. 105-107. C. legrasi genitalia. 105, 5* ; 106, aedeagus ; 107, $. 



82 D. S. FLETCHER 

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 
legrasi closely to the preceding species, rostella. 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 $. Madagascar centre, Tananarive, Pare de 
Tsimbazaza, alt. 1200 m., 3.11.1952 {P. Viette), in Museum national d'Histoire 
naturelle, Paris. 

Madagascar : Diego Suarez, ii,iii,iv,vii.ic)i7 (G. Melon), 9 <$, 1 $ ; N.E. 
Madagascar, base de la montagne Tsaratanana (ex Lamberton, 1923), 1 $ ; E. 
Madagascar, Marcantsetra, 2 $ ; Central Madagascar, 2500 ft. (F. B. Pratt), 2 <$, 
1 $ ; Tananarive, 1 $ ; Station Perinet, 149 km. east of Tananarive, i,ii,iii,x,xi,xii 
(N. & G. Olsoufieff), 78 <$, 18 $ ; Ambinanindrano, 50 km. west of Mahonoro, i,v 
(G. K. Kendall-Cornish), 2$, 1 $ ; Mananjary, 1918 (G. Melon), 8^, 1 $ ; Mananjara, 
xi.1918, 4 c? ; Ankarampotsy, near Fianarantsoa, col. de Tantamaly, xii. 1933-1.1934 
(R. Catala), 5 <$, 1 $ ; Betsileo, 3000-4000 ft., 1881 {Deans Cowan), 1 <$ ; S. 
Madagascar {Me. Lamberton), 2 <$. 



Cleora angustivalvis (Herbulot) comb. n. 
(Text-figs. 108-110 ; PI. 10, figs. 321-324 ; Map 5) 

Neocleora angustivalvis Herbulot, 1965 : 121. 

o*. 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 (PI. 10, fig. 321). Underside white, patterned with fuscous (PI. 10, fig. 322). 

c?. 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. 

$. 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 M±, the smaller between 
veins M% and Cu± ; veins on both wings interruptedly warm buff distad of postmedial fascia 
(PI. 10, fig. 323). Underside differs in the reduction of fuscous pattern, especially on hind wing 
(PL 10, fig. 324). 

$. Genitalia (Text-fig. no). 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 : o* 35 mm. ; $ 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, rostella 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. 



F . : 





Figs. 108-110. C. angustivalvis genitalia. 108, aedeagus ; 109, $ ; no, 9- 



84 D. S. FLETCHER 

The species is distinguished externally from both rothkirchi insulamm 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 angitstivalvis a 
grey appearance by comparison with males of transversaria 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 <$, 11 $ ; ibid., 1894 (L. Humblot), 6 $ ; Grande 
Comoro, viii.1911 (G. F. Leigh), 1 $. 



Cleora serena sp. n. 

(Text-figs. 111-113 ; PI. 11, figs. 337-340 ; Map 5) 

o". 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 (PI. 11, fig. 338). 

o*. 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. 

$. 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. 6* 34-38 mm. ; $ 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 rostella, 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 <$. 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, 1 $, 6 $ ; ibid., 12-16. iv.1934, 1 <$, 1 $ ; 
ibid., 17-22.iv.1934, 1 (J, 1 $ ; Quirimbo, 75 km. E. of P. Amboim, 300 m., 7-12.V. 
1934 (Dr. K. Jordan), 1 <$, n $ ; N'Dalla Tando, 2700 ft., 21. xi. 1908 (Dr. W. J. 
Ansorge), 1 $. Congo : 1 $. 




Figs. 111-113. C. serena genitalia. 111, $ ; 112, $ ; 113, aedeagus. 



86 



D. S. FLETCHER 



Cleora oligodranes (Prout) comb. n. 

(Text-figs. 114-116 ; PI. 11, figs. 34 x -344 ; Map 5) 

Chogada oligodranes Prout, 1922 : 358. 

Neocleora oligodranes (Prout) Janse, 1932 : 271, pi. 8 : 8 ; text-fig. 100. 

o*- Vestiture : first abdominal segment white, remainder pinkish cinnamon, irrorate with 
fuscous. Wings pinkish cinnamon, irrorate and patterned with fuscous (PI. 11, fig. 341). 
Under surface of wings tilleul buff evenly, but very sparsely irrorate and patterned with fuscous 
(PI. 11, fig. 342). 




fe : -j 



116 




Figs. 114-116. C. oligodranes genitalia. 114, $ ; 115, 5* ; 116, aedeagus. 



REVISION OF AFRICAN SPECIES OF CLEORA 87 

cj. 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 
(PI. 11, fig. 343). Underside differs in the greyer ground colour of the wings and less well 
defined pattern (PI. 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. <J 32-37 mm. ; $ 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 SC2 are 
stalked ; in a further example they are coincident, and in two other examples SC2 
anastomoses with the stalk of SC3-5. 

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 quadrimaculata, 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 <$. [Orange Free State] : Thaba'nchu, i.1915 
(G. Edelstori), genitalia slide Geometridae No. 2138. 

Orange Free State : type data, 6 <$, 1 $. Bechuanaland : Upington, 14.n1. 
1950 (H. B. Kettlewell), 1 $. Cape Province : Vredendal, 23-30. vii. 1927 (G. van 
Son), 1 (J, 1 $ ; Karoo, Richmond, 16. iv. 1949 (H. B. D. Kettlewell), 1 <$ ; Ceres, 
ii.1925 (R. E. Turner), 1 $ ; Kokstad, 27.iii.1952 (C. G. C. Dickson), 1 <$, 1 $ in 
Transvaal Museum ; Pondoland, Port St. Johns, 7-13. viii. 1923, 1 <J. 

Cleora macracantha (Herbulot) comb. n. 
(Text-figs. 117-119 ; PI. 11, figs. 33°-334 > Map 5) 

Neocleora macracantha Herbulot, 1959 : 39, fig. I. 

o*. 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 



D. S. FLETCHER 




Figs. 117-119. C. macracantha genitalia. 117, $ ; 118, <J ; 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 (PI. n, fig. 330). Underside of wings white, heavily 
patterned with fuscous (PI. 11, fig. 331). 

<£. 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. 

$. Similar to male, but upperside of wings usually less densely irrorate (PI. 11, figs. 333, 334). 

$. 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, o* 3°-4° mm. ; $ 39-50 mm. 

Variation in pattern similar to that of legrasi ; among a series of 160 specimens 
are nine examples (PL 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 Chora 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 <$. Madagascar centre, Tananarive, Pare de 
Tsimbazaza, alt. 1200 m., 2.11.1952 (P. Viette), in Museum national d'Histoire 
naturelle, Paris. 

Madagascar : Diego Suarez, ii-ix. (G. Melon), 52 £, 32 $ ; Kulau (G. Melon), 
2 <J ; Tamatave et forets d'Alahakato (Edouard Perrot), 1 <$ ; Imarina, 1892 (R. P. 
Camboue), 1 <$ ; Ambinanindrano, 50 km. W of Mahonoro, iii,v,viii,x,xii (G. K. 
Kestell-C Ornish), 5 <$, 3 $ ; Tananarive, 1933, 1 <$ ; Mananjary, 1918 (G. Melou), 
12 <J, 8 $ ; Mananjara, xi.1918, 37 <J, 4 $ ; S. Madagascar, iv.1922 (Lamberton), 1 $. 



Cleora derogaria (Snellen) comb. n. 

(Text-figs. 120-122 ; PI. 12, figs. 345-348 ; Map 5) 

Boarmia derogaria Snellen, 1872 : 73, pi. 6 : 36. 
Boarmia obsitaria Mabille, 1890 : 47. Syn.n. 
Chogada acaciaria sensu Warren, 1898 : 248. 



go 



D. S. FLETCHER 




Figs. 120-122. C. derogaria genitalia. 120, $ ; 121, o" ; !22, aedeagus. 



REVISION OF AFRICAN SPECIES OF CLEORA 91 

Chogada acaciaria ab. innsitata Warren, 1898 : 248. Syn.n. 

Chogada subspnrcata Warren, 1898 : 248. 

Boarmia derogaria Snellen ; Swinhoe, 1904 : 532 [synonymy]. 

o*. First abdominal segment white ; remainder of vestiture buffy brown, irrorate with bister. 
Wings buffy brown, irrorate and patterned rather obscurely with bister (PI. 12, fig. 345). Under- 
side of wings tilleul buff densely suffused and heavily patterned with fuscous (PI. 12, fig. 346). 

o*. 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. 

$. Lighter in colour and with better defined pattern than male, ground colour of vestiture 
and upperside of wings tilleul buff (PI. 12, fig. 347). Ground colour of underside of wings white 
and much less suffused with fuscous than male (PI. 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, <J 35-40 mm. ; 9 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 $ of Boarmia derogaria Snellen (without abdomen) : 
Banana, Afrika, v. Woerden, in Rijksmuseum van Natuurlijke Historie, Leiden. 

Lectotype $ of Boarmia obsitaria Mabille. From two male syntypes I select as 
LECTOTYPEthe specimen labelled : Landana, Boarmia obsitaria Mabille, ex. Musaeo 
P. Mabille 1923. Oberthiir Coll. Brit. Mus. 1927-3, genitalia slide Geometridae No. 

2135- 

Lectotype $ of Chogada subspnrcata Warren. From a series of 5 <$ and 4 $ 
syntypes I select as LECTOTYPE the female specimen labelled : Warri, ix.1897 
(Dr. Roth), Chogada subspnrcata Warren type $, Rothschild Bequest B.M. 1939-1. 

Holotype $ of Chogada acaciaria ab. inusitata Warren : Warri, vi.1897 (Dr. Roth), 
genitalia slide Geometridae No. 2152. 

Gambia (A. Moloney), 2 <$, 3 $. Sierra Leone (C. R. Bartlett), 1 £. Ivory 
Coast : Bingerville, 1-7.vii.1915 (G. Melou), 1 <$. At Sea : Between Old Calabar 
and Accra (G. E. Bergman), 1 $. Nigeria : Warri, iv,vii-ix (Dr. Roth), 6 <$, 6 $ ; 
Degema (Ansorge), 3^, 2 $ ; Niger Coast, 1 $ ; Opobo, 1 $ ; Rio del Ray, 12.x. 1931 
(D. R. Rosevear), 1 $. N. Angola : Landana, 1 <$. 



92 D. S. FLETCHER 

Cleora pavlitzkiae (Fletcher) comb. n. 
(Text-figs. 123-135 ; PI. 12, figs. 349-362 ; Map 6) 

Neocleora pavlitzkiae Fletcher, 1958a : 139, pi. 2 : 8, 10, pi. 4 : 26-28. 

o*. 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. 

o*. 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. 

$. 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 lamella, 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 (Leopold ville), 
Mutwanga, are intermediate between subsp. lamella 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 (Leopold ville) ; Cameroun. 

Cleora pavlitzkiae pavlitzkiae (Fletcher) 

(Text-figs. 123-125 ; PI. 12, figs. 349~354 ', Map 6) 

6* $. In the 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 (PI. 12, fig. 353) ; a female from Vumba has the upper surface of the wings patterned with 
fuscous black (PI. 12, fig. 354). 

o*. 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. 




123 




Figs. 123-125. C. pavlitzkiae pavlitzkiae genitalia. 123, aedeagus ; 124,6*; 125,$. 



94 D. S. FLETCHER 

$. 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, o* 37 _ 45 mm. ; $ 41-45 mm. 

Distribution (Map 6). Rhodesia ; Malawi. 

Material examined. Holotype <J. S. Rhodesia : Vumba, 7.XL1936 (/. E. 
Drysdale), genitalia slide Geometridae No. 2 171. 

S. Rhodesia : Vumba Mts., Umtali, ii,xii, 7 <$, 1 $ and Vumba, 16.iii.1925 
(P. AS.), 1 <$, in National Museum of Rhodesia ; Vumba, ii,xi (G. van Son), 2 $, 
2 $ and Mt. Selinda, xii.1935 (G. van Son), 2 $, in Transvaal Museum ; type data, 
4 £. Malawi : Port Herald, Nr. Zambezi, vi.1926 (H. Barlow), 1 <J. 

Cleora pavlitzkiae lamella ssp. n. 

(Text-figs. 126, 127 ; PI. 12, figs. 355, 356 ; Map 6) 

(J $. Wings in most examples patterned and densely irrorate with bister to fuscous black 
(PI. 12, figs. 355, 356). 

o*. 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. 

$. Genitalia (Text-fig. 126). Similar to those of the nominate subspecies, but with a slightly 
broader posterior margin to the colliculum. 

Measurements. 0* 38-50 mm. ; $ 44-46 mm. 

Distribution (Map 6). Kenya ; Tanzania. 

Holotype $. Kenya : Nanyuki, x.1961 (R. H. Carcasson), genitalia slide 
Geometridae No. 5288. 

Paratypes : Kenya : Nairobi, v. 1905 (Jackson), 1 <$ ; Nairobi, Ngong, vii.1954 
(Fowler & Coulson), 1 <$ ; Mt. Kenya, Naro Moru, 7000 ft., 20.viii.1949 (J. A. Riley), 

1 <y ; Mt. Kenya, Meru Distr., ix.1930 (Mrs. H. Young), 1 $ ; Between Nanyuki 
and Meru (E. Barnes), 1 $ ; Nakuru, bred (A. L. H. Townsend), ix.1936, 1 <$ ; ibid., 
i.1937, 1 (J ; ibid., vii.1937, 1 $ ; ibid., ix.1937, 2 £ ; ibid., i.1938, 1 $ ; ibid., 
9. i. 1943, 1 (J ; ibid., 13. i. 1944, 1 <$ ; Mt. Elgon, i.1953 (T. H. E. Jackson), 1 <$ ; 
ibid., i.1959, 1 (J in Coryndon Museum. 

Other material. [Tanzania] Tanganyika : Morogoro, Kibuku, 1 $ ; Njombe, 
6000-7000 ft., 19. x. 1947 (W.Peters), ig ; ibid., 19.X.1951, 1^ ; Mufindi (P. Burdon), 

2 <£, 2 $ ; Marangu, 1-20.iii.1959 (Lindner), 1 $ in Zoological Museum, Stuttgart. 

Cleora pavlitzkiae etesiae ssp. n. 

(Text-figs. 128-130 ; Map 6) 

^ $. Externally similar to pavlitzkiae lamella in having the upper surface of the wings densely 
irrorate and patterned with bister. 

cT. 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. 

$. 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. ; $ 44-48 mm. 




Figs. 126-127. C. pavlitzkiae lamella genitalia. 126, ? ; 127, o*- 



96 



D. S. FLETCHER 




Figs. 128-130. C. pavlitzkiae etesiae genitalia. 128, aedeagus ; 129, o* I 130, ?. 



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 PI. 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 <$. [Ethiopia] Abyssinia : Harar, zi.vi.1939 (R. E. Ellison), genitalia 
slide Geometridae No. 5322. 





Figs. 131-132. C. pavlitzkiae oriadelpha genitalia. 131, $ ; 132, $. 



98 D. S. FLETCHER 

Paratypes ; [Ethiopia] Abyssinia : Harar, 18.iii.1939 (R. E. Ellison), 1 $ ; 
ibid., 9.VL1939, 1 $ ; ibid., ix-x.1939, 1 g, 14.X.1939, 1 $, 30.X.1938, 1 <$, 26. xi. 1938, 
1 $, 5.XU.1937, 1 S > Addis Ababa, 21.iii.1939 (T. Wikely), 1 <J ; Addis Ababa, 
20.iii.1948 (K. M. Guichard), 1 £ ; ibid., 7^.1948, 2 $. 

Cleora pavlitzkiae oriadelpha ssp. n. 

(Text-figs. 131, 132 ; PI. 12, figs. 357, 358 ; Map 6) 

$ $. Variable in pattern, but fairly densely irrorate with fuscous as in the subspecies lamella 
and etesiae (PI. 12, figs. 357, 358). 

o*. Genitalia (Text-fig. 131). Scobinate medial plate of gnathus as innominate 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 etesiae. 

$. 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. ; $ 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. 

HolotypecJ. Kenya : Kaptagat, [7867 ft.], 26.xi.1948 {Walker). 

Paratypes : Kenya : Kaptagat [7867 ft.], hi. 1948 [Walker), 1 <$ ; ibid., 3i.vii. 
1948, 1 <j> ; Mile 478 on Uganda Rly. [Molo, 8065 ft.], 14.xi.1900 (C. 5. Betton), 1 <J ; 
Nakuru, bred, 27.vii.1948, 1 $ ; ibid., 18.vii.1948, 1 £. 

Cleora pavlitzkiae saltuensis ssp. n. 

(Text-figs. 133-135 ; PI- 12, figs. 359-362 ; Map 6) 

o* $. 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 (PI. 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 
(PI. 12, figs. 361, 362). 



REVISION OF AFRICAN SPECIES OF CLEORA 



99 



o*. Genitalia (Text-figs. 133,134). Scobination of medial plate of gnathus broad and extending 
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. 




Figs. 133-135. C. pavlitzkiae saltuensis genitalia. 133, aedeagus ; 134, 0* ; 135, $. 



ioo D. S. FLETCHER 

$. 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, o* 36-46 mm. ; $ 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 hisitanica in W. Uganda. 

Distribution (Map 6). Burundi ; Rwanda ; Central Eastern Congo (Leopold- 
ville) ; S.W. Uganda, Kigezi. 

Holotype $. Burundi : Kitega, 17.viii.1g62 {Dr. M. Fontaine), in Musee Royal 
de l'Afrique Centrale. 

Paratypes : Burundi : Kitega, 9-21.iii.1962 {Dr. M. Fontaine), 2 $ ; ibid., 
17.iv.1962, 1 <$, 8.VL1962, 1 $, 13-28.vii.1962, 7 $, 2 $, 1-29.viii.1962, 10 g, 5 $, 
9-15.X.1962, 2 $, all in Musee Royal de l'Afrique Centrale. [Rwanda] : Lake Kivu, 
Rugege Forest, 7000 ft., x.1921 (T. A. Barns), 1 <$ in British Museum (Natural 
History). [Congo (Leopold ville)] Belgian Congo : Rutschuru, i.1928 (Ch.Seydel), 
1 cJ : Kivu, Terr. Lubero, Mulo, 9.X.1954 (R. P. M. J. Celis), 1 $ ; N. Lac Kivu, 
Rwankwi, iv-v.1948 (/. V. Leroy), 2<J, 1 $ ; ibid., 16-29.viii.1947, 1^,1$, 6.1X.1947, 

1 $, 19.ix.1947, 1 <$, i.xi.1947, 1 $, xii.1947, 1 $ ; Kivu, Nyamunyunga (Mulungu), 
v.1960 (/. Hecq), 1 <$, all in Musee Royal de l'Afrique Centrale ; Mt. Hoyo, 31.x. 1956 
(Ch. Seydel), 1 $. Uganda : Kigezi, Mafuga Forest, hi. 1952 (T. H. E. Jackson), 

2 $ ; Mafuga Forest, 17.vii.1961, ex Pinus patula (W. K. Brown), 1 <$ ; ibid., 
14.vii.1962, 1 $, 23.vii.1962, 1 9 I Mafuga, 9.viii.i962, ex Cupressus hisitanica 
(W. K. Brown), 1 $. 

Cleora lima sp. n. 

(Text-figs. 136, 137 ; PI. 12, figs. 363, 364 ; Map 8) 

o*. Vestiture and upper surface of wings closely similar to pavlitzkiae saltnensis . 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 . 

o*. 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. 

$. Not known. 

Measurement, q* 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^. [Congo (Leopoldville)] : Rwankwi, iv.1948 (/. V. Leroy), in Musee 
Royal de l'Afrique Centrale. 

Paratypes : Burundi : Kitega, 25.viii.1962 (Dr. M. Fontaine), 1 $ in Musee 
Royal de l'Afrique Centrale. [Rwanda] : Lake Kivu, Rugege Forest, 8000 ft., 
xii.1921 (T. A. Barns), 1 J. [Congo (Leopoldville)]: N.W. Kivu, Upper Oso River, 
4000 ft., forest with some grass, ii.1924, wet season (T. A. Barns), 1 <$. 




Figs. 136-137. C. lima genitalia. 136, $ ; 137, aedeagus. 



Cleora scobina sp. n. 

(Text-figs. 138-140 ; PI. 9, figs. 303, 304 ; PL 11, figs. 335, 336 ; Map 5) 

o*. 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 D. S. FLETCHER 

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 Mi of fore wing dilate to large spot. (PI. n, fig. 335). Under surface of wings 
white, sharply patterned with bister (PL 11, fig. 336). 

(J. Genitalia (Text-figs. 138, 139). Uncus slender with minute thorn-Ike tip ; scobinate 
medial plate of gnathus one and one-fourth times as broad as cucullus, slightly produced medio- 




Figs. 138-140. C. scobina genitalia. 138, aedeagus ; 139, $ ; 140, $. 



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, o* 37-42 mm. ; $ 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 <$. Uganda : Kampala, 22.viii.1932 (H. Hargreaves), genitalia slide 
Geometridae No. 2291. 

Paratypes : Uganda : Kampala, 7.VH.1927 (H. Hargreaves), 1 $ ; ibid., pupated 
13.V.1931, emerged 22. v. 1931, 1 $ ; ibid., 11.vii.1932, 1 <$ ; Kampala, 29.vii.1925, 

1 cJ ; Kampala, 26. xi. 1929 (G. H. E. Hopkins), i<J ; Entebbe, i.iv.1900 (Capt. H. B. 
Rattray), 1 $ ; Bwamba, ii-iii.1957 (R. Carcasson), 1 $ ; Mpanga Forest, 22. xi. i960, 
ex Cupressus lusitanica (W. K. Brown), 1 $ ; Mpanga Forest, 21.vii.1962, ex 
Eucalyptus torelliana (W '. K.Brown), 1 $. Burundi : Usumbura, 900 m., 8.VU.1961 
(Dr. M. Fontaine), 1 $, in Musee Royal de l'Afrique Centrale. [Congo (Leopoldville)] 
Belgian Congo : Ruwenzori, 18-20.V.1950 (H. B. D. Kettlewell), 3 <$, 2 $ ; Takalu, 
W. of Lake Albert, iv, 1 <$ ; E. Ituri Valley, 30 miles south of Irumu, 3000 ft., 
vii.1924 (T. A. Bams), 1 $ ; Upper Uelle distr., Dungu, iv,v,vii, 3 <$ ; Upper Uelle 
distr., Sabuni, v, 1 $, all in British Museum (Natural History) ; Kibali-Ituri, Nioka, 
10.vii.1953 (/. Hecq), 1 $ ; Haut-Uele, Moto, 1921, 2 $ ; Rutschuru, 2.L1928 (Ch. 
Seydel), 1 <$ ; N. Lac Kivu, Rwankwi, 4.ix.i947 (/. V. Leroy), 1 $ ; ibid., iv.1948, 

2 <J ; ibid., v.1948, 1 $ ; Lulua Kapanga, xii.1932 (F. G. Overlaet), 1 <§, all in Musee 
Royal de l'Afrique Centrale. 

Other material. Ivory Coast : Bingerville, 1915 (G. Melou), 1 $, 1 $ ; ibid., 
1-7.vi.1915, 1 $ ; ibid., n-14.viii.1915, 1 <J, 1 $. 



104 



D. S. FLETCHER 



Cleora radula sp. n. 

(Text-figs. 141-143 ; PI. 13, figs. 365-379 \ M &P 7) 

o*. Colours of vestiture and of upper and under surfaces of wings vary geographically. 

o*. Genitalia (Text-figs. 141, 142). Uncus with minute thorn-like tip ; scobinate medial 
plate of gnathus semicircular, diameter equal to one-half width of cucullus ; apex of juxta 
rugose ; 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. 





141 



Figs. 141-143. C. radula genitalia. 141, aedeagus ; 142, 5* ; 143, $. 



REVISION OF AFRICAN SPECIES OF CLEORA 105 

$. 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 (PI. 13, fig. 367). Underside of wings white, patterned with 
bister (PI. 13, fig. 368). 

$. 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 
copulatnx 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 radula 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 radula from scobina ; 
the female of lima is not yet known. 

Distribution (Map 7). Guinea ; Ghana ; Nigeria ; Fernando Po ; Cameroun ; 
Angola ; Congo (Leopoldville) ; Uganda ; Kenya. 

Cleora radula radula ssp. n. 

(Pi. 13, figs. 365-369 ; Ma P 7) 

c?. 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 (PI. 13, fig. 366). 

$. (PI. 13, figs. 367, 368). See under description of species. 

Measurements, o* 38-44 mm. ; $ 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 (PI. 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. 



io6 D. S. FLETCHER 

Holotype <$. [Congo (Leopoldville)] : Lusambo, 29.vi.1949 {Dr. M. Fontaine), 
in Musee Royal de l'Afrique Centrale. 

Paratypes : Gabon : Lastourville, i.1959 (P. Rougeot), 1 $, in British Museum 
(Natural History). [Congo (Leopoldville)] : Leopoldville, Binza, 29. ix. 1954 {Dr. 
M. Fontaine), 1 $ ; Stanleyville, 9.VL1948 {Dr. Faniel), 1 $ ; Sankuru, Djeka, 
17.xii.1952 {Dr. M. Fontaine), 1 $ ; Sankuru, Komi, vi.1939 (/. Ghesquiere) , 1 <$ ; 
Sankuru, Lusambo, 7. v. 1950 {Dr. M. Fontaine), 1 $ ; ibid., 14. vi. 1949, 1 £ ; 17.VL 
1949, 1 (J, 1 $ ; 20.vi.1949, 1 (J ; 2.vii.i949, i<J ; 14.vii.1950, i<J ; 18.vii.1950, i<J ; 
23.vii.1950, 1 (J, 1 ? ; 2.viii.i95o, 1 $ ; 6.viii.i95o, i<J ; 7.viii.i95o, 2<$ ; 8.viii.i95o, 
1 $ ; 9.viii.i95o, 1 <J ; 14.viii.1950, 1 $ ; 31.viii.1950, 1 $ ; ix.1950, 1 ^ ; 5,6,8, 
14. ix., each 1 <$ ; x.1950, 1 <$ ; 11. x. 1950, 1 <$ ; Sankuru, Katako-Kombe, 16. hi. 
1952 {Dr. M.Fontaine), 1$ ; ibid., 19. hi. 1952, i£ ; 6JX.1952, ij ; 15.ix.1952, 1^ ; 
8.X.1952, 1 (J ; 14. xh. 1952, 1 (J ; Uele, Paulis, 19. i. 1956 {Dr. M. Fontaine), 1 $ ; 
ibid., 2.U.1956, 1 ? ; 4JV.1956, 1 £ ; 8JV.1956, 1 ^ ; 12.iv.1956, 1 ^ ; 4.^.1956, 
icJ ; 28. xi. 1952, i(J ; i.xh.1956, 1 ^ ; ibid., 28. xh. 1955, 1 $ ; Ituri, Nduye-Makara, 
x.1921 {A. Pilette), 1 <$, all in Musee Royal de l'Afrique Centrale ; Kondolola 
District, Lindi Valley, 1600-1700 ft., v.1921 {T. A. Barns), 2 $ in British Museum 
(Natural History). 

Other material. Angola : Quicolungo, 120 km. N. of Lucala, 800 m., iv.1936 
{R. Braun), 11 <$. 



Cleora radula leptopasta ssp. n. 

(PL 13, hgs. 370, 371 ; Map 7) 

tj. 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. (PI. 13, fig. 370). Under surface of wings white, suffused and patterned 
with bister (PI. 13, fig. 371). 

Measurements, o* 38-43 mm. 

Externally closely similar to dargei with which it occurs and only reliably 
determined by reference to the genitalia. 
Distribution (Map 7). Ghana ; Nigeria. 

Holotype <$. [Nigeria] : Lagos, genitalia slide Geometridae No. 5469. 

Paratypes : [Ghana] Gold Coast : Coomassie [Kumasi] {H. Whiteside), 2 £ ; 
Nsuaem, i.1922 {N. E. Bell), 1 <$. Nigeria : Lagos, 1 <$ ; Ilesha {L. E. H. Humfrey), 
1 (J ; S. Nigeria, 1 $. 

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 $. 
Fernando Po : (W. Cooper), 2 <$, 1 $. [Cameroun] : Afriq. Occid., Station 
Kamerun, Lolodorf, 1894-1895 (L. Conradt), 1 $ ; Johann-Albrechts Hohe (L. 
Conradt), 2 (J, 1 $ ; Bitje, Ja River, 1 <$. 

Cleora radula arenosa ssp. n. 

(PI. 13, figs. 372-376 ; Map 7) 

o*. 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 (PL 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 
(PI. 13, fig. 372). 

$. 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 M 2 and 
Cui and between postmedial and subterminal fasciae (PI. 13, figs. 375, 376). 

Measurements. £ 38-41 mm. ; $ 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 (Leopold ville), Mt. Hoyo and Kivu ; W. Uganda. 

Holotype <$. Uganda : Bwamba, vi.1956 (R. Carcasson). 

Paratypes : [Congo (Leopoldville)] Belgian Congo : Mount Hoyo, 10.ii.1956 
(Ch. Seydel), 1 $ ; ibid., i.iii.1956, 1 <$ ; Ituri Forest, 3800-4000 ft., iii-iv.1930, 
beginning wet season (Lord Howard de Walden Exp.), 1 <$ ; Escarpment West 
Semliki Valley, 20 mis. S.W. of Boga, 3000-4000 ft., vii.1924, borders of tropical 
forest and long grass (T. A. Barns), 1 $ ; N. Lac Kivu, Rwankwi, iv.1948 (/. V. 
Leroy), 2 $ and 19.viii.1947, 1 $ in Musee Royal de l'Afrique Centrale ; Takulu, W. 
of Lake Albert, 1 $ ; W. Kivu, south side middle Lowa Valley, south of Walikali, 
3000 ft., forest, hi. 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 £ ; W. Kivu, Lowowo Valley, South Lowa District, 4000 ft., mountain 
forest, hi. 1924, wet season (T. A. Barns), 1 <$. Uganda : Bwamba, v. 1956 (R. 
Carcasson), 3^ ; ibid., vi.1956, 1 $ ; Kigezi District, Impenetrable Forest, Kanungu, 
4500 ft., v.1952 (/. A. Burgess), 1 <$. 



Cleora radula eumelana ssp. n. 

(Pi. 13, figs. 377-379 ; Ma P 7) 

o*. 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. (J 40-41 mm. ; $ 40 mm. 



io8 D. S. FLETCHER 

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 (PI. 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 <$. W. Kenya : [Kakamega], Malaba Forest, vi.1957 (C. R. Howard), 
genitalia slide Geometridae No. 5360. 

Paratypes : data as holotype, 1 <$, 1 $ in British Museum (Natural History), 
2 <J in Coryndon Museum. 

Cleora rothkirchi (Strand) comb. n. 
(Text-figs. 144-146 ; PI. 14, figs. 380-391 ; Map 8) 

Boarmia rothkirchi Strand, 1914 : 44. 
Boarmia acaciaria sensu Hampson, 1903 : 330. 

o* ?. 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. 

o*. 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. 

$. 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 rothkirchi Strand, 1914 : 44. 

cj. 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 







145 






146 

Figs. 144-146. C. rothkirchi genitalia. 144, aedeagus ; 145, <$ ; 146, $. 



no D. S. FLETCHER 

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 M% of forewing dilate to form spot (PL 14, fig. 380) . Under surface of wings white, 
patterned with bister (PI. 14, fig. 381). 

$. Pattern similar to that of male but less strongly defined and dark irroration usually 
sparse (PI. 14, fig. 382). Underside white, tinged with light buff on veins and margins and pat- 
terned with bister (PL 14, fig. 383). 

Measurements. 6* 33~38 mm. ; $ 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 $. Cameroun : Duala, 22.x. (v. Rothkirch), in 
Deutsches Entomologisches Institut. 

Ivory Coast : Bingerville, 1915 (G. Melon), 2 $. Ghana : N. Territorie Ketes, 
Krachi (A. W. Cardinall), 2 $. Nigeria : R. Niger, 7 miles S. of Baro, 7.U.1911 
(G. B. Simpson), 1 $ ; Bonny, 3.1.1902 (Ansorge), 1 $. Angola : Quirimbo, 75 km. 
E. of P. Amboim, 300 m., 7-12. v. 1934 (Dr. K. Jordan), 1 $, 1 $, aU in British Museum 
(Natural History). Congo (Leopoldville) : Stanleyville a Coquilhatville, xi.1921 
(L. Verlaine), 1 $ ; Yumbi [i°53"S. i6°34"E.], i.x.1929 (A. J. Bredo), 1 $ ; Bolombo 
[3°59"S. 2i°24"E.], vii.1938 (/. Ghesquiere) , 1 $ ; Rutschuru, i.1928 (Ch. Seydel), 
1 <$ ; N. Kivu, Ngesho, ix.1937 (/. Ghesquiere), 1 $ ; Lusambo, vii,viii (Dr. M. 
Fontaine), 3 ^ ; Katanga, R. Lufira, 14.x. 1925 (Ch. Seydel), 1 <$ ; Elisabeth ville, 
iii,v (Ch. Seydel), 1 <$, 2 $. Rwanda : Kisenyi, 20.iv.1957 (Dr. M. Fontaine), 1 $. 
Burundi : Usumbura, 900 m., vi,vii,viii (Dr. M. Fontaine), 3 $, all in Musee Royal 
de l'Afrique Centrale. Uganda : Kampala, 26.vii.1925, 1 $ ; Entebbe (F. J. 
Jackson), 3 $. Kenya : (G. W. Jeffery), 1 $ ; Kibwezi, v.1922 (W. Feather), 1 $. 
Suna, Kavirondo, iv.1932 (W. Feather), 1 J. Tanzania : Shinyanga, Mwandui, 
hi. 1952 (Capt. Croft), 1 $. Mozambique : Lorenzo Marques (Distant coll.), 1 <$. 
S. Rhodesia : Victoria Falls, rain forest, 22. vii. 1949 (H. B. D. Kettlewell), 1 <$. 



Cleora rothkirchi amydropa ssp. n. 

(PI. 14, figs. 388-391 ; Map 8) 

Boarmia acaciaria sensu Hampson, 1903 : 330. 

o* $. Differs from the nominate subspecies and from subsp. insularum 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, a" 34-38 mm. ; $ 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 in 

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 J. Socotra : Deneghan, 300 ft., 14. hi. 1953 (G. Popov), genitalia slide 
Geometridae No. 2194. 

Paratypes : data as holotype, 6 $, 3 $ ; Ahdo Dimellus, 3000 ft., 8-15. ii. 1899 
(W. R. 0. Grant), 1 <J, 2 $. 

Other material. Kenya : Mombasa Island, 1-7. x. 1903 (F. J. Jackson), 1 $ ; 
Mombasa, vi.1916 (van Someren), 1 $ ; Diani, 15 mis. S. of Mombasa, iv.1953 (N. 
Mitton), 1 <$, all in British Museum (Natural History) ; Kenya coast, Shimba Hills, 
xii.1961 (R. H. Carcasson), 1 $ ; Kenya coast, Gazi Forest, xii.1961 (R. H. Carcasson) 
1 $, both in Coryndon Museum. 

Cleora rothkirchi insularum ssp. n. 

(PI. 14, figs. 386, 387 ; Map 8) 

o* ?. 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 (PI. 14, figs. 386, 387). 
Measurements. $ 30-35 mm. ; $ 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. 
amy drop a. 

Distribution (Map 8). Madagascar ; ? Comoro Is. 

Holotype $. Madagascar : Diego Suarez, iii-iv (G. Melon), genitalia slide 
Geometridae No. 2181. 

Paratypes : Madagascar : Diego Suarez, 21-23. iv.1917, 1 <$, 4 $ ; ibid., 3. v. 
1917, 1 $ ; ibid., 8-24.vii.1917, 1 <$, 2 $ ; ibid., 23.viii.1917, 1 $ ; Lambomakandro, 
Sakaraha, Tulear, hi. 1935 (R. Catala), 1 $ ; Mananjara, xi.1918, 6^. 

Other material. Comoro Is. : Anjouan I., 26-30. vi.1911 (G. F. Leigh), 1 <$ ; 

ibid., iv-vii.1951 (Behagel), 1 <$ ; Grande Comore, 1884 (L. Humblot), 1 J, 1 $ ; 

Moheli, Fomboni, 10 m., ix.1958 (P. Griveaud), 1 $ ; Mayotte, Chingoni, 70 m., 
x.1958 (P. Griveaud), 1 $. 



D. S. FLETCHER 
CHECK LIST OF SPECIES INCLUDED IN CLEORA 



CLEORA, Curtis, 1825 

Cerotricha Guenee, 1857 

Bar sine Meyrick, 1883 

Meyrickia Butler, 1884 

Chogada Moore, 1887 

Carecomotis Warren, 1896 

Neocleora J arise, 1932 
cinctaria cinctaria (Denis & Schiffermiiller, 

1775) 

pascuaria (Brahm, 1791) 

consimilaria (Duponchel, 1829) 
cinctaria bowesi Richardson, 1952 
cinctaria insolita (Butler, 1878) 
sublunaria (Guenee, 1857) 

transfixaria (Walker, i860) 

atrolinearia Hulst, 1888 

areataria Broadwell, 1907 
projecta (Walker, i860) 

■Manitoba (Grossbeck, 191 1) syn. n. 
nigrofasciaria (Leech, 1897) 
leucophaea (Butler, 1878) 

elegans (Oberthur, 1884) 

pagina (Wileman, 191 1) 
venustaria (Leech, 1891) 
alienaria alienaria (Walker, i860) 
alienaria gelidaria (Walker, 1863) 
alienaria rasanaria (Swinhoe, 1915) 
alienaria fumipennis Prout, 1929 
fratema (Moore, 1888) 
determinata (Walker, i860) 
yakushimana (Inoue, 1956) 
minutaria (Leech, 1891) 
hermaea Prout, 1929 
subbarbaria Prout, 1929 
sevocata Prout, 1929 
hospita (Prout, 1916) 
illustraria illustraria (Walker, 1863) 
illustraria anestiaria (Swinhoe, 1915) 
illustraria crina Prout, 1929 
illustraria aequivoca Prout, 1929 
apista Prout, 1929 
meceoscia Prout, 1929 
mecistoscia Prout, 1929 
paepalima West, 1932 
tella (West, 1929) 
pendleburyi Prout, 1929 
xanthorrhages Prout, 1929 
mjoebergi Prout, 1926 
neomenia Prout, 1932 
repetita (Butler, 1882) 

inflexaria (Snellen, 1881) 



epistictis (Meyrick, 1889) 
proletaria (Swinhoe, 1915) 

lipotera West, 1920 

processaria (Walker, i860) 

nigronotaria (Wileman, 191 1) comb. n. 

inject aria injectaria (Walker, i860) 
sublectaria (Walker, 1863) 
compactaria (Walker, 1863) 

injectaria fuliginosa (Warren, 1894) 

injectaria vittata (Warren, 1899) 

injectaria dobboensis Prout, 1929 

injectaria anidryta Prout, 1929 

idiocrossa Turner, 19 18 

concentraria concentraria (Snellen, 1877) 
? invalidaria (Snellen, 1895) 

concentraria praia Prout, 1928 

concentraria inobeda Prout, 1929 

godeffroyi (Butler, 1886) 

psectra Fletcher, 1957 

samoana (Butler, 1886) 

collenettei Prout, 1929 

esoterica Prout, 1929 

f. pusillanimis Prout, 1935 

myrmidonaria (Guenee, 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 

licornaria Guenee, 1857 

dodonaeae Prout, 1929 

cheesmanae Prout, 1929 

nausori (B. -Baker, 1905) 

perstricta Prout, 1934 

scriptaria (Walker, i860) 

panagrata (Walker, 1862) 
stigmaticata (Walker, 1862) 
menanaria (Walker, 1863) 
antipodaria (Felder, 1874) 
arenacea (Butler, 1879) 
desiccata (Butler, 1879) 

perlepidaria (Warren, 1900) 

compectinata (Warren, 1906) 

displicata (Walker, i860) 

pheucta Prout, 1937 

goldfinchi Prout, 1937 

diphasia diphasia Prout, 1937 

diphasia refota Prout, 1937 



REVISION OF AFRICAN SPECIES OF CLEORA 



113 



rhadia rhadia Prout, 1929 
rhadia frigescens Prout, 1937 
perbona Prout, 1937 
lacteata (Warren, 1897) 
decisaria (Walker, 1866) 
amphidoxa Prout, 1937 
callicrossa (Meyrick, 1889) 
nigristigma nigristigma Prout, 1937 
nigristigma talaseensis Prout, 1937 
hemiopa hemiopa Prout, 1928 
hemiopa quirosi Prout, 1929 
hemiopa ecdees Prout, 1929 
ictaibasis Prout, 1937 
ciicullata 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 cultrata (Fletcher, 1953) 
onycha serrata (Fletcher, 1953) 
onycha limitata (Fletcher, 1953) 
onycha acuata (Fletcher, 1953) 
taprobana (Fletcher, 1953) 
kalisi (Fletcher, 1953) 
biclavata (Fletcher, 1953) 
inornata (Fletcher, 1953) 
fasciata (Fletcher, 1953) 
olivata (Fletcher, 1953) 
cornaria (Guenee, 1857) 

invectaria (Walker, i860) 

proper ata (Walker, i860) 
repulsaria (Walker, i860) 
propulsaria propulsaria (Walker, i860) 
propulsaria fieldi (Fletcher, 1953) 
olivomaculata (Fletcher, 1953) 
rostrata rostrata (Fletcher, 1953) 
rostrata moniliata (Fletcher, 1953) 
falculata (Fletcher, 1953) 
tenebrata tenebrata (Fletcher, 1953) 
tenbrata acntiorata (Fletcher, 1953) 
tenebrata arcnata (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 ftaviorata (Fletcher, 1953) 
costiplaga f err ata (Fletcher, 1953) 
costiplaga eichhorni (Fletcher, 1953) 
costiplaga umbrata (Fletcher, 1953) 
pupillata pnpillata (Walker, i860) 
pupillata fitliginala (Fletcher, 1953) 
pupillata luzonensis (Fletcher, 1953) 
buxtoni (Fletcher, 1953) 
perfumosa extendata (Fletcher, 1953) 
perfumosa perfumosa (Warren, 1896) 
braeckeli Debauche, 1941 
forfictilata (Fletcher, 1953) 
levata (Fletcher, 1953) 

albobrunneata albobrnnneata (Fletcher, 1953) 
albobrunneata cordata (Fletcher, 1953) 
sabulata sabulala (Fletcher, 1953) 
sabulata inconspicnata (Fletcher, 1953) 
contiguata contiguata (Moore, 1867) 
contiguata brooksi (Fletcher, 1953) 
contiguata imparata (Fletcher, 1953) 
contiguata bigladiata (Fletcher, 1953) 
acaciaria (Boisduval, 1833) 
Imiisversaria (Pagenstecher, 1907) 
belularia (Warren, 1897) 

fnnesta (Warren, 1905) 
jhtrivenala sp. n. 
melanochorda (Fletcher, 1958) 
papillifer Prout, 1934 
cancer sp. n. 
tamsi sp. n. 

viettei (Herbulot, 1958) 
oculata sp. n. 
prosema Prout, 1927 
anacantha sp. n. 
epiclithra sp. n. 
carcassoni sp. n. 

herbuloti herbuloti (Fletcher, 1958) 
herbuloti phaea ssp. n. 
acerata sp. n. 

subcincta subcincta (Warren, 1901) 
subcincta longifibulata (Fletcher, 1958) 
lacrymata sp. n. 
echinodes sp. n. 
raphis sp. n. 
aculeata sp. n. 
panarista sp. n. 
quadrimaculata (Janse, 1932) 
boetschi (Herbulot, 1961) 
bicornis sp. n. 
dargei (Herbulot, 1961) 
dactylata sp. n. 



ii4 



D. S. FLETCHER 



thy vis sp. n. 

nigrisparsalis (Janse, 1932) 

tulbaghata (Felder, 1875) 

plax sp. n. 

munda (Warren, 1899) 

lamottei (Herbulot, 1954) 

toulgoetae (Herbulot, 1961) 

rostella sp. n. 

legrasi (Herbulot, 1955) 

angustivalvis (Herbulot, 1965) 

serena sp. n. 

oligodranes (Prout, 1922) 

macracantha (Herbulot, 1959) 

derogaria (Snellen, 1872) 

obsitaria (Mabille, 1890) 



subspurcata (Warren, 1898) 
pavlitzkiae pavlitzkiae (Fletcher, 1958) 
pavlitzkiae lamella ssp. n. 
pavlitzkiae etesiae ssp. n. 
pavlitzkiae oriadelpka ssp. n. 
pavlitzkiae saltuensis ssp. n. 
lima sp. n. 
scobina sp. n. 
radula radula sp. n. 
radula leptopasta ssp. n. 
radula arenosa ssp. n. 
radula eumelana ssp. n. 
rothkirchi rothkirchi (Strand, 1914) 
rothkirchi amydropa ssp. n. 
rothkirchi insularum ssp. n. 



SPECIES TO BE MOVED FROM CLEORA WHEN GENERA ARE AVAILABLE 



Cleora albitrigonis Prout, 1927, Trans, ent. Soc. Lond. 75 : 195, pi. 20 : 13. 

Cleora argicerauna Prout, 1929, Treubia 7 : 448, pi. 9 : 14. 

Cleora atriclava Prout, 1926, Novit. zool. 33 : 183. 

Cleora bathyscia Turner, 1917, Proc. Linn. Soc. N.S.W. 42 : 371. 

Cleora cnephaea Prout, 1915, Novit. zool. 22 : 359. 

Cleora cryptogonia Prout, 1927, /. Bombay nat. Hist. Soc. 31 : 941, pi. 2 : 7. 

Cleora discipuncta Joicey & Talbot, 1917, Ann. Mag. nat. Hist. (8) 20 : 74, pi. 4 : 8. 

Cleora euboliaria (Walker), Turner, 1917, Proc. Linn. Soc. N.S.W. 42 : 372. 

Tephrosia? euboliaria Walker, i860, List Lep. Ins. B.M. 21 : 419. 
Cleora euplates Prout, 1925, Trans, ent. Soc. Lond. 1925 :31s, pi. 36 : 26. 
Cleora expleta Prout, 1932, Jl. fed. Malay St. Mus. 17 : 105, pi. 11 : 25. 
Cleora fenestrata Prout, 1916, Novit. zool. 23 : 52. 
Cleora 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, pi. 4 : 16. 

Alcis cinnamomea Rothschild, 1915, Lepidoptera Brit. Ornith. Union Exped. S. Dutch New 
Guinea, 86. 
Cleora gypsochroa Turner, 1947, Proc. R. Soc. Qd 58 : 91. 
Cleora hoplogaster Prout, 1916, Novit. zool. 23 : 51. 

Cleora inaequipicta Prout, 1921, Bull. Hill Mus. Witley 1 : 150, pi. 18 : 14. 
Cleora incompletaria (Guenee), Vinson, 1938, Bull. Maurit. Inst. 1(4) : 38. 

Boarmia incompletaria Guenee, 1862, in Maillard, Notes sur Vile de la Reunion, Annexe G, 27. 
Cleora nesiotis Turner, 1926, Pap. Proc. R. Soc. Tasm. 1925 : 99. 
Cleora orygaria (Guenee), Vinson, 1938, Bull. Maurit. Inst. 1(4) : 38. 

Neocleora orygaria brunneata (Warren) Herbulot, 1956, Naturaliste malgache 8 : 252. 

Xylopteryx brunneata Warren, 1912, Novit. zool. 9 : 523. 
Cleora polymiges Prout, 1926, Novit. zool., 33 : 22. 
Cleora proemia Prout, 1917, Novit. zool. 24 : 435. 
Neocleora refulgens Herbulot, 1965, Bull. Soc. ent. Fr. 69 : 258. 

Boarmia (Cleora) rhamphoides Wehrli, 1943, in Seitz, Macrolepidoptera of the World 4, Suppl., 494. 
Cleora russoi Prout, 1932, Rev. Zool. Bot. afr. 21 : 244. 

Cleora scripta Joicey & Talbot, 1917, Ann. Mag. nat. Hist. (8) 20 : 74, pi. 3 : 7. 
Cleora semidiscata (Warren) Warren, 1907, Novit. zool. 14 : 172. 

Chogada semidiscata Warren, 1906, Novit. zool. 13 : 141. 



REVISION OF AFRICAN SPECIES OF CLEORA 115 

Chora subnigrata (Warren) Prout, 1927, /. Bombay nat. Hist. Soc. 31 : 942, pi. 2 : 7. 

Scotopteryx? subnigrata Warren, 1901, Novit. zool. 8 : 34. 
Chora tora Prout, 1926, Novit. zool. 33 : 183. 
Chora trigrapta Prout, 1927, /. Bombay nat. Hist. Soc. 31 : 940. 
Chora trisinuata Warren, 1898, Novit. zool. 5 : 248. 
Chora vatia Prout, 1927, /. Bombay nat. Hist. Soc. 31 : 941. 

SPECIES MOVED FROM CLEORA TO OTHER GENERA 

Chora aargostigma Prout, 1927, /. Bombay nat. Hist. Soc. 31 : 939 moved to A his. Comb. n. 
Chora aeglophanes Prout, 1926, Sarawak Mus. J . 3(2) : 203 ; 1928, op. cit., pi. 16 : 9, moved to 

Aids. Comb. n. 
Cleta albipunctaria Schaus, 1901, Trans. Am. ent. Soc. 27 : 241, treated by Prout as Chora, 

moved to Stellidia in the Noctuidae. Comb. n. 
Chora amictozona Prout, 1932, Novit. zool. 38 : 114 moved to Alcis. Comb. n. 
Chora antelmaria (Mabille) ; Vinson, 1938, Bull. Maurit. Inst. 1(4) : 38. 
Boarmia antelmaria Mabille, 1893, Ann. Soc. ent. Belg. 37 : 64. 

Chogada martnorata Warren, 1897, Novit. zool. 4 : 247 moved to Ascotis. Comb. n. 
Chora athola Prout, 1926, Novit. zool. 33 : 183 moved to Alcis. Comb. n. 
Chora bianqnita Schaus, 1901, Trans. Am. ent. Soc. 27 : 182 moved to Vinemina. Comb. n. 
Chora chionospila Turner, 1947, Proc. R. Soc. Qd 58 : 90 moved to Parathetnis and placed in the 

synonymy of Tephrosia externaria Walker, 1866. Syn. n. 
Chora cockaynei Prout, 1916, Novit. zool. 23 : 53 moved to Alcis. Comb. n. 
Chora colorifera Prout, 1916, Novit. zool. 23 : 53 moved to Alcis. Comb. n. 
Chora coniozona Prout, 1927, /. Bombay nat. Hist. Soc. 31 : 940, pi. 2 : 3 moved to A his. Comb. 

n. 
Chora decussata Moore, 1867, Proc. zool. Soc. Lond. 1867 : 628, pi. 33 : 4 moved to Alcis. Comb. 

n. 
Chora derivata Prout, 1926, Sarawak Mus. J. 3(2) : 203 moved to Alcis. Comb. n. 
Chora dolichoptila Turner, 1947, Proc. R. Soc. Qd 58 : 91 moved to Chhnias. Comb. n. 
Boarmia (Chora) flavolinearia (Leech) ; Wehrli, 1954, * M Seitz, Macrohpidoptera of the World 4, 
Suppl., 720. 

Boarmia flavolinearia Leech, 1891, Entomologist 24, Suppl., 47 moved to Alcis. Comb. n. 
Boarmia (Chora) fortunata (Blachier) ; Wehrli, 1943, in Seitz, Macrohpidoptera of the World 4, 
Suppl., 494. 

Boarmia fortunata Blachier, 1887, Feuille jeun. Nat. 17 : 103. 

Boarmia obscura B. -Baker, 1891, Trans, ent. Soc. Lond. 1891 : 218. 

Boarmia wollastoni B. -Baker, 1891, Trans, ent. Soc. Lond. 1891 : 217 moved to Ascotis. 

Comb. n. 
Boarmia buechlei Kilian, 1897, Societas ent. 12 : 41. 
Chora glaucotoxa Prout, 1927, Trans, ent. Soc. Lond. 75 : 195, pi. 20 : 12 moved to Ascotis. 

Comb. n. 
Chora godmani Druce, 1892, Biologia cent. -am., Zool., Lep. Het. 2 : 72, pi. 48 : 9 moved to 

Semiothisa. Comb. n. 
Chora gracilis Warren, 1904, Novit. zool. 11 : 107 moved to Iridopsis. Comb. n. 
Chora hemichroma Turner, 1947, Proc. R. Soc. Qd 58 : 89 moved to Chhnias. Comb. n. 
Chora hemiphanes Prout, 1925, Novit. zool. 32 : 56 moved to Alcis. Comb. n. 
Chora hypopoecilia Prout, 1928, Bull. Hill Mus. Withy 2 : 151 moved to Alcis. Comb. n. 
Chora imbecilis (Moore) ; Prout, 1927, /. Bombay nat. Hist. Soc. 31 : 939. 

Pseudocoremia imbecilis Moore, 1888, Descr. New Indian Lepidopterous Insects in coll. 
Atkinson, 241 moved to Alcis. Comb. n. 
Chora irrita Prout, 1928, Bull. Hill Mus. Withy 2 : 152 moved to Alcis. Comb. n. 
Chora irrita f. (?sp.) obruta Prout, 1928, loc. cit. 



n6 D. S. FLETCHER 

Chora latifasciata (Warren) ; Prout, 1927, /. Bombay nat. Hist. Soc. 31 : 938. 

Poecilalcis latifasciata Warren, 1893, Proc. zool. Soc. Lond. 1893 : 427. 

Boarmia enryzona Hampson, 1895, Fauna Br. India, Moths 3 : 281. 

Boarmia eurydiscaria Hampson, 1902, /. Bombay nat. Hist. Soc. 14 : 507 moved to Aids. 
Comb. n. 
Chora megaspilaria Moore, 1867, Proc. zool. Soc. Lond. 1867 : 629 moved to Alcis. Comb. n. 
Chora nigridorsaria Guenee, 1857, Histoire naturelle des Insectes, Species general des Lepidopteres 

9 : 232 moved to Alcis. Comb. n. 
Chora nigriscripta (Warren) ; Prout, 1929, Bull. Hill Mus. Withy 3 : 38, 51. 

Poecilalcis nigriscripta Warren, 1903, Novit. zool. 10 : 401. 

Chora nigriscripta gavisa Prout, 1929, Bull. Hill Mus. Withy 3:38. 

Chora nigriscripta phnimedia Prout, 1929, Bull. Hill Mus. Withy 3 : 51 moved to Alcis. 
Comb. n. 
Chora pachydesma Turner, 1947, Proc. roy. Soc. Qd 58 : 90 moved to Symmetroctena. Comb. n. 
Chora periphracta Prout, 1926, Sarawak Mus. J. 3(2) : 202 moved to Alcis. Comb. n. 
Chora phaeocala Turner, 1947, Proc. R. Soc. Qd 58 : 91 moved to Chlenias. Comb. n. 
Chora praecisa Turner, 191 7, Proc. Linn. Soc. N.S.W. 42 : 372 moved to Syneora. Comb. n. 
Chora praevariegata Prout, 1926, Sarawak Mus. J. 3(2) : 202; 1928, 3(3), pi. 16 : 8 moved to 

Alcis. Comb. n. 
Chora rufomarginata Moore, 1867, Proc. zool. Soc. Lond. 1867 : 628 moved to Alcis. Comb. n. 
Boarmia (Chora) scortea (Bastelberger) ; Wehrli, 1943, in Seitz, Macrohpidoptera of the World 4, 
Suppl., 494. 

Boarmia scortea Bastelberger, 1909, Ent. Z. Frank. a. M. 23 : 33 moved to Alcis. Comb. n. 
Chora semiochrea Prout, 1917, Novit. zool. 24 : 315 moved to Alcis. Comb. n. 
Chora semipullata Prout, 1925, Novit. zool. 32 : 57 moved to Alcis. Comb. n. 
Chora sericea (Warren); Prout, 1927, /. Bombay nat. Hist. Soc. 31 : 942. 

Apophyga sericea Warren, 1893, Proc. zool. Soc. Lond. 1893 : 418 removed to Apophyga. 
Chora terebravia (Guenee) Vinson, 1938, Bull. Maurit. Inst. 1(4) : 38. 

Hypopalpis terebraria Guenee, 1862, in Maillard, Notes sur Vile de la RSunion, Annexe G, 29. 

Boarmia perf or aria Guenee, 1862, op. cit., p. 30, pi. 23 : 4. 

Boarmia rousseli Oberthiir, Etudes Lip. comparies 7 : 275, pi. 169 : 1652, 1653 moved to 
Ascotis. Comb. n. 
Chora variegata (Moore) ; Prout, 1927, /. Bombay nat. Hist. Soc. 31 : 938. 

Pseudocoremia variegata Moore, 1888, Descr. New Indian Lepidopterous Insects in coll. 
Atkinson, 240. 

Chora nebulosa Swinhoe, 1891, Trans, ent. Soc. Lond. 1891 : 488. 

Chora variegata convariata Prout, 1935, Novit. zool. 39 : 231 moved to Alcis. Comb. n. 
Chora versicolor Prout, 1915, Sarawak Mus. J. 2(2) : 181 moved to Alcis. Comb. n. 
Chora venustularia Walker, 1866, List Lep. Ins. BM. 35 : 1579 moved to Alcis. Comb. n. 

REFERENCES 

Anon. 1963. Notes and Comments. Jl S. Afr. For. Ass. 44 : 1-6, 32-33. (Editor J. R. 
Sime). 

Boisduval, J. A. 1833. Descriptions des Lepidopteres de Madagascar (title omitted from 
beginning of paper, but included in index). Nouv. Ann. Mus. Hist. nat. Paris. 2(2) : 149— 
268, 8 pis. ; 2 (3): 269-270. 

1833a. Faune ent. Madagascar, Bourbon et Maurice, Lepidopteres. 122 pp., 16 pis. 

Paris. 

Butler, A. G. 1883. On a small series of Lepidoptera from the Hawaiian Islands. Ento- 
mologist's mon. Mag. 19 : 176-180. 

1884. On the duplication of generic names employed by Walker. Entomologist's mon. 

Mag. 21 : 133. 



REVISION OF AFRICAN SPECIES OF CLEORA 117 

Curtis, J. 1825. British Entomology, 2, pis. 51-98. London. 

1826. British Entomology, 3, pis. 99-146. London. 

Felder, R. & Rogenhofer, A. F. 1 874-75. Reise der Osterreichischen Fregatte Novara um 

die Erde in 1857-59, unter den Befehln des Commodore B. von W idler storf-Urbair. Zoo- 

logischer Teil. Band. 2. Abteilung 2 (Heterocera), 20 pp. 46 pis. (75-120), 1874 ; 20 pis. 

(121-140), 1875. Wien. 
Fletcher, D. S. 1953. A revision of the genus Carecomotis (Lep. Geometridae) . Ann. Mag. 

nat. Hist. (12) 6 : 100-142, pis. 3-6. 

1958. Rnwenzori Expedition 1952, 1 (6) (Geometridae) : 77-176, 32 pis. London. 

1958a. Geometridae from Tanganyika collected by Dr. Christa Lindemann and Nina 

Pavlitzki. Veroff. zool. StSamml., Miinch. 5 : 1 17-144, 5 pis. 

Guenee, A. 1857. Histoire naturelle des Insectes, Species general des Lipidopteres 9. i-lvi -f- 

514 pp. Paris. 
1862. In Maillard, L., Notes stir Vile de la Reunion (Bourbon), Annex. G. 1-72 pp., 

pis. 22-23. Paris. 
Hampson, G. 1903. In Forbes, H. O., 1903, The Natural History of Socotra and Abd-el- 

Kuri. xlvii + 598 pp., 27 pis. London. 
Herbulot, C. 1954. La Reserve Naturelle Integrate du Mont Nimba, Fascicule II, Lepi- 

dopteres Geometrides. Mim. Inst. fr. Afr. noire 40 : 301-333, 1 pi. 
1955- Deux nouveaux Neocleora malgaches (Lep. Geometridae). Bull. Soc. ent. Fr. 

60 : 38-40, figs. 
1957- L6pidopteres Geometridae de l'lle de la Reunion. Mim. Inst, scient. Madagascar 

(E) 8 : 227-236, 1 pi. 

1958. Mission du Museum dans les iles du golfe de Guinee. Entomologie : VIII Lepi- 

doptera Geometridae. Bull. Soc. ent. Fr. 63 : 100-109, figs. 

1961. Trois nouveaux Neocleora de Guinee et du Cameroun (Lepidoptera Geometridae). 

Bull. Inst. fr. Afr. noire (A) 23 : 493-495, figs. 

1965. Nouveaux Geometridae de Madagascar et des Comores. Bull. mens. Soc. linn. 

Lyon 34 (4) : 119-121. 
Hubner, J. i8i6-(i827). Verzeichniss bekannter Schmettlinge (sic). 431 pp. Anzeiger 

72 pp. Augsburg. 
Janse, A. J. T. 1932. The Moths of South Africa. Vol. 1, Sematuridae and Geometridae. 

xi + 376 pp., 15 pis., 130 figs. Durban. 
Kettlewell, H. B. D. 1957. Industrial melanism in moths and its contribution to our know- 
ledge of evolution. Proc. R. Instn Gt Br. 36, No. 164. 14 pp., 6 pis. 
McDunnough, J. Ff. 1920. Studies in North American Cleorini (Geometridae). Bull. Dep. 

Agric. Can. ent. Brch 18. 64 pp., 11 pis. 
Meyrick, E. 1883. Monograph of New Zealand Geometrina. N.Z. Jl Sci. 1 : 526-531. 

1917. Revision of New Zealand Notodontina. Trans. N.Z. Inst. 49 : 248-273. 

Moore, F. 1884-87. The Lepidoptera of Ceylon. 3. 578 pp. London. 

Pagenstecher, A. 1907. In Voeltzkow, A. Reise in Ostafrika in den Jahren 1903-1905 

2 : 93-146, 1 pi. Stuttgart. 
Prout, L. B. 1922. New and little-known Geometridae. Novit. zool. 29 : 327-363. 
1927. List of the Geometridae (Lep. Ffet.) known to occur in the Island of Sao Thome, 

with descriptions of some new species collected by Mr. T. A. Barns. Trans, ent. Soc. 

Lond. 75 : 187-200, 1 pi. 

1928. Insects of Samoa 3 (3) : 11 7-168, pi. 5. London. 

1929. Two new Polynesian species of Cleora (Lepidoptera Geometridae). Entomologist 

62 : 255-256. 
1929a. A revision of the Indo-Australian Cleora of the alienaria group. Bull. Hill. 

Mus. Witley 3 : 179-222, pis. 5-7. 

1929b. Geometridae of the " St. George " Expedition from French Oceania. Trans, ent. 

Soc. Lond. 77 : 265-277. 



n8 



D. S. FLETCHER 



Prout, L. B. 1929c. On the Geometric! genus Catoria Moore. Novit. zool. 35 : 132-141. 

1929-38. In Seitz, A., The M acrolepidoptera of the World 16 : 1-160, pis. 1-18. Stutt- 
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1934- Geometridae from the Marquesas Islands. Pubis Pacif. ent. Surv. 7 (13) : 171-177. 

1934- New Congo Geometridae. Revue Zool. Bot. afr., 26 : 82-97. 

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Strand, E. 1914. Neue Lepidoptera aus Kamerun gesammelt von Herrn Leutnant v. 
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Swinhoe, C. 1904. On the Geometridae of tropical Africa in the National Collection. Trans, 
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Hist. nat. Paris. (2) 26 : 506-509. 

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Inst. 1 (4) : 1-69. 

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from the Papuan region, collected by Mr. Albert S. Meek. Novit. zool. 3 : 272-306. 

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Museum. Novit. zool. 4 : 12-130. 

1898. New Species and Genera of the Families Drepanulidae, Thyrididae, Uraniidae, 

Epiplemidae and Geometridae from the Old-World Regions. Novit. zool. 5 : 221-258. 

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midae and Geometridae from the Old-World Regions. Novit. zool. 6 : 1-66. 

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Novit. zool. 8 : 6-20. 

— — 1904. New Drepanulidae, Thyrididae, Uraniidae and Geometridae from the Aethiopian 
Region. Novit. zool. 11 : 461-482. 

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Wehrli, E. 1939-54. In Seitz, A., Die Gross-Schmetterlinge der Erde 4, Suppl. 254-766, 
pis. 19-53. Stuttgart. 

Zimmerman, E. C. 1958. Insects of Hawaii 7. 542 pp., 423 figs. Honolulu. 



INDEX 



Index to nominal genera and to Ethiopian species and subspecies of Cleora (synonyms in italics) 



acaciaria Boisduval, 16 
acerata sp. n., 41 
aculeata sp. n., 49 
Alcis Curtis, 7 
amydropa ssp. n., no 
anacantha sp. n., 34 
angustivalvis Herbulot, 82 
arenosa ssp. n., 107 
Ascotis Hiibner, 7 



cancer sp. n., 28 
carcassoni sp. n., 36 
Carecomotis Warren, 5, 6, 7 
Catoria Moore, 8 
Cerotricha Guenee, 4 
Chogada Moore, 5 
Cleora Curtis, 4 
Colocleora Prout, 7 
Cusiala Moore, 8 



Barsine Meyrick, 5 
betularia Warren, 20 
bicornis sp. n., 57 
boetschi Herbulot, 54 



dactylata sp. n., 60 
dargei Herbulot, 58 
derogaria Snellen, 89 



INDEX 



119 



echinodes sp. n., 46 
epiclithra sp. n., 36 
etesiae ssp. n., 94 
eumelana ssp. n., 107 



flavivenata sp. n., 22 
funesla Warren, 20 



herbuloti Fletcher, 38 
Hypopalpis Guenee, 7 



Ophthalmodes Guenee, 
oriadelpha ssp. n., 98 



panarista sp. n.,51 
papillifer Prout, 25 
Paradromulia Warren, 8 
pavlitzkiae Fletcher, 92 
phaea ssp. n., 40 
plax sp. n., 71 
prosema Prout, 33 
Pseudalcis Warren, 8 



insularum ssp. n., 111 



quadrimaculata Janse, 53 



lacrymata sp. n., 44 
lamella ssp. n., 94 
lamottei Herbulot, 73 
legrasi Herbulot, 80 
leptopasta ssp. n., 106 
lima sp. n., 100 
longifibulata Fletcher, 43 



macracantha Herbulot, 87 
melanochorda Fletcher, 24 
Meyrickia Butler, 5 
munda Warren, 71 



radula sp. n., 104 
raphis sp. n., 48 
rostella sp. n., 77 
rothkirchi Strand, 108 



saltuensis ssp. n., 98 
scobina sp. n., 101 
Scotorythra Butler, 7 
serena sp. n., 84 
subcincta Warren, 41 
subspurcata Warren, 91 



Neocleora Janse, 5, 6, 7 
nigrisparsalis Janse, 64 



obsitaria Mabille, 89 
oculata sp. n., 31 
oligodranes Prout, 86 



tamsi sp. n., 28 
thyris sp. n., 62 
toulgoetae Herbulot, 76 
transversaria Pagenstecher, it 
tulbaghata Felder, 67 



viettei Herbulot, 30 





PLATE 


1 






Cleora 




Fig 








147 


acaciaria $ 


uppersid 


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148 


s 


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149 


9 


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150 


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151 


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152 


transversari 


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153 




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154 




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155 




$ 


B.M.Neg.34878 


156 




$ 


B.M.Neg.34869 


157 




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158 




$ 


B.M.Neg.34874 


159 




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160 




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161 




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B.M.Neg.34872 


162 




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B.M.Neg.34867 


163 




9 


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164 




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165 




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Hull Br. Mus. not. Hist Entomology, Suppt. 8 



PLATE i 





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57 




58 




59 






160 



161 



62 










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165 



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Chora 
Fig. 

166. betularia $ upperside B.M.Xeg. 35238 

167. g underside B.M.Neg. 35236 

168. $ upperside B.M.Neg. 35240 

169. $ underside B.M.Neg. 35239 

170. $ upperside B.M.Neg. 35241 

171. 3 underside B.M.Neg. 35237 

172. <J upperside B.M.Neg. 35229 

173. $ underside B.M.Neg. 35232 

174. (J upperside B.M.Neg. 35231 

175. $ underside B.M.Neg. 35227 



Bull. Br. Mus. nat. Hist. Entomology, Suppt. 8 



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PLATE 14 



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Map i. Distribution of Ethiopian Cleora, species-group acaciaria 




o anacantha 
▼ carcassoni 
• epiclithra 
a oculata 
■ prosema 



Map 2. Distribution of Ethiopian Chora, species-group oculata 




Map 3. Distribution of Ethiopian Chora, species-group r aphis 




Map 4. Distribution of Ethiopian Cleora, species-group tulbaghaia 




♦ angustivalvis 
■ derogaria 

v lamottei 
d legrasi 
" a macracantha 
^oligodranes 
» rostella 

• scobina 
© serena 

o toulgoetae 



Map 5. Distribution of Ethiopian Cleora, species-group rostella 




Map 6. Distribution of Ethiopian Cleova, species-group rostella, species pavlitzkiae 




Map 7. Distribution of Ethiopian Cleora, species-group rostella, species radula 



' 




■ rothkirchi rothkirchi 
o rothkirchi insularum 
e rothkirchi ? insularum 
• rothkirchi amydropa 
©rothkirchi ? amydropa 
' a lima 



Map 8. Distribution of Ethiopian Cleora, species-group rostella, species rothkirchi and lima 




d acerata 

* herbuloti herbuloti 
a herbuloti phaea 

• subcincta subcincta 
o subcincta longifibulata 
▼ tamsi 
v viettei 
■ bicornis 



Map 9. Distribution of arbitrarily placed species of Ethiopian Chora 




**M *<& 



A LIST OF SUPPLEMENTS 
TO THE ENTOMOLOGICAL SERIES 
OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



1. 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. 

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

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

4. 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. 

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

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129 : 328 Text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera : Coccoidea). Pp. 168 : 43 Text-figures. January, 
1967- £3 3*- 



2mk^ Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 

P.