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THE TAXONOMY OF THE

DREPANINAE REPRESENTED IN

CHINA, WITH AN ACCOUNT OF

THEIR WORLD DISTRIBUTION

(LEPIDOPTERA : DREPANIDAE)

<&■

A. WATSON

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Supplement 12

LONDON: 1968

THE TAXONOMY OF THE DREPANINAE

REPRESENTED IN CHINA, WITH AN

ACCOUNT OF THEIR WORLD DISTRIBUTION

(LEPIDOPTERA : DREPANIDAE)

BY

A. WATSON

14 Plates, 293 Text-figs.

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ENTOMOLOGY Supplement 12

LONDON: 1968

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. 12 of the Entomo-
logical series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

World List abbreviation :
Bull. Br. Mus. nat. Hist. (Ent.)

Trustees of the British Museum (Natural History) ic

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 13 November, 1968 Price £5

THE TAXONOMY OF THE DREPANINAE

REPRESENTED IN CHINA, WITH AN

ACCOUNT OF THEIR WORLD DISTRIBUTION

(LEPIDOPTERA : DREPANIDAE)

By A. WATSON

CONTENTS

Synopsis
Introduction
Distribution .
Key to Genera
Main Taxonomic Part
References .
Index

Page

3
3
6

*4

16

145
149

SYNOPSIS

A review is given of the taxonomy and distribution of the 76 species of Chinese Drepaninae,
and the taxonomic status of a further 8 species is examined. The genera Agnidra, Albara,
Betalbara, Callicilix, Didymana, Nordstroemia and Pseudalbara are revised. A new genus,
Par albara, 14 new species and 10 new subspecies are described. 30 names are newly placed
in synonymy, three names are extracted from synonymy, and 31 changes in the combination
of species-group and generic names are made. A key to the genera of Drepaninae found in
China is given. The world distribution of the genera, species and subspecies of Drepaninae
represented in China is discussed. The species distribution is compared with that of the
Oretinae, the other subfamily of Drepanidae present in China. For purposes of this paper
China, Formosa and Tibet are included in the term ' China '.

INTRODUCTION

The British Museum (Natural History) has for several years possessed Drepaninae
material from Szechwan and adjacent provinces of China, chiefly from the collections
of Charles Oberthur and J. H. Leech. In the early 1960's I was able to study the
Drepaninae from the valuable collection made by the late Dr. H. Hone (see Gross,
1962), now housed in the Museum Alexander Koenig, in Bonn, Germany, which
includes examples from eastern and central provinces of China where little or no
previous collecting had been carried out. The collection at Bonn together with the
BM(NH) material and specimens from several European museums and the United
States National Museum have made possible this survey of the Drepaninae of China.
The opportunity has been taken to broaden the scope of this paper to include
related species from India, Japan and other areas associated zoogeographically with

4 A. WATSON

China wherever sufficient material was available or where knowledge of these species
is particularly relevant to the identification and distribution of the Chinese species.
This inclusion of non-Chinese species anticipates, to some extent, future records of
species not yet known from China. Seven of the Drepaninae genera represented in
China have been revised as a result of this wider treatment.

The nominal genera Ditrigona Moore (1887 : 258), Peridrepana Butler (1889 : 43)
and Leucodrepana Hampson (1892 : 333) are excluded from discussion in the
following paper as they are currently being revised by Dr. C. Wilkinson. A revision
of Teldenia Moore (1882 : 119) was published by Wilkinson (1967) during the final
stages of the preparation of this paper. Two species of Teldenia have been recorded
from China.

Apart from the extensive collection in the Museum Koenig, Bonn, important
material has been lent to me by the following museums and institutes : Hope
Department, University Museum, Oxford, U.K. ; Landhauptstadt Wiesbaden
Stadtisches Museum, Germany ; Museum national d'Histoire naturelle, Paris,
France ; Naturhistorisches Museum, Vienna, Austria ; Naturhistoriska Riksmuseet,
Stockholm, Sweden ; Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands ;
United States National Museum, Washington, D.C., U.S.A. ; Zoological Institute,
Academy of Sciences of the U.S.S.R., Leningrad, U.S.S.R. Further material was
borrowed from the private collections of Drs. F. Daniel (Munich), H. Inoue (Japan),
and H. Yamamoto (Japan). The types of nearly all the nominal species mentioned
in the text have been examined by the author either in the BM(NH) or in European
museums. The types of Matsumura species have been kindly studied for me by
Dr. H. Inoue in Japan. Lectotypes have been selected where necessary and where
it has been possible to examine the syntypic material.

I should like to acknowledge the generous co-operation of the following workers
who have helped by arranging loans of material or in other ways : Dr. C. Besuchet,
Geneva, Switzerland ; Dr. F. Daniel, Munich, Germany ; Dr. D. Davis, Washington,
D.C., U.S.A. ; Dr. W. D. Duckworth, Washington, D.C., U.S.A. ; Dr. M. Falkovitch,
Leningrad, U.S.S.R. ; Mr. D. S. Fletcher, London, U.K. ; Dr. W. Forster, Munich,
Germany ; Dr. E. Franz, Frankfurt-am-Main, Germany ; Dr. F. J. Gross, Wies-
baden, Germany ; Dr. H. J. Hannemann, Berlin, D.D.R. ; the late Dr. B. Hanson,
Stockholm, Sweden ; Dr. H. Inoue, Fujisawa, Japan ; Dr. F. Kasy, Vienna,
Austria ; Dr. A. I. Kurentzov, Vladivostok, U.S.S.R. ; Dr. C. Lemaire, Paris,
France ; Dr. B. Mannheims, Bonn, Germany ; Dr. I. W. B. Nye, London, U.K. ;
Dr. E. C. Popham, Salford, U.K. ; Dr. U. Roesler, Bonn, Germany ; Dr. K. Sattler,
London, U.K. ; Dr. H. Schroder, Frankfurt-am-Main, Germany ; Mr. E. Taylor,
Oxford, U.K. ; Dr. E. Todd, Washington, D.C., U.S.A. ; Professor G. C. Varley,
Oxford, U.K. ; Dr. P. Viette, Paris, France ; Mr. P. E. S. Whalley, London, U.K. ;
Dr. C. Wilkinson, Portsmouth, U.K. ; Dr. H. Yamamoto, Fukuoka, Japan. The
technical assistance of Miss K. Brookes, Miss R. Hauenstein and Mrs. J. E. Saunders
is also gratefully acknowledged.

It would be difficult to overestimate the magnanimous help given to me by the
late Dr. H. Hone of the Museum Koenig, Bonn, who made available his superb

CHINESE DREPANINAE 5

Chinese collection. My thanks are also to his widow whose many kindnesses during
my work on the collection at Bonn are not forgotten.

The photographic work was done in the Photographic Section of the British
Museum (Natural History) under the supervision of Mr. M. G. Sawyers.

Text-figures i, 5, 9, 12, 18, 19, 20, 27, 57, 61, 108, 114, 128, 132, 133, 138, 143,
149, 153, and 158 were drawn by Mr. Arthur Smith. The remaining drawings,
except for text-figures 89-96, 101-104, 165-182, 203-218, 251-274 and 291-293,
which were drawn by the author, were prepared by Mrs. J. E. Saunders.

The term ' comb, rev.' is used to denote a change in the combination of a specific
or subspecific name and a generic name which restores a previously published
combination ; ' comb, n.' is used in the usual way to denote a new combination ;
sp. rev.' and ' ssp. rev.' are used to denote, respectively, species and subspecies
names which have been removed from synonymy. BM(NH) is an abbreviation of
British Museum (Natural History). Names of other institutions have been short-
ened, not abbreviated in the strict sense, the full titles being given above. Biblio-
graphical references in the text are given mainly in a shortened form, the full refer-
ence appearing at the end of the paper.

The form of the descriptions is similar to that in Watson (1965 : 7) except that
the fore wing measurements are given in the following sequence : range of measure-
ments in the material examined from apex to centre of mesoscutum, followed, in
parentheses, by the number of specimens measured. Some of the more unusual
morphological characters merit comment here. For example, in Cilix, Sc + Ri
(vein 8) anastomoses with the base of the cell a short distance after it arises from the
base of the hind wing, in contrast with most other Drepanidae in which Sc -\- Ri
anastomoses with or approximates to Rs for a short distance distal to the end of the
cell. The only other exceptions found so far are Phalacropsis carnosa Swinhoe,
Phalacra edentata Hampson, Phalacra kerara Swinhoe and Phalacra tenera Swinhoe
(see Gaede 1931, for references to original descriptions), all of which are Oriental
species that have not been recorded from China. Also unusual in the Drepaninae
is the presence in the males of Pseudalbara of a vestigial frenulum, which is present
as a short costal process concealed by scales. All other Drepaninae have a reason-
ably well-developed frenulum in the male, as in the Madagascan Nidarinae, but in
contrast with the Oretinae, the males and females of which lack a frenulum.
Modification of the seventh as well as the eighth abdominal sternum in the male is
present in the abdomen of Paralbara, Albara, Betalbara, Agnidra, Thymistida,
Nordstroemia, Didymana and Strepsigonia. The seventh sternite may be sym-
metrical or asymmetrical bilaterally. In Strepsigonia two separate medial sclerites
seem to have been derived from the seventh abdominal sternum ; both sclerites
are bilaterally asymmetric. The medial eighth abdominal sternite is usually
bilaterally symmetric, but can be asymmetric as in Paralbara. Lateral sacs are
found associated with the pleural region of the eighth segment in some species of
Betalbara and Agnidra, and in one species of Nordstroemia. In some species of
Betalbara and Agnidra separate lateral sclerites are placed on either side of the
normal medial eighth sternite.

6 A. WATSON

The key to genera and most of the keys to species are based on males alone.
There are two main reasons for this : firstly that the females of some species are
not yet known, and secondly that it is difficult or impossible at this stage to identify
the females of some groups of closely related species.

Reference to a particular country or area in the paragraph on distribution in each
species description indicates that material from there has been identified during
the present study.

DISTRIBUTION

Generic distribution. (See Table i.) Most of the genera of Drepaninae which
occur in China have been recently revised, or are fairly well known in that their
taxonomy appears to be reasonably satisfactory and that identification of the
included species presents little difficulty. It has been possible, therefore, to show
in Table i the world distribution of these genera. Where genera have been revised
in the present paper, details of the distribution of all the included species have been
given in the Table ; the species of recently revised genera are listed in the Table
only if they are known to occur in China, but the full world distribution of each
genus is indicated. The species of other genera whose taxonomy is in an acceptable
state but which have not been revised recently or in this paper are listed fully. It
has been possible to give some idea in the Table of the distribution of the remaining
genera only when the generic placement of the included species is considered to be
probably correct.

Nine of the 26 genera of Drepaninae present in China are either endemic to the
Indo-Chinese Subregion or have a high proportion of Indo-Chinese species with
incursions chiefly into the south-eastern limits of the Palaearctic Region and into
the Malayan Subregion. Tridrepana and Canucha extend to the Papuan Sub-
region, including the Solomons {Tridrepana). Palaeodrepana, Drepana and Cilix do
not occur east of the Indo-Chinese Subregion but extend westwards into Europe
(including Britain). Although some taxonomic reappraisal of Strepsigonia, Drape-
todes and Hyalospectra is needed, it seems likely that the former two will prove to be
chiefly Malayan, while the range of Hyalospectra will probably prove to be com-
parable with that of Canucha or Tridrepana.

Callidrepana is unique in the Drepaninae in that it is represented both in the
Oriental Region and in the Ethiopian Region where three West African species are
known (see Watson, 1965). No comment can be made on the details of its Oriental
distribution until a generic revision has been carried out.

Although the overall pattern of distribution may be distorted by the possibility
of differential extinction and the certainty that the areas involved have not been
uniformly covered by collectors, it seems reasonable to suppose that the apparent
high percentages of endemism in the Indo-Chinese Subregion represent the real
pattern of distribution. Thymistida, Didymana and Thymistadopsis, for example,
are unknown beyond the limits of the Indo-Chinese Subregion, while in Paralbara,
Agnidra, Betalbara, Nordstroemia, Deroca and Auzata the percentages of specific
endemism in this Subregion range from 60% in Betalbara to 83% in Auzata. This

CHINESE DREPANINAE 7

high degree of endemism possibly reflects the enhanced opportunities for speciation
in the varied ecological conditions resulting from the Cenozoic elevation of the
Himalayas and suggests that this part of south-eastern Asia can reasonably be
considered as the probable centre of origin for several Drepaninae genera. The
Papuan Subregion forms another centre of endemism in Tridrepana and Canucha —
possibly a secondary centre at least in Tridrepana which has a greater proportion
of endemic Indo-Chinese species than Papuan species. A pattern of distribution
similar to that in Tridrepana occurs in Oreta (Drepanidae, Oretinae) (see Watson,
1967).

The small genus Cilix is known from Western Europe, the Mediterranean area
(including North Africa), the Middle East, Afghanistan, northern India, China,
Korea, Japan and south-eastern Russia ; a pattern which suggests a dispersal route
for the genus from a possible Indo-Chinese centre, where the greatest degree of
endemism occurs. The pre-Glacial pattern of distribution might, however, have
revealed a more northerly route or a much broader North-South distribution.

Specific distribution. (See Table 1.) A total of 76 species of Drepaninae have so
far been described from China. A further one or perhaps two species of the genus
Drapetodes occur there but have not yet been described. Fifty-three species are
endemic to the Indo-Chinese Subregion. Sixteen species are shared by the Indo-
Chinese Subregion and the Manchurian Subregion of the Palaearctic Region, with
two of these species, Drepana curvatula and Palaeodrepana harpagula, extending
into Western Europe and the British Isles. Three species are found in both the
Indo-Chinese and Malayan Subregions ; one species occurs in the Indo-Chinese,
Malayan and Indian Subregions ; one species is common to the Indo-Chinese and
Malayan Subregions and Celebes, and one to the Indo-Chinese, Indian and Malayan
Subregions and Celebes. One species is Manchurian but is not known from else-
where in China. The species of Drepaninae found in China are thus predominantly
endemic to the Indo-Chinese Subregion, with incursions chiefly into the adjacent
Malayan and Indian Subregions of the Oriental Region and into the Manchurian
Subregion of the Palaearctic Region. Only four of the Chinese species extend
beyond these limits ; they are Drepana curvatula and Palaeodrepana harpagula
which are found in Western Europe, and Tridrepana fulvata and Canucha specularis
whose ranges extend as far east as Celebes. Except for specularis, each of the latter
four species is represented at the periphery of its range by a subspecies different
from that occuring in China.

Within China, judging from the high degree of endemism and the presence of
several groups of closely related species, the provinces of Szechwan and Yunnan
apparently form a centre of evolutionary activity for many genera (the Yunnan
Centre of de Lattin, 1957), with a second, less well defined centre in the hilly eastern
provinces of Chekiang and Fukien.

The distribution of the Chinese species of Oreta Walker and Cyclura Warren
(Drepanidae, Oretinae) (see Watson, 1967), is comparable with that of the Drepaninae
except that no species of Oreta, or indeed Oretinae, occurs in Western Europe and
there is apparently no zoogeographical match in the Drepaninae for the Nearctic

8 A. WATSON

Oreta rosea Walker which is closely allied to the Chinese species 0. pulchripes Butler.
One genus of Drepaninae, Drepana Schrank, is represented both in China and the
Nearctic Region, but in contrast with Oreta the two Nearctic species of Drepana,
arcuata Walker (1855 : 164) and bilineata Packard (1864 : 376), apparently have
their closest relatives in Western Europe. However arcuata is not taxonomically
distant from curvatula Borkhausen, which is found not only in Western Europe but
also in the Oriental Region (including China) and the eastern limits of the Palae-
arctic Region.

Table i

The World Distribution of the genera, species and subspecies of Drepaninae represented in

China

p

)

Oriental Region [Subdivisions

m

Z

is

are those of Gressitt (1956)]

3"

5'

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P

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0'

Cfi

5"

3

3

cr
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(TO

5'
3

3

(Col. 1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(IO)

(11)

(12)

Albara Walker

X

X

X

reversaria Walker

X

X

X

reversaria reversaria

X

reversaria opalescens

Warren

X

X

Par albara gen. n.

X

X

X

muscularia Walker

X

X

perhamata Hampson

X

spicula sp. n.

X

X

X

pallidinota sp. n.

X

CHINESE DREPANINAE

(Col. I)

Thymistida Walker

nigritincta Warren
tripunctata Walker
undilineata Warren

Agnidra Moore

hoenei sp. n.

fulvior sp. n.

furva sp. n.

fenestra Leech

specular ia Walker

corticata Warren

corticata corticata
corticata francki ssp. n.

vinacea Moore

scabiosa Butler

scabiosa scabiosa
scabiosa fix seni Bryk

fuscilinea Watson

discispilaria Moore

Betalbara Matsumura

acuminata Leech

manleyi Leech

manleyi manic vi
manleyi prolatior ssp. n.

prunicolor Moore

leucoslicta Hampson

flavilinea flavilinea Leech

flavilinea flavilinea Leech
flavilinea shensiensis ssp. n.

cupreogrisea Hampson

rectilinea sp. n.

rugosa sp. n.

violacea Butler

robusta Oberthiir

Pseudalbara Inouc
parvalu Leech
fusci fascia sp. n.

Nordstroemia Bryk
vira Moore
bicostata Hampson

bicoslata bicostata

bicostata opalescens Oberthiir
agna Oberthiir
problematica Bryk
argenticeps Warren
sumatrana Roepke
japonica Moore
grisearia Staudinger

(2)

(3)

4)

(5

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

■

X

X

X

X

X

X

X

X

X

X

(6)

(7)

(8)

(9)

(io)

(12)

A. WATSON

(Col. i)

recava sp. n.
lilacina Moore
simillima Moore
siccifolia Roepke
ochrozona Bryk
duplicata Warren
humerata Warren
undata sp. n.

Didymana Bryk
bidens Leech

Palaeodrepana Inoue
harpagula Esper

harpagula harpagula Esper
harpagula olivacea Inoue
harpagula emarginata
ssp. n.
harpagula bitorosa ssp. n.
binaria Hufnagel
cultraria Fabricius

Strepsigonia Warren [Generic
revision needed. 6 species at present
recognized]

diluta Warren

other material examined

Canucha Walker

curvaria Walker
sublignata Warren

ST
n
P

5'

3

5"

3

7;

(3)

Oriental Region [Subdivisions
are those of Gressitt (1956)]

(4)
x

p

p

3

m

c
cr

CTQ

o'

(6)

o

►5'

►T3

(8)

(9)

p

c
p

3
C/J

c
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3"

p

O

H

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T)

r+

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5S
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12

CHINESE DREPAXIXAK

(Col. i)

specularis Moore

duplexa Moore

duplexa duplexa Moore
duplexa birmana Moore

bouvieri Oberthiir

miranda Warren

Drepana Schrank [Generic revision
needed. 1 1 species are at present
recognized : generic distribution
doubtful.]

curvatula Borkhausen
curvatula curvatula Borkhausen

curvatula acuta Butler
rufofasciata Hampson
pallida Moore

pallida pallida Moore
pallida cretacea Moore
pallida flexuosa ssp. n.
pallida nigromaculata Okano
dispilala Warren

dispilata dispilata Warren
dispilata rufata ssp. n.
dispilata grisearipennis Strand
other species

Tridrepana Swinhoe [See revision by
Watson (1957) f° r distribution of
remaining 24 species.]
fulvata Snellen

tulruta fulvata Snellen

fulvata brevis Watson

fulvata, undescribed subspecies
arihana Matsumura

arikana arikana Matsumura

arikana falcipen nis
crocea Leech
unispina Watson
finita Watson
rubromarginata Leech

; ubromarginata rubromarginata
Leech

rubromarginata indica Watson
thermopasta Hampson
maculosa Watson
marginata Watson
fulva Hampson
other species

Callidrepana Hampson [Generic revision
needed. 26 species at present
recognized.]

(2)

(3)

(4)

(5)

X

X

X
X

X

X

X

X

X

X

X

X

X

X

X

X

X
X

X

X

X

X

x

X
X

X

X

X

X

X

X

X
X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

(6)
X

(7)
x

(8)
X

(9)

(10)

(11)

(12)

XXX

XXX

A. WATSON

(Col. i)

patrana Moore

patrana patrana Moore
patrana palleollus Motschulsky

ovata sp. n.

gemina sp. n.

gemina gemina
gemina curta ssp. n.

hirayamai Nagano

hirayamai hirayamai Nagano
hirayamai forcipulaia Nagano

other species

Drapetodes Guenee [Generic revision
needed : 10 species at present recognized]

i or 2 unidentified species (Chinese)

other species

Thymistadopsis Warren

albidescens Hampson

trilinearia Moore

trilinearia trilinearia Moore
trilinearia pidvis Oberthiir

undulifera Hampson

Deroca Walker [Generic revision by
Watson (1959).]

hyalina Walker

hyalina hyalina Walker
hyalina latizona Watson
hidda Swinhoe

hidda hidda Swinhoe

Oriental Region [Subc

ivisions

p

are those of Gressitt (1956)]

O'

^3

p_

3
St

n

CD

a

hd

fD

°s

5'

3

-1

0R3 O

5' E

3 3

to

p"
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p

3

5'

3

w.

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0"

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3'

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2
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(2)

(3)
X

X

X
X

(4)

X
X

X
X

X

X

(5)

X
X

X

(6)

(7)

(8)

(9)

(IO)

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X
X

x

X
X

X

x

X

X

X

X
X

X

X
X

X
X

w

c

*2
p'
3

°2
5"

p

7C
ft

OfQ

(II)

CHINESE DREPANIXAE

13

(Col. 1)

huida bifida Watson

pulla Watson

inconclusa Walker

inconclusa inconclusa Walker
inconclusa concinna Warren
inconclusa carinata Watson
inconclusa phasma Butler

Callicilix Butler

abraxata Butler

abraxata abraxata Butler
abraxata nguldoe Oberthiir

Auzata Walker [Genus revised by
Watson (1959)]

semipavonaria Walker

chinensis Leech

chinensis chinensis Leech
chinensis prolixa Watson
chinensis arcuata Watson

simpliciata Warren

superba Butler

superba superba Butler
superba cristata Watson

minuta Leech

minuta minuta Leech
minuta spiculata Watson

ocellata Warren

Macrocilix Butler

mysticata Walker

mvsticata mysticata Walker
mysticata watsoni Inoue
mysticata brevinotata Watson
maia Leech
taiwana Wileman
orbiferata Walker

orbiferala orbiferata Walker
orbiferata cilicoides Snellen
sericea Warren
Hyalospectra Warren [Generic revision
needed. Generic distribution doubtful]
hyalinata Moore

Cilix Leech

glaucata Scopoli
asiatica Bang- Haas
depalpata Strand
filipjevi Kardakoff

filipjevi filipjevi Karkakoff
filipjevi malivora Inoue
palula sp. n.

(2)

(3)

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A. WATSON

(Col. i)

danieli sp. n.
tatsienluica Oberthur

Macrauzata Butler [Genus in need of
revision]

fenestraria Moore
minor Okano
maxima Inoue

maxima maxima Inoue
maxima chinensis Inoue
other material examined

Phalacra Walker [Genus in need of
revision. Generic distribution doubtful.
12 species at present recognized.]

strigata Warren

undescribed species

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Key to Genera based on the Oriental Species

Males

Apex of fore wing falcate (e.g. PI. i, fig. 294) ....... 5

Apex of fore wing not falcate (e.g. PL 12, fig. 373) ...... 2

Ground-colour of upper surface of wings white ....... 3

Ground-colour of upper surface of wings not white . . DRAPETODES (p. 124)

Anal angle of upper surface of hind wing with some yellow coloration

MACROCILIX (p. 131)
Anal angle of upper surface of hind wing with no yellow coloration ... 4

Wings transparent, sparsely scaled ...... DEROCA (p. 127)

Wings not transparent, ground-colour opaque white .... CILIX (p. 137)

CHINESE DREPANINAE 15

5 Frenulum vestigial, not visible without removal of surrounding scales. Colour-

pattern as in PI. 3, figs. 320 or 321 . . . . PSEUDALBARA (p. 65)

- Frenulum well-devloped. Colour-pattern not as in PI. 3, figs. 320 or 321 . . 6

6 Ground-colour of wings white .......... 7

- Ground-colour of wings not white . . . . . . . . . 11

7 Wings with large transparent areas as in PI. 13, fig. 378 . HYALOSPECTRA (p. 136)

- Wings not as in PI. 13, fig. 378 ......... 8

8 Transverse lines on upper surface of hind wing lunulate or dentate ... 9

- Transverse lines on upper surface of hind wings not indented

THYM1STADOPS1S (p. 124)

9 Antennae lamellate. Hind tibia with two pairs of well-developed spurs

AUZATA (p. 130)

- Antennae not lamellate. Hind tibia with one pair of spurs (Drepana) ; or with two

pairs of spurs, in which case the outer spur of the proximal pair is vestigial and

not visible without removal of the surrounding scales (Callicilix) . . . 10

10 Hind tibia with one pair of spurs ... . . DREPANA (part) (p. 103)

- Hind tibia with two pairs of spurs ..... CALLICILIX (p. 127)

11 Hind wing with tail or angulate process at outer angle or anal angle ... 12

- Hind wing without tail or angulate process at outer angle or anal angle . . 13

12 Antenna bipectinate. Outer margin of fore wing with process THYMISTIDA (p. 24)

- Antenna lamellate. Outer margin of fore wing without process . PHALACRA (p. 143)

13 Highly lustrous scales present on upper surface of wings ; especially along costa,

outer margin and at distal end of cell .... CALLIDREPANA (p. 113)

- Upper surface of wings without highly lustrous scales ..... 14

14 Outer margin of fore wing with process (e.g. PI. 4, fig. 331) . . . . 15

- Outer margin of fore wing without process ....... 16

15 Ground-colour of upper surface of fore wing the same as that of hind wing. Distal

third of hind wing yellowish brown, not paler than rest of wing (e.g. PI. 4, fig. 331)

PALAEODREPANA (p. 92)

- Ground-colour of upper surface of fore wing much darker than that of hind wing.

1 >istal third of hind wing yellowish white, paler than rest of wing (PI. 11, fig. 357)

DIDYMANA (p. 92)

16 Base and proximal part of shaft of antennae clothed in highly lustrous scales

NORDSTROEMIA (part) (p. 69)

- Antennae without highly lustrous scales . . . . . . . . 17

17 Ground-colour of upper surface of wings yellow . . . TRIDREPANA (p. m)

- Ground-colour of upper surface of wings not yellow ...... 18

18 Postmedial fascia on upper surface of fore wing straight (e.g. PI. 3, fig. 315) . . 19

- Postmedial fascia on upper surface of fore wing not straight .... 24

19 Postmedial fascia on upper surface of hind wing arcuate ; dentate or non-dentate . 20

- Postmedial fascia on upper surface of hind wing straight ; not dentate . . 21

20 Hind tibia with two pairs of spurs ...... AGNIDRA (p. 29)

- Hind tibia with one pair of spurs ..... DREPANA (part) (p. 103)

21 Proximal half of costa of fore wing concave or straight ; strongly convex near apex

(e.g. PI. 11, fig. 366) CANUCHA (p. 99)

- Proximal half of costa convex ; not strongly convex near apex .... 22

22 Antenna uniserrate ...... AGNIDRA (part) (p. 29)

- Antenna bipectinate or lamellate ......... 23

23 Subterminal fascia on upper surface of fore and hind wings represented by series of

dots ; fascia darker than rest of wing ...... ALBARA (p. 16)

- Subterminal fascia absent or represented on fore and hind wings by markings other

than dots BET ALBARA (p. 46)

24 Hind tibia with one pair of spurs ......... 25

16 A. WATSON

- Hind tibia with two pairs of spurs ......... 26

25 Postmedial fascia on upper surface of hind wing lunulate ; the convex side of each

lunula faces base of wing STREPSIGONIA (p. 97)

- Postmedial fascia on hind wing not lunulate, or, if lunulate, the convex side of each

lunula faces outer margin of wing ..... DREPANA (part) (p. 103)

26 Hind wing without markings or with trace of pattern at anal margin

NORDSTROEMIA (part) (p. 69)

- Hind wing with well-developed pattern . . . . . . . .27

27 Antemedial fascia on fore and hind wings straight (PI. 13, fig. 385)

MACRAUZATA (p. 142)

- Antemedial fascia on fore and hind wing not straight ...... 28

28 Arms of uncus in male genitalia robust and widely separated . PARALBARA (p. 19)

- Male genitalia not as above ...... AGNIDRA (part) (p. 29)

ALBARA Walker
(PI. 1, figs. 296-297 ; Text-figs. 1-7)

Albara Walker, 1866 : 1566. [Name adopted from multiple original spelling by the first
reviser, Kirby, 1892 : 734.] Type-species, by monotypy, Albara reversaria Walker, 1866 :

1567-
Albara Walker ; Gaede, 1931 : 31. [Partim.]
' Albaria ' ; Walker, 1866 : 1567. [An incorrect original spelling of Albara Walker.]

o". Palp extends to just above labrum ; proximal three-fifths of antenna bipectinate.
Upper surface of wings, thorax and abdomen dark violet-grey ; fore wing with weakly marked
antemedial fascia, strongly marked oblique postmedial fascia, short arcuate line proximal to
postmedial near wing apex, and with ill-defined interrupted subterminal fascia ; hind wing
similar to fore wing. Under surface of wings, thorax and abdomen very pale violet-grey,
paler and more yellowish at anterior and posterior margins. Vein i?i arises from near distal
end of cell and R2 from areole in fore wing ; Sc + i?i approximates to Rs for short distance
distal to end of cell in hind wing. Mesothoracic tibia with one pair of terminal spurs ; meta-
thoracic tibia with two pairs of spurs.

o* genitalia : valves short, with processes ; arms of uncus widely separated ; socii small ;
diaphragma with strongly sclerotized medial structure ; eighth abdominal tergite and sternite,
and asymmetric seventh sternite forming part of genital apparatus.

$. As for male but with weakly biserrate antennae.

§ genitalia with asymmetrically placed ostium ; corpus bursae without signum ; eighth
and ninth tergites lobate, well developed.

Albara is probably most closely allied to Paralbara gen. n. It can be separated
from the latter by differences in the wing-pattern and in the male and female
genitalia.

Albara Walker, sensn Gaede (1931), has been partly restricted and revised by
Bryk (1943) and Inoue (1953, 1962). The only species which can be placed correctly
in Albara is in fact the type-species. The remaining species are transferred in this
paper to one or other of the following genera : Paralbara gen. n., Agnidra Walker,
Pseudalbara Inoue, Betalbara Matsumura, Nordstroemia Bryk, and Thymistadopsis
Warren.

Distribution (see Table 1) : N. India, China, Formosa, Malaysia and Indonesia.

CHINESE DREPANINAE 17

Albara reversaria Walker

(PI. i, figs. 296, 297 ; Text-figs. 1-7)

' Albaria ' reversaria Walker, 1886 : 1567.

Two subspecies are known : the nominate subspecies (Sumatra), and opalescens
Warren (India, Formosa, China). Two males and eleven females in the BM(NH),
from Malaya, differ from the Sumatran material in minor genitalic characters and
may prove to represent a new subspecies.

Albara reversaria reversaria Walker
(Text-figs. 1-4)
Albara reversaria Walker ; Gaede, 1931 : 33.

Readily distinguished from opalescens Warren by the <$ genitalia (Text-figs. 1-3),
particularly by the shape of the anellus, basal valve processes, medial gnathus
process, socii and uncus.

Measurements. <$ 160 mm. (1) ; $ 15-0-18-5 mm. (8).

Material examined. Holotype $, Sumatra ; in the Hope Department Museum,
Oxford. An examination of the genitalia slide made from the abdomen found
attached to the type (Drepanidae slide No. 263) has shown that this is not the
original abdomen. However the BM(NH) possesses a male and female from
Sumatra, the abdomens of which had not been glued on and are doubtless genuine.

Other material. BM(NH). Sumatra : 1 <$, Barisan Range, Western slopes,
2500 ft., x-xi.1961 (Pratt) ; 1 $, Lebong Tandai, 6.xii.i92i (Brooks).

Albara reversaria opalescens Warren stat. n.
(PI. 1, figs. 296, 297 ; Text-figs. 5-7)

Albara opalescens Warren, 1897 : 12.

Albara opalescens Warren ; Warren, 1922 : 468. [Fig.]

Albara opalescens Warren ; Gaede, 1931 : 32.

A Ibara griseotincta Wileman, 1914 : 268. syn. n.

A Ibara griseotincta Wileman ; Gaede, 1931 : 31.

Albara horishana Matsumura, 1921 : 948. syn. n.

Apparently indistinguishable externally from the nominate subspecies, but with
distinctive male genitalia. The female is unknown.

Wing, <J 15-0-19-5 mm. (9).

I have not seen the type of horishana Matsumura, but this has been kindly
examined for me in Japan by Dr. H. Inoue.

Material examined. Types. opalescens. LECTOTYPE <£, here selected,
labelled : Khasis, Oct. 1895, Nat. Coll. ; Albara opalescens Warr. Type $ ;
Drepanidae genitalia slide No. 782. In the BM(NH).

i8

A. WATSON

Figs. 1-7. Albara, genitalia. 1-4, reversaria reversaria. 1, $ ; 2, aedeagus ; 3, seventh
and eighth sternites, and eighth tergite ; 4, $. 5-7, reversaria opalescens, $. 5, ^ ; 6,
aedeagus ; 7, seventh and eighth sternites, and eighth tergite.

CHINESE DREPAMN \l 19

griseotincla. Holotypecy, Formosa : Kanshirei, 1,000 ft., 20,29^.1906 (Wileman).
In the BM(NH).

Other material. BM(NH). Formosa : 1 $, Central Formosa, 1959. India :
6 o*. Khasis, x.1895 (2 ex.). Museum Koenig, Bonn. China : 1 $, Kwangtung,
Linping, i.iv.1924 {Hone).

PARALBARA gen. n.

(PI. 1, figs. 294, 295, 298, 299 ; Text-figs. 8-20)
(Gender : feminine)

Type-species Fascellina muscularia Walker, 1866 : 1554.

cJ. Palp extends to just above labrum. Antenna bipectinate from base to about three-
quarters <>i its length. Upper surface of wings dull greyish brown or buff. Vein 7?i arises
from distal end of cell in fore wing and Rt from distal end "I areole. Fore wing with poorly
marked, lunulate antemedial and postmedial fasciae ; usually with ill-defined subterminal
fascia or spots ; and with spot, or group of spots or pit. ttes, al posterior angle of cell. In
hind wing Sc -\ Rj approximates to Rs for short distance distal to end of cell ; similar to fore
wing m pattern but with subterminal fascia very weakly marked. I'nder surface of wings
pale lustrous grey or greyish butt, with diffusely marked postmedial and subterminal fascia ;
wings darkest proximal to postmedial fascia.

Thorax and abdomen similar in colour to adjacent surface of wings. Mesothoracic tibia
with one pair of spurs, metathorai u tibia with two pairs of spurs.

<J genitalia : seventh abdominal sternum modified into two sclerites, asymmetric ; eighth
tergite tapered or truncate posteriorly ; eighth steimte narrow anteroposteriorly ; valve
small with large inwardly directed process at base ; sao as well-developed ; diaphragma with
medial scleroti/ation, very Btrongly developed in spicula ; uncus bifid, robust; aedeagus
sinuous.

$. As for J but with weakly biserrate antennae and apex of fore win- more strongly pro-
duced.

$ genitalia : seventh abdominal sternite emarginate posteromedially ; bin tlatrix

with single ovate, concave signurn, or without signum ; ostium surrounded by nearly circular
plate ; eighth and ninth tergites moderately well si Unitized.

Paralbara can be distinguished from its closest apparent relative, Albara Walker,
by the wing-pattern and by the genitalia of both sexes.

I include in this genus two species transferred from Albara Walker and two new
species.

Distribution. N. India, Sikkim, Bhutan, Burma, China {muscularia, pallidinota,
spicula) and Malaysia. (See Table 1.)

Key to species. Both Sexes

Single, large, pale, diffusely marked patch distal to end of cell on fore wing (PI. 1,

fig. 295). $ genitalia as in Text-fig. 20 . . . . pallidinota (p. 24)

Fore wing with pale patch absent distal to end of cell or if present then associated with

other patches immediately posterior to it. $ genitalia not as in Text-fig. 20 . 2

Collar and base of antenna orange-yellow ; anterior part of subterminal fascia on

20 A. WATSON

fore wing dark brown ; costa orange-yellow ; postmedial fascia on hind wing
simple (PI. i, fig. 294). Genitalia as in Text-figs. 8-1 1 . . muscularia (p. 20)

- Collar and base of antenna without orange-yellow scales ; anterior part of sub-

terminal fascia on fore wing pale grey (lighter than surrounding area of wing (PI. 1,
figs. 298, 299) ; costa not orange-yellow ; postmedial fascia of hind wing double.
Genitalia not as in Text-figs. 8-1 1 ......... 3

3 Genitalia as in Text-figs. 12-15 ....... perhamata (p. 22)

- Genitalia as in Text-figs. 16-18 ........ spicula p. 22)

Paralbara muscularia (Walker) comb. n.
(PI. 1, fig. 294 ; Text-figs. 8-11)

Fascellina muscularia Walker, 1866 : 1554.

Albara muscularia (Walker) Warren, 1922 : 468.

Albara muscularia (Walker) ; Gaede, 1931 : 32.

Drepana orphnina Hampson, [1893] : 337. syn. n.

Albara orphnina (Hampson) Warren, 1922 : 468.

Albara orphnina (Hampson) ; Gaede, 1931 : 32. [The male and female syntypes of Albara

orphnina ab. subpallida Warren are conspecific with the neotype of muscularia.']
Albara inaequidiscata Warren, 1922 : 469. [Good figs.] syn. n.

Distinguished from the closely allied perhamata by the following characters:
collar and base of antenna orange-yellow ; fore wing with anterior part of sub-
terminal fasciae dark brown ; costa orange-yellow ; postmedial fascia of hind wing
simple, all fascia poorly marked ; distal half of wing only slightly paler than
proximal half. The male and female genitalia are characteristic (Text-figs. 8-1 1).

Wing. $ 14-5-20-5 mm. (24) ; $ 17-0-20-5 mm. (10).

There is some variation in the size of the whitish cell markings on the upper
surface of the wings. In the lectotype and paralectotype of inaequidiscata and in
one other male specimen these markings are large and conspicuous (see Warren,
1922 : pi. 4gf). In the remaining specimens the cell markings are either minute or
absent.

Distribution. N. India, N. Burma, and China.

Material examined. Types, muscularia. No trace can be found of the original
type material which was stated by Walker (1866 : 1554) to be male, from ' North
Hindostan ', and to be deposited in the collection of A. E. Russell. The Russell
collection, however, is apparently lost (see Horn and Kahle, 1937 : 380). There is
no trace of the type material either in the BM(NH) or in the Hope Department
Museum, Oxford, where much of Walker's original material is deposited. I there-
fore select as NEOTYPE a $ in the collection of the BM(NH) labelled: Darjeeling,
22.viii.1886 (H. J. Elwes) ; Rothschild Bequest B. M. 1939-1 ; B. M. negative
No. 29101.

orphnina. Holotype $, India, Naga Hills, 5500-7000 ft., viii-ix.1899 (Doherty) ;
Drepanidae genitalia slide No. 948. In the BM(NH).

inaequidiscata. LECTOTYPE $, here selected Khasis Nat. Coll. ; H. J. Elwes ;
Albara inaequidiscata Type $ Warr. ; Rothschild Bequest B. M. 1939-1. In the
BM(NH).

CHINESE DREPANINAE

21

Figs. 8-ii. Paralbara muscularia, genitalia. 8, $ ; 9, ^ ; 10, <$ seventh and eighth
sternites and eighth tergite ; 11, aedeagus.

22 A. WATSON

Other material. BM(NH). India : 2 <$, 2 $, Assam, Khasis, 1894 ; 1 <$, 1 $
Cherrapunji, vii.1893 ; 4 (J, Darjeeling, 20.vii.1886, 6.iii.i889 (Elwes, Pitcher) ;
1 <$, Shillong, 26.x. 1918. 1 $, Bengal ; 1 £, N. India ; 1 <$. Sikkim : 3 J, 3 $,
14.ix.1888, 1889, 8.vii.i89i, ix.1909 (Elwes, Dudgeon, M oiler). Bhutan : 1 $.
Burma : 1 $, 1$, Upper Burma, Htawgaw, 6000 ft. (Swann) ; 1 <$. N. E. Burma,
Kambaiti, 7000 ft., 9.VL1934 (Malaise) ; 3 <J, Mt. Victoria, Pakokku, Chin Hills,
2200 m., 5-30. vi. 1938 (Heinrich). China : 1 $, Kwanhsien [this may prove to
represent a new subspecies] .

Paralbara perhamata (Hampson) comb. n.
(PI. 1, fig. 299 ; Text-figs. 12-15)

Drepana perhamata Hampson, [1893] : 33°-

Albara perhamata (Hampson) Warren, 1922 : 468. [Figs.]

Albara perhamata (Hampson) ; Gaede, 1931 : 32.

This species apparently forms a superspecies with spicula, from which it is distin-
guished only by the genitalia. It is separable from muscularia by the absence of
orange-brown scaling on the costa, collar and antenna, by the very pale postmedial
fascia on the fore wing, and by the usually yellowish brown distal half of the hind
wing, bordered proximally by a double postmedial fascia. The male and female
genitalia are also diagnostic.

Wing. <$ 16-0-19-9 mm. (12) ; $ 21-5 mm. (1).

Distribution. N. E. India and Sikkim.

Material examined. Type. I select as LECTOTYPE a $ syntype in the
BM(NH) labelled : Jaintia [N. E. India], 2-4000, Oct. 87 ; Drepana perhamata
Hmpsn $ ; Coll. H. J. Elwes ; Drepanidae genitalia slide No. 950.

Other material. BM(NH). India : 7 <$, Assam, Khasia Hills ; 1 $, Naga
Hills, 1500 ft., ix-x.1889 (Doherty). Sikkim : ix.1909 (M oiler).

Paralbara spicula sp. n.

(PI. 1, fig. 298 ; Text-figs. 16-19)

The male and female genitalia distinguish this species from perhamata ; in particular, the
shape of the seventh sternite, uncus, and the elongate gnathus processes and socii in the male,
and the ostial plate in the female.

Wing. 6* 14.5-17° mm - (5); ? 4 2 -5 m m - (*)■

Holotype £. S. China : [Kwangtung], Linpging, v. 1922 (Hone) ; Drepanidae
genitalia slide No. 947. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 2 $, Kwangtung, Lingping,
3. v. 1922 (Hone) ; 1 (J, Kwangtung, Canton (Hone).

Other material. 1 $, without abdomen, from Fukien, Kuatan, in the Museum

CHINESE DREPANINAE

23

Figs. 12-15. Paralbara perhamata, genitalia. 12, <$ ; 13, aedeagus ; 14, <J seventh and
eighth sternites and eighth tergite ; 15, $.

24 A. WATSON

Koenig, Bonn, probably represents this species, i <$, Borneo, in the Zoologisches
Museum, Berlin (see below).

This species apparently replaces perhamata in China, the two forming a super-
species. The single male from Borneo, Kina Balu, probably represents a new sub-
species of spicula.

Paralbara pallidinota sp. n.

(PI. i, fig. 295 ; Text-fig. 20)

Readily distinguished from its close allies perhamata and muscularia by the colour-
pattern and the distinctive female genitalia, especially the shape of the ostial plate
and the presence of a signum.

<$. Vertex of head and outer surface of palp dark brown ; front of head dark brown, but
brown just above labrum ; collar pale yellow-brown; antenna pale greyish brown.

Thorax and abdomen greyish brown dorsally, much paler ventrally. Wing-pattern of
upper surface as in PI. 1, fig. 295 : palest areas grey ; ground-colour brown with proximal
half of fore wing reddish brown ; slightly lustrous ; costa of fore wing dull yellowish orange.
Under surface of both wings very pale brownish grey, each with broad, slightly darker band
along outer margin ; fore wing darker at base costad and with dull yellowish orange costal
area distad. Outer surface of prothoracic leg dark brown ; outer surface of mesothoracic leg
pale greyish brown ; legs otherwise very pale brownish grey.

$ genitalia as in Text-fig. 20.

Wing. $ 20-0-22-0 mm. (2).

o*. Not known.

Holotype $. China : N. Yunnan, Likiang, c. 2000 m., 8.viii.i934 (Hone) ;
Drepanidae genitalia slide No. 953. In the Museum Koenig, Bonn.

Paratype. Museum Koenig, Bonn. China : 1 $, N. Yunnan, Likiang, 8.viii-
16.ix.1934 (Hone).

THYM1STIDA Walker
(PI. 1, fig. 300, PI. 14, figs. 389-392 ; Text-figs. 21-24)

Thymistida Walker, 1865 : 515. Type-species Thymistida tripunctata Walker, 1865 : 515, by

monotypy.
Thymistida Walker ; Warren, 1923 : 474.
Thymistida Walker ; Gaede, 1931 : 39.
Hybodrepana Bryk, 1943 122. Type-species Hybodrepanagrotesca Bryk, 1943 : 23, by monotypy

syn. n.

Thymistida appears to be most closely allied to Agnidra. Its most distinctive
feature is the presence of a short tail on the hind wing (see Plate 1).

Three species are known : nigritincta Warren (India, Burma), tripunctata Walker
(India, Burma, China) and undilineata Warren (1923 : 474) (N.E. India). (See
Table 1.) The species nigritincta together with the sole Chinese representative,
tripunctata, are dealt with below and the opportunity taken to unravel their
synonymy. The latter two species both occur in India and Burma and are similar
in colour pattern.

CHINESE DREPANINAE

25

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26

A. WATSON

Figs. 19, 20. Paralbara, $ genitalia : 19, spicula ; 20, pallidinota.

CHINESE DREPANINAE 27

Thymistida nigritincta Warren
(PI. 1, fig. 300 ; Text-figs. 21-24)

Thymistida nigritincta Warren, 1923 : 474. [Published simultaneously with rufa ; here

selected as the valid name for this taxon.]
Thymistida nigritincta Warren ; Gaede, 1931 : 39.
Thymistida rufa Warren ; 1923 : 475. syn. n.
Hybodrepana grotesca Bryk, 1943 : 23. syn. n.

This species can be distinguished from tripunctata Walker (q.v.) by the shorter
antennal pectinations in the male (longest pectination equal to about three quarters
greatest width of eye), the distinctively shaped costa of the fore wing, and by the
male and female genitalia.

Distribution. N.E. India and N. Burma.

Types, nigritincta. LECTOTYPE $, here selected, in the BM(XH), labelled :
Khasis, June 1895, Nat. Coll. ; Thymistida nigritincta Type $ Warr. ; Rothschild
Bequest B.M. 1939-1 ; B.M. negative No. 29118.

rufa. LECTOTYPE <$, here selected, labelled " Khasis ". ' In the BM(NH).

grotesca. Holotype $, N.E. Burma, Kambaiti ; Drepanidae genitalia slide No.
1047 ; in Naturhistoriska Riksmuseet Stockholm.

Thymistida tripunctata Walker

(PI. 14, figs. 389-39 2 )

Thymistida tripunctata Walker, 1865 :51s.

Thymistida tripunctata Walker ; Warren, 1923 : 474. [Good fig.]

Thymistida tripunctata Walker ; Hampson, [1893] : 343.

Thymistida tripunctata Walker ; Gaede, 1931 : 39.

Erosia cervinaria Moore, 1867 : 646. [Synonymized by Hampson [1893].]

Thymistida nigritincta divisa Bryk, 1943 : 2}>- syn. n.

This species is readily separated from the similarly marked nigritincta by the
much larger antennal pectinations in the male (longest pectination equal to over
twice greatest width of eye), the less sinuous costa of the fore wing, and by the male
and female genitalia.

Distribution. N.E. India, N. Burma and China (2 $, from Siao-Lou, Szechwan,
in the BM(NH)).

Types, tripunctata. Holotype <$, labelled ' E. India '. In the BM(NH). The
locality is given as ' Hindustan ' in the original description. The abdomen of the
holotype is missing.

cervinaria. The syntypes stated to be from Bengal are lost according to Horn
and Kahle (1937 : 380), but there is a short series in BM(NH) from the Moore Coll.
labelled cervinaria.

divisa. Holotype $, N.E. Burma, Kambaiti ; Drepanidae genitalia slide No.
1049. I n Naturhistoriska Riksmuseet, Stockholm.

28

A. WATSON

Figs. 21-24. Thymistida nigritincta, genitalia. 21, <J ; 22, ^ seventh and eighth sternites,
and eighth tergite ; 23, aedeagus ; 24, $.

CHINESE DREPANINAE 29

AGNIDRA Moore gen. rev.
(Pis. 1, 2, figs. 301-311 ; Text-figs. 25-71)

Agnidra Moore, [1868] : 618. Type-species, here designated, Fascellina specidaria Walker,

1866 : 1553.
Zanclalbara Inoue, 1962 : 27. Type-species Drepana scabiosa Butler, 1877 : 478, by monotypy.

syn. n.
Albara Walker sensn Gaede, 1931 : 31. [Partim.]

<$. Palp extends to just above labrum. Antenna bipectinate from base to between one
half and four-fifths of its length, except in fenestra which has uniserrate ciliate antenna.

Mesothoracic tibia with one pair of spurs, metathoracic tibiae with two pairs of spurs.
Prothoracic tibia with well-developed brush-organ in scabiosa, hoenei and fuscilinea.

Vein i?i in fore wing arises from distal end of cell ; i?2 arises from just distal to end of areole
in fenestra, otherwise from distal end of areole. In the hind wing Sc + i?i approximates to
Rs except in fenestra in which 5c -+- i?i anastomoses with Rs for some distance distal to end of
cell. Upper surface of wings buff, yellowish grey or yellowish brown ; sinuous antemedial
fascia simple or double ; discocellular cell-spot and posterior cell-spot usually present ; pale
patches at end of cell in some species ; postmedial fascia double, lunulate, sinuous or straight,
well-marked in most species ; subterminal fascia double, usually ill-defined except for short
distance just before apex. Under surface of wings yellow, buff or yellowish grey ; cell-spots
present or absent ; postmedial fascia well-marked in some species, hardly visible in others ;
subterminal variously marked.

o* genitalia : valve elongate in specidaria, vinacea and corticata, short in other species, with
variously shaped processes or setae at base ; socius well-developed ; diaphragma sclerotized
medially ; uncus simple in vinacea and corticata, absent in specularia, bifurcate to varying
extent in remaining species ; seventh abdominal sternum modified except in fenestra, asymmetric
in corticata francki ; eighth abdominal tergite emarginate or convex posteriorly ; eighth
sternite a narrow transverse plate in discispilaria, otherwise elongate, with lateral sclerite on
either side except in fenestra ; long eversible sac on either side of eighth sternite in corticata
and specularia.

$. As for male but antennae very weakly uniserrate (all except fenestra) or uniserrate and
cilate (fenestra), apex of fore wing more strongly produced, and prothoracic tibia without
brush-organ.

$ genitalia : signum ovate in fenestra, otherwise an elongate band ; eighth abdominal
segment and ninth tergum moderately well sclerotized.

Agnidra probably has closest affinities with Thymistida Walker and Betalbara
Matsumura. It can be separated from the latter by the coloration and colour-
pattern of the wings in both sexes and by the shape of the seventh and eighth
abdominal sternites in the male. (The seventh sternite is unmodified in fenestra.)
Agnidra is readily distinguished from Thymistida by the shape of the wings (see
Plates).

Ten species are now included in Agnidra and are dealt with below. Three of
these are new ; two have been transferred from Drepana Schrank {fenestra and
corticata), four from Albara Walker {discispilaria, fuscilinea, specidaria, vinacea),
and one from the monotypic Zanclalbara Inoue {scabiosa).

Distribution. Ceylon {specularia), N. India {specularia, corticata, vinacea,
discispilaria) , Sikkim {specularia) , Bhutan {specidaria) N. Burma {vinacea, specidaria,
fenestra), Thailand {discispilaria), Vietnam {specularia), China {fulvior,furva, hoenei,
corticata, fenestra, scabiosa), Korea {scabiosa), Japan {scabiosa) and Malaya {fuscilinea).

3 o A. WATSON

Seven of the ten species of Agnidra are endemic to the Indo-Chinese Subregion,
specularia is shared between the Indo-Chinese and Indian Subregions, scabiosa is
shared between the Indo-Chinese Subregion and the Manchurian Subregion, and
fuscilinea is endemic to the Malayan Subregion. (See Table i.)

Certain tentative species groupings can be made in this genus : specularia,
vinacea and corticata form one group ; scabiosa, fuscilinea, fulvior, furva, hoenei, and
probably discispilaria, a second group. The latter could be subdivided by ex-
tracting fulvior, furva and hoenei as one unit, discispilaria as a second, and scabiosa
and fuscilinea as a third. On the evidence of the total external and genitalic
characters studied, the species fenestra is probably most satisfactorily placed in
Agnidra in spite of the venational and antennal difference between it and the rest
of the genus.

Key to the species of AGNIDRA
Males

Antenna uniserrate ; postmedial fascia on upper surface of fore and hind wing

straight (PI. 2, fig. 307) ; Sc + 7?i anastomosed with Rs distal to cell in hind wing ;

7th sternum of abdomen unmodified ...... fenestra (p. 34)

Antenna bipectinate ; postmedial fascia on upper surface of fore and hind wings

straight, lunulate or sinuous ; Sc + Ri approximated to Rs distal to cell in hind

wing ; 7th sternum of abdomen modified ....... 2

Postmedial fascia on upper surface of fore wing straight or nearly so (e.g. PI. 1, fig. 305) 3

Postmedial fascia of upper surface of fore wing lunulate or sinous, not straight . . 5

Prothoracic femur with large brush organ. Genitalia : valve not elongate : uncus

bifid ........... fuscilinea (p. 44)

Prothoracic femur without large brush organ. Genitalia : valve elongate ; uncus

simple ............. 4

Area of pale medial patches on fore wing as in PI. 1, figs. 303, 304. Genitalia :

gnathus not heavily spinose (Text-fig. 50) .... corticata (p. 39)

Area of pale medial patches on fore wing as in PI. 1, fig. 305). Genitalia : gnathus

heavily spinose (Text-fig. 52) ....... vinacea (p. 40)

Area of pale patches on hind wing as in (PI. 2, fig. 306). Genitalia : uncus absent

(Text-fig. 41) ......... specularia (p. 36)

Area of pale patches small (e.g. PI. 2, fig. 309). Genitalia : uncus present . . 6

Pale patches on each wing encircled by ring of dark scales (PI. 2, fig. 308)

discispilaria (p. 44)
Pale patches on each wing not encircled by ring of dark scales .... 7

Transverse fasciae lunulate on both wings ........ 8

Transverse fasciae non-lunulate on both wings. (PI. 1, figs. 301, 302) scabiosa (p. 42)

Prothoracic femur with well-developed brush-organ. Genitalia as in Text-figs. 68-71

hoenei (p. 31)
Prothoracic femur without brush-organ. Genitalia not as in hoenei ... 9

Fore wing moderately falcate (PI. 2, fig. 311) ; ground-colour of upper surface

brownish buff. Genitalia as in Text-figs. 30-33 .... furva (p. 33)

Fore wing weakly falcate (PI. 2, fig. 310) ; ground-colour of upper surface buff.

Genitalia as in Text-figs. 25-28 ....... fulvior (p. 31)

CHINESE DREPANINAE 31

Agnidra hoenei sp. n.

(PI. 2, fig. 309 ; Text-figs. 29, 68-71)

o". Vertex of head and base of antenna dark reddish brown ; rest of antenna, front of head
and outer surface of palp brownish buff ; antenna bipectinate from base to about four-fifths
of its length. Collar yellow.

Thorax and abdomen similar in colour to corresponding surface of wing. Colour-pattern
of wings as in PI. 2, fig. 309. Ground-colour of upper surface of fore wing buff, variable in
tone, moderately lustrous ; pale whitish medial patches sometimes strongly irrorate with dark
brown ; remaining markings pale purplish brown, except for dark brown edge to medial patches
and dark brown anterior markings of subterminal fascia. Ground-colour of hind wing usually
slightly paler than fore wing, moderately lustrous ; markings pale purplish brown, except for
patch at end of cell (as for fore wing).

Under surface of both wings brownish yellow-orange, slightly lustrous. Both wings with
brownish grey subterminal and postmedial fasciae anteriorly, or in some specimens (e.g. holotype)
with subterminal moderately well-marked anteriorly but with only a trace of postmedial
fascia. Well-marked dark brown discocellular spot on fore wing and smaller faintly marked
spot at posterodistal angle of cell ; hind wing with similarly placed but poorly defined cell-
spots.

Prothoracic leg with brush-organ ; outer surface of femur, tibia and tarsus dark greyish
brown ; legs otherwise as for colour of under surface of wings. Mesothoracic tibia with fringe
of long hair-scales on inner surface.

o* genitalia as in Text-figs. 68-71.

$. Similar to male but with ciliate, very weakly biserrate antennae, and prothoracic and
mesothoracic legs with normal vestiture of scales.

$ genitalia as in Text-fig. 29.

Wing, o* 17-5-20-0 mm. (14) ; $ 19-5-21-0 mm. (14).

Closely related to furva and fulvior but separated from both by the genitalia
(especially the aedeagus) and the presence of a brush-organ on the male fore leg,
and from furva by the coloration, colour-pattern of both wings and the less strongly
falcate fore wing.

Distribution. Known only from the type-locality, which is also the only known
locality for its closest relative fulvior.

Holotype <$. China : N. Yunnan, Likiang, 8.vi.i934 {Hone) ; Drepanidae
genitalia slide No. 968. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China: 14^ and 14 $, N. Yunnan, Likiang,
8.vi.34, 2.V-3.X.1935 {Hone). Daniel Collection, Munich. China : 2 $, 1 $, N.
Yunnan, Likiang, 6.vi, 28.ix. 1934, 5. ix. 1935 {Hone). BM{NH). China; 3 <J, 1 $,
N. Yunnan, Likiang, Lvii-7.ix.1934, 19JX-3.X.1935 {Hone).

Agnidra fulvior sp. n.

(PI. 2, fig. 310 ; Text-figs. 25-28, 34)

o". Whole of head and outer surface of palp dark reddish brown. Antenna greyish brown,
darkest at base. Bipectinate from base to four-fifths of its length. Collar light buff.

Thorax and abdomen reddish buff dorsally, pale buff ventrally.

Colour-pattern of upper surface of wings as in PI. 2, fig. 310 ; coloration as for hoenei but
with ground-colour of both wings usually duller, more brownish, and with medial patches of

A. WATSON

Figs. 25-29. Agnidra, genitalia. 25-28, fulvior, $. 25, seventh sternite ; 26, eighth
tergite and sternite ; 27, $ ; 28, aedeagus. 29, hoenei, $.

CHINESE DREPANINAE 33

fore wing invariably irrorate with brown. Under surface of both wings dull brownish yellow
with only cell-spots well-marked ; fasciae similar to hoenei, but diffusely marked. Outer
surface of fore leg dark brown ; legs otherwise pale buff.

cj genitalia as in Text-figs. 34a-34d.

$. Similar to male but with very weakly biserrate, ciliate antenna.

$ genitalia as in Text-fig. 34.

Wing, o* 19-5-20-5 mm. (7) ; ? 20-5 (1).

Readily distinguished from the closely related furva by the colour-pattern and by
the less strongly falcate fore wings ; from hoenei, probably its closest ally, by the
lack of a brush-organ on the male fore leg ; and from both species by differences in
the male genitalia (especially in the aedeagus).

Distribution. Known only from the type locality (China, Yunnan).

Holotype <$. China : N. Yunnan, Likiang, 22. vi. 1935, (Hone) ; Drepanidae
genitalia slide No. 964. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 3 $, 2 $, N. Yunnan, Likiang,
22.vi-9.viii.1935 (Hone). Daniel Collection, Munich. China. : 1 <$, N. Yunnan,
Likiang, 3.ix.i935 (Hone). BM(NH). China : 2 <$, N. Yunnan, Likiang, 30.VU-
7.viii.i935 (Hone).

Agnidra furva sp. n.

(PI. 2, fig. 311 ; Text-figs. 30-33)

o*. Head and outer surface of palp greyish brown. Antenna greyish brown at base, paler
distally ; bipectinate from base to four-fifths of its length. Collar brownish white.

Thorax brownish buff dorsally, buff ventrally. Colour-pattern of upper surface of wings as
in PL 2, fig. 311. Ground-colour of upper surface of fore wing brownish buff ; fasciae greyish
brown, anterior part of subtcrminal fascia very dark brown ; pale brown medial patches
irrorate with greyish brown. Hind wing slightly more yellowish buff on upper surface but
brownish buff at base and distal to subterminal fascia, colour of markings as on fore wing.
Under surface of both wings buff, but greyish brown medially and distally in fore wing and
antero-distally in hind wing. Under surface of fore wing with moderately well-marked, greyish
brown, double postmedial fascia and trace of subterminal fascia ; well marked, dark brown
discocellular spot and similar but smaller spot at posterior angle of cell. Under surface of
hind wing similar to fore wing but with discocellular spot smaller than posterior cell-spot.
Legs buff but with outer surface of fore leg greyish brown.

Abdomen greyish buff dorsally, paler posteriorly ; buff ventrally.

o* genitalia as in Text-figs. 30-33.

$. Not known.

Wing. 6* 37 -0-38 -o mm. (3).

Separated from the closely related hoenei and fulvior by the more strongly falcate
fore wings, the brownish buff ground-colour of the wings, the more strongly marked
postmedial fascia on the upper surface of both wings and by the male genitalia
(particularly the aedeagus).

Holotype $. China : Tsekou, 1900 (Dubernard) ; Drepanidae genitalia slide
No. 966. In the BM(NH).

Paratypes. BM(NH). China: 2 (J, N. Yunnan, Tsekou, 1898-1900 (Dubernard).

34

A. WATSON

Agnidra fenestra (Leech) comb. n.
(PI. 2, fig. 307 ; Text-figs. 35-38)

D rep ana fenestra Leech, 1898 : 368.

Drepana fenestra Leech ; Strand, 191 1 : 202. [Good fig.]

Drepana fenestra Leech ; Gaede, 1931 : 26.

In contrast to each of the remaining species of the genus, fenestra has uniserrate
antennae in both sexes, R% of the fore wing arises from just distal to the end of the
areole, and in the hind wing Sc + R\ anastomoses with Rs for some distance distal
to the end of the cell. The male genitalia differ in possessing an unmodified seventh

Figs. 30-34. Agnidra, genitalia. 30-33, furva. 30, q" eighth sternite
seventh sternite ; 33, aedeagus. 34, fulvior, $.

31. 6" ; 32, 6*

CHINESE DREPANINAE

35

abdominal sternum. In coloration and colour-pattern fenestra resembles a small
corticata or vinacea, but is easily distinguished by the translucent patches at the
distal end of the cell and by the well-defined confluent postmedial fascia on the
upper surface of the fore and hind wing.

Wing. <$ 12-0-14-0 mm. (10) ; $ 13-0-15-5 mm. (6).

Distribution. N. E. Burma, China (Szechwan, Yunnan, Shensi).

Figs. 35-38. Agnidra fenestra, genitalia. 35, $ ; 36, <$ eighth tergite and sternite ; 37,

aedeagus ; 38, $.

36 A. WATSON

Material examined. Type. I select as LECTOTYPE a <$ from the syntypic
series of two males and one female in the BM(NH), labelled : Wa-Shan [China,
Szechwan], 6000 ft. A. E. Pratt coll. May 1889 ; Leech Coll. 1900-64 ; Drepana
fenestra sp. n. Type $ ; Drepanidae genitalia slide No. 803.

Paralectotypes. BM(NH). China : 1 <J, 1 $, [Szechwan], Wa-shan, 6000 ft.,
v.1889 (Pratt).

Other material. BM(NH). China: i <$ [Szechwan], Frontiere orientale, Tibet,
1905 (Dejean) ; 1 <$, S. Shensi, Tsinling, Tapaishan, 23. vi. 1935 (Hone) ; 1 $, N.
Yunnan, Likiang, 2.vii.i934 (Hone). Burma : 1 <$, Kambaiti, 7000 ft., 15. vi. 1934
(Hone). Naturhistorisch.es Museum, Vienna. China: i $, Szechwan, Ta-tsien-lou,
1910 (Chasseurs indigenes). Museum Koenig, Bonn. China : 7 ex., N. Yunnan,
Likiang (Hone) ; 2 ex., N. Yunnan, A-tun-tse (Hone) ; 13 ex., S. Shensi, Tsinling,
Tapaishan (Hone). U.S. National Museum. China : 1 $, Szechwan, Beh Luh
Din (30 miles N. of Chengtu) (Graham).

Agnidra specularia (Walker) comb. rev.

(PI. 2, fig. 306 ; Text-figs. 39-43)

Fascellina specularia Walker, 1866 : 1553.
Agnidra specularia (Walker) Moore, [1868] : 618. [Fig.]
Agnidra specularia (Walker) ; Butler, 1886 : 17. [Figs.]
Drepana specularia (Walker) Hampson, [1893] : 335.
Drepana specularia (Walker) ; Strand, 191 1 : 202. [Fig.]
Albara specularia (Walker) Swinhoe, 1892 : 242.
Albara specularia (Walker) ; Gaede, 193 1 : 33.
Albara ochracina Bryk, 1943 : 17. [Fig.] syn. n.

Distinguished from the rest of the genus, including its close allies vinacea and
corticata, by the large sparsely scaled patches on both wings (see Plate), and in the
male genitalia (Text-figs. 40-43) by the absence of an uncus, the shape of the eighth
sternite and the presence of a robust two-spined process at the base of the valve.

Wing. $ 19-0-22-5 mm. (33) ; $ 24-0-25-5 mm. (10).

This species, as in scabiosa, hoenei and fuscilinea, has a brush-organ on the femur
of the prothoracic leg in the male.

Distribution. N.E. India (Darjeeling, Khasia Hills), Sikkim, Bhutan, Ceylon and
Vietnam.

Material examined. Types, specularia. Walker (1866 : 1553) described this
species from a single male specimen from ' North Hindostan ' in the collection of
A. E. Russell. The Russell collection is stated to be lost by Horn and Kahle
( I 937 : 38°) an d no trace of this collection has been found by the present author.
I therefore select as NEOTYPE a male in the BM(NH) labelled : Darjeeling,
4 August, 1886. H. J. Elwes ; Coll. H. J. Elwes ; Rothschild Bequest 1939-1.

ochracina. Holotype <$, N. E. Burma, Kambaiti, 2000 m. ; Drepanidae genitalia
slide No. 792 ; in the Naturhistoriska Riksmuseet, Stockholm.

Other material. BM(NH). India : 8 $, 4 $, Darjeeling (Elwes, M oiler, Pilcher) ;

CHINESE DREPANINAE

37

Figs. 39-43. Agnidra speciriaria, genitalia. 39, $ ; 40, aedeagus ; 41, $ ; 42, £ seventh
sternite ; 43, $ eighth tergite, and eighth sternite showing right lateral sac.

3»

A. WATSON

4 <J, i $, Khasis ; I $, Assam, Cherrapunji. Sikkim : II <J, 2 $, 7000-10,000 ft.,
viii.1886, 23.iv-ix.1889, viii.1909 (M oiler, Elwes, Pilcher). Bhutan : 1 <J, 2 $.
Ceylon : 3 $, 1 $, Haputale. Daniel Collection, Munich. There is a single male
from Vietnam (Tonking) which may prove to represent a new subspecies of specularia.

44

Figs. 44-48. Agnidra corticata corticata, genitalia. 44, $ ; 45, ^ ; 46, aedeagus ; 47, $
seventh sternite ; 48, £ eighth tergite and sternite.

CHINESE DREPANINAE 39

Agnidra corticata (Warren) comb. n.
(PI. 1, figs. 303, 304 ; Text-figs. 44-51)
Drepana corticata Warren, 1922 : 464. [Good fig.]

Separable from the closely allied vinacea, and from fuscilinea which it resembles
externally, by the much smaller area of pale patches at the end of the cell on the
fore wing. These patches are pale buff; not sparsely scaled as in specularia, another
closely allied species. The male and female genitalia are also diagnostic, particularly
the short uncus.

Two subspecies are known : the nominate subspecies (N.E. India), and francki
(China).

Agnidra corticata corticata (Warren)
(PI. i, fig. 304 ; Text-figs. 44-48)
Drepana corticata Warren ; Gaede, 1931 : 26.

49

51

Figs. 49-51. Agnidra corticata francki, 0* genitalia ; 49, seventh sternite ; 50, o* ) 51.
eighth tergite, and eighth sternite showing right lateral sac.

4 o A. WATSON

Distinguished from francki by difference in the shape of the pale patches on the
upper surface of the fore and hind wings (see Plate) and by the male genitalia, in
particular the seventh and eighth abdominal tergites.

Wing. <J 15-5 mm. (1) ; $ 17-5 mm. (1).

The specimen illustrated in PI. 1, fig. 304 (a female from the Khasia Hills in
Assam) is the only known specimen apart from the holotype.

Material examined. Type. Holotype <$, Darjeeling, (M oiler) ; Drepanidae
genitalia slide No. 738. In the BM(NH).

Other material. BM(NH). India : 1 $, Khasis, vi.1895 (Nat. Coll.).

Agnidra corticata francki ssp. n.

(PI. i, fig. 303 ; Text-figs. 49-51)

Similar to the nominate subspecies in both sexes, but with the pale medial patches on the
fore wing only faintly marked and the medial shade on the hind wing narrower. The male
genitalia (Text-figs. 49-51) differ from those of the nominate subspecies, particularly in the
shape of the seventh and eighth abdominal sternites.

Wing. (J 18-5 mm. (1) ; $ 19-0 mm. (1).

Holotype $. China : Kwanhsien, 10.vii.1926 (Franck) ; Drepanidae genitalia
slide No. 799. In the BM(NH).

Paratype. BM(NH). China : 1 $, Szechwan, Kwanhsien, 24.vii.1926 (Franck).

Agnidra vinacea (Moore) comb. n.
(PI. 1, fig. 305 ; Text-figs. 52-56)

Drepana vinacea Moore ; 1879 : 85.

Albara vinacea (Moore) Warren, 1922 : 468. [Good fig.]

Albara vinacea (Moore) ; Watson, 1961 : 326.

Albara birmanica Bryk, 1943 : 18. [Good fig.] [Synonymized by Watson, 1961 : 326.]

Distinguished from fuscilinea, which vinacea most closely approaches externally,
by the colour-pattern (see Watson, 1961 : 326) and by the male genitalia (particu-
larly the heavily spinose gnathus). The species specularia and corticata are also
close allies of vinacea but are easily distinguished from it by differences in the
colour-pattern (see fig. in Warren, 1922) and by the genitalia of both sexes (Text-
figs. 52-56).

Wing. £ 17-5-21-0 mm. (27) ; $ 19-5-23-5 mm - ( I0 )-

Distribution. Sikkim, N.E. India, N.E. Burma.

Material examined. Types, vinacea. A lectotype was selected by the present
author (Watson, 1961) from syntypic material (two examples in the BM(NH) and
four examples in the Zool. Museum, Berlin). Lectotype <J, Darjeeling, 1864 ;
Drepanidae genitalia slide No. 724 ; in the BM(NH).

birmanica. Holotype <$, N.E. Burma, Kambaiti, 2000 m. ; Drepanidae genitalia
slide No. 725 ; in the Naturhistoriska Riksmuseet, Stockholm.

CHINESE DREPAXIXAK

4i

Figs. 52-56. Agnidra vinacea, genitalia. 52, <$ ; 53, aedeagus ; 54, $ eighth tergite and
sternite showing right lateral sac ; 55, <$ seventh sternite ; 56, $.

42 A. WATSON

Other material. BM(NH). India : 5 <J, 3 $, Darjeeling, 20.vii.1886, v,vi.i88o.
(Moller, Elwes) ; 3 <$, 1 $, Darjeeling, Gopaldhara, vi.1918, 3400-5800 ft. (Stevens,
Nat. Coll.) ; 10 ^, 3 $, Khasis, xi.1894, iv., vi.1895 ; 2^, Naga Hills, 5000-8000 ft.,
vii,viii.i889 (Doherty). Sikkim : 2 <$, 1 $, Pedong (Desgodins) ; 4 cJ, 3 ?, 7000 ft.,
20.xi.1889, vii,ix.i909 (Moller, Pilcher). Burma : 2 <J, N.E. Burma, Kambaiti,
7000 ft., i.iv., 15.V.1934 (Malaise).

Agnidra scabiosa (Butler) comb. n.

(PI. 1, figs. 301, 302 ; Text-figs. 57-63)

Drepana scabiosa Butler, 1877 : 478.

Drepana scabiosa Butler ; Strand, 191 1 : 20.

Drepana scabiosa Butler ; Gaede, 1931 : 27.

Albara scabiosa (Butler) Bryk, 1949 : 27.

Albara scabiosa (Butler) ; Inoue, 1956 : 369.

Zanclalbara scabiosa (Butler) Inoue, 1962 : 27. [Good figs.]

Distinguished from fuscilinea, probably its closest ally, and from the rest of the
genus by the distinctive colour-pattern of the wings (see Inoue, 1962) and in the
male genitalia by the shape of the gnathus, socii and uncus.

Two subspecies are known : the nominate subspecies (S.E. Russia and Japan)
and fixseni (China and Korea).

Agnidra scabiosa scabiosa (Butler)
(PI. 1, fig. 301 ; Text-figs. 57-59)

Albara scabiosa (Butler) ; Inoue, 1956 : 368. [Partim.] [Good figs.]

Separated from fixseni by the narrow, usually poorly marked, proximal sub-
terminal line and by the male genitalia (see fixseni).
Wing. <$ 14-0-17-5 mm. (40) ; $ 17-5-20-0 mm. (3).
Distribution. Japan (see Inoue, 1956, 1962) and S.E. Russia.

Type. I select as LECTOTYPE a $ syntype labelled : 77.9 Japan ; Drepana
scabiosa Butler Type. This is one of a pair of syntypes, which according to the
registration details recorded in the Department of Entomology, BM(NH) (Registra-
tion No. 1877-9), were collected by F. M. Jonas in Yokohama (Japan). The type-
locality given by Butler in the original description is ' Yokohoma (Jonas) ', which
confirms the supposition that the present lectotype and paralectotype (labelled
77.9 Japan) form part or whole of Butler's original material.

Agnidra scabiosa fixseni (Bryk) ssp. rev., comb. n.
(PI. 1, fig. 302 ; Text-figs. 60-63)

Albara scabiosa fixseni Bryk, 1949 : 27.

Albara scabiosa (Butler) ; Inoue, 1956 : 368. [Partim.] [Synonymy of fixseni with scabiosa.]

Drepana scabiosa (Butler); Fixsen, 1887 : 347.

Most specimens of fixseni can be distinguished from the nominate subspecies by

CHINESE DREPANINAE

43

Figs. 57-59. Agnidra scabiosa scabiosa, <$ genitalia. 57, <$ ; 58, aedeagus ; 59, seventh
and eighth sternites, and eighth tergite.

44 A. WATSON

the broad, strongly marked, proximal subterminal line between Cui a and M% on
the fore wing. In the male genitalia the characteristic shape of the seventh ab-
dominal sternite, which can be seen without dissection, and structural differences in
the diaphragma, eighth sternite and aedeagus (Text-figs. 61-63) separate the two
subspecies.

Wing. <J 15-5-17-0 mm. (10) : $ 17-5-19-0 mm. (5).

Distribution. Korea, China (Manchuria, Chekiang, Hunan, Hupeh, Kiangsu).

Material examined. Type. Holotype <$, Korea, Kariuzawa ; Drepanidae
genitalia slide No. 719. In the Naturhistoriska Riksmuseet, Stockholm.

Other material. BM(NH). Korea: 2 <$, Gensan, vii. 1887 {Leech). 7 g, Seoul,
Chungyangri, 13. vii. 1956, 8.v.-i2.viii.i96o (Pak) ; 1 $, Ori Dong, 1-7. ix. 1953
(Thompson). China : 1^,1$, Chekiang, West Tien-mu-shan, 1600 m., 3. vii. 1932,
28. ix. 1933 (Hone) ; 1 g, Chekiang, Mokanshan, 28.viii.1930 (Hone) ; 3 ^, 1 $,
[Hupeh], Changyang, vi,viii.i888 (Pratt) ; 1 <$, [Kiangsu], Nanking, Lungtan,
Berg Pao-Hwa, iv ; 2 <$, 1 $, Hunan, Hoeng-shan, 900 m., 9 # v.-26.vi.i933 (Hone).
Museum Koenig, Bonn. China : 6 ex., Hunan (Hone) ; 21 ex., Chekiang (Hone).

Agnidra fuscilinea (Watson) comb. n.
Albara fuscilinea Watson, 1961 : 326. [Figs., including genitalia.]

The male genitalia of this species (see Watson, 1961) indicate possible close
relationships with scabiosa. It is easily separable from scabiosa by the colour-
pattern of the wings and by several differences in the male genitalia. Externally,
fuscilinea is closest to vinacea, from which it differs by the dark-edged patches on
the upper surface of the wings and by the presence on the under surface of the
hind wing of a conspicuous, dark brown postmedial fascia and a large brown patch
at the distal end of the cell.

Wing. <J 20-5-21-0 mm. (2).

Distribution. Malaya, and probably Sumatra (1 $ in the Ent. Lab., Wageningen).

Material examined. Type. Holotype^, Malaya, Selangor, Bukit Kutu, 8,500 ft.,
22.iii.1931 (Pendlebury). Drepanidae genitalia slide No. 891. In the BM(NH).

Other material. (See Watson, 1961.)

Agnidra discispilaria Moore comb. rev.
(PI. 2, fig. 308 ; Text-figs. 64-67)
Agnidra discispilaria Moore, 1867 : 619.

Albara discispilaria (Moore)
Albara discispilaria (Moore)
Albara discispilaria (Moore)

Swinhoe, 1892 : 242.

Warren, 1922 : 467. [Fig. (as ' discipilavia ').]

Gaede, 1931 : 31.
Drepana discispilaria (Moore) Hampson, [1893] : 336.

Agnidra usta Butler, 1886 : 17. [Synonymized with discispilaria by Warren, 1922 : 467.]
Albara magnidiscata Warren, 1922 [German edition] : 468. [Fig.] [Selected by Gaede,
1931 : 32, from a multiple original spelling.] syn. n.

CHINESE DREPANINAE

45

Figs. 60-63. Agnidra scabiosa fixseni, genitalia. 60, $ ; 6i, <$ ; 62, aedeagus ; 63, <J
seventh and eighth sternite, and eighth tergite.

46 A. WATSON

Albara ' magnadiscata ' ; Warren, 1922 [English edition] : 468. [Incorrect original spelling

(see above).]
Albara discispilaria macularis Bryk, 1943 : 18. [Good fig.] syn. n.

The affinities of this species are doubtful, but similarities in the male genitalia
suggest that scabiosa and fuscilinea are its closest allies. The colour-pattern of
both sexes and the extremely large gnathus in the male genitalia are highly diagnostic.

Seven of the twenty-five examined specimens of this species match the lectotype
of magnidiscata and the holotype of macularis in that the dark-edged grey patch at
the distal end of the cell on the fore wing is distinctly larger (about 2 mm. in diameter)
than in the neotype of discispilaria (just over 1 mm. in diameter). Where pin-
label data was available, it was found that the large-spotted specimens (these are
also lighter in coloration) were taken in April, whereas the small-spotted specimens
were taken in June, July or August.

Wing. ^ 19-0-23-0 mm. (14) ; <j> 21-5-22-5 mm. (6).

Distribution. N.E. India, Sikkim, and Thailand (1 $ in the B.M.(NH)).

Material examined. Types, discispilaria. The male type material of disci-
spilaria cited originally as from ' Bengal, in Coll. A. E. Russell ' is apparently lost,
together with the rest of the Russell collection (see Horn and Kahle, 1937 : 380).
No trace of this material can be found in the BM(NH) nor in several other European
Museums which have been consulted. I therefore select as NEOTYPE a $ in the
BM(NH) labelled : 7.86 Darjeeling. H. J. E. ; Coll. H. J. Elwes ; Albara
discispilaria <$ Moore ; Rothschild Bequest B. M. 1939-1.

usta. LECTOTYPE $, in the BM(NH), here selected labelled : Darjiling 79.57
[ex Lidderdale coll.] ; Agnidra usta Butler.

magnidiscata. LECTOTYPE <$, in the BM(NH), here selected, labelled :
Darjeeling (Pilcher) ; 18.3.89 ; Albara magnidiscata Type J Warr. [in Warren's
handwriting] ; Rothschild Bequest B. M. 1939-1 ; Drepanidae genitalia slide No. 728.

macularis. Holotype <$> N.E. Burma, Kambaiti, 7000 ft. ; Drepanidae genitalia
slide No. 727 ; in the Naturhistoriska Riksmuseet, Stockholm.

Other material. BM(NH). India : 4 <$, 1 $, Darjeeling (Moller, Lidderdale) ;
1 (J, 2 $, Darjeeling, Gopaldhara, 4720 ft., 3440-5800 ft., vii.1918 (Stevens) ; 1 $,
Khasis, ii.1896 (Nat. Coll.). Sikkim : 10 <$, 6 $, 6.vi.i888, 14.iv.-14.viii. 1889,
vi-viii.1909 (Pilcher, Moller). Thailand : 1 $, Chiengmai Mt., 5800 ft., 19.iii.1928
(McKean).

BET ALBARA Matsumura
(Pis. 2, 3, figs. 312-319 ; Text-figs. 72-117)

Betalbara Matsumura, 1927 : 47. Type-species Drepana manleyi Leech, 1898 : 366, by original

designation.
Betalbara Matsumura ; Inoue, 1962 : 23.
Microblepsis Warren, 1922 : 461. Type-species Problepsis cupreogrisea Hampson, 1895 : 288,

by monotypy. syn. n.

<J. Palp extends to just above labrum. Antenna strongly bipectinate in most species but
weakly bipectinate in acuminata and robusta and lamellate in manleyi.

CHINESE DREPANINAE

" 70

Figs. 64-71. Agnidra, genitalia. 64-67, discispilaria. 64, <$ eighth tergite and sternite ;
65, aedeagus ; 66, £ ; 67, $. 68-71, hoenei, (J. 68, seventh sternite ; 69, aedeagus ;
70, eighth tergite and sternite ; 71, <$.

48 A. WATSON

Thorax and abdomen similar in colour to adjacent surface of wing. Upper surface of wings
pale buff in manleyi, yellowish white in acuminata and grey or brownish grey in remaining
species. i?i in fore wing arises from areole in prunicolor, from near end of cell in rest of genus ;
Rs arises from areole. Antemedial fascia of fore wing absent only in leucosticta, postmedial
fascia oblique, strongly marked ; subterminal fasciae absent in violacea, broad and diffusely
marked in robusta, narrow and weakly marked in rest of genus. Sc + i?i approximates to Rs
distal to end of cell in hind wing. Antemedial fascia and strongly marked postmedial fascia
present on hind wing except in robusta ; weakly marked subterminal fascia present. Under
surface of wings dull white in acuminata, pale brownish yellow in manleyi, buff in robusta, and
yellow, brownish grey or neutral grey in remaining species. Under surface of both wings
unmarked in robusta ; with postmedial and subterminal fasciae in manleyi and acuminata,
but with only diffusely marked postmedial fascia in the other five species. Mid tibia with one
pair of spurs in violacea, but with two pairs in rest of genus. Hind tibia with two pairs of spurs.

o* genitalia : saccus with medial dorsal process in rectilinea and rugosa and with long lateral
processes in flavilinea ; valve small, with processes at base ; socius simple or with processes ;
diaphragma with medial sclerotization, best developed in flavilinea ; uncus bifurcate in manleyi
and acuminata, absent in violacea, bifid in rest of genus ; aedeagus variously shaped and
ornamented ; seventh abdominal sternite with two lateral anterior apodemes in violacea,
otherwise with single medial apodeme ; eighth tergite truncate or concave posteriorly ; eighth
sternite variously shaped, with lateral sclerite on either side in violacea and robusta and with
pair of long eversible setose sacs in violacea.

9- As for male but with apex of fore wing slightly more strongly produced, and antennae
very weakly biserrate or uniserrate except in leucosticta which has weakly bipectinate antennae.

$ genitalia : signum an elongate band in robusta, absent in manleyi, flavilinea, cupreogrisea
and leucosticta, ovate in remaining species ; eighth and ninth segments variously sclerotized,
without processes.

Included in this revision are manleyi and acuminata, listed by Inoue (1962) ;
prunicolor, flavilinea, leucosticta and violacea, transferred from Albara Walker ;
robusta, transferred from Drepana Schrank, cupreogrisea transferred from Micro-
blepsis ; and two new species, rectilinea and rugosa. The classification of violacea
and, in particular, robusta in Betalbara is tentative, but I believe that there are
sufficient similarities between them and the rest of the genus to justify their inclusion.

Betalbara is probably most closely related to Albara. It can be distinguished
externally from Albara by the continuous subterminal fascia on the fore and hind
wing, or by its absence, whereas in Albara this fascia is represented by a series of
spots. In the male genitalia the uncus is not massive and bifid as in Albara, and
the seventh sternite is symmetrical in contrast with the strongly asymetric seventh
sternite of Albara. The females of Betalbara lack the flattened, paired, dorsal lobes
of the ninth segment found in Albara. The species of Pseudalbara differ in the
presence of a vestigial frenulum in the male and the absence of pattern on the upper
surface of the hind wing.

Distribution. N. India {prunicolor, rugosa, leucosticta, cupreogrisea, violacea) ;
Sikkim [prunicolor, leucosticta) ; Burma (prunicolor, cupreogrisea) ; China (manleyi,
acuminata, prunicolor , leucosticta, flavilinea, rectilinea, violacea, robusta) ; Formosa
(violacea) ; Japan (manleyi, acuminata) ; Malaya (leucosticta, rugosa). Six species
are Indo-Chinese endemics, two are shared between the Indo-Chinese and Malayan
Subregions and two between the Indo-Chinese and Manchurian Subregions (see
Table 1).

CHINESE DREPANINAE 49

Key to Species
Males

1 Ground-colour of upper surface of wings yellowish buff or white .... 2

- Ground-colour of upper surface of wings brown or grey ..... 3

2 Antenna bipectinate ; upper surface of fore wing with two conspicuous spots at distal

end of cell .......... acuminata (p. 49)

- Antennae weakly biserrate ; upper surface of fore wing with single dark marking at

distal end of cell ......... tnanleyi (p. 52)

3 Single well-marked transverse fascia present on hind wing ; outer margin of fore wing

strongly convex (PI. 3, fig. 319) ....... robusta (p. 65)

- At least two well-marked transverse fasciae present on hind wing ; outer margin of

fore wing straight or weakly convex ........ 4

4 Hind tibia with one pair of spurs. Postmedial fascia on fore wing nearly parallel to

outer margin of wing (PI. 3, fig. 318) ...... violacea (p. 62)

- Hind tibia with two pairs of spurs. Postmedial fascia on fore wing not nearly

parallel to outer margin of wing ......... 5

5 Antemedial fascia absent on upper surface of fore wing (PI. 2, fig. 314)

leucosticta (p. 55)

- Antemedial fascia present on upper surface of fore wing ..... 6

6 Broad, greyish white, transverse band present distal to postmedial fascia on hind

wing (PI. 3, fig. 316) ........ cupreogrisea (p. 58)

- Upper surface of hind wing without white band distal to postmedial fascia on hind

wing .............. 7

7 Antemedial fascia on fore wing acutely angled at middle (PI. 2, fig. 312)

prunicolor (p. 53)

- Antemedial fascia on fore wing not angled at middle ...... 8

8 Genitalia : saccus with long lateral process on either side, without dorsal process

(Text-fig. 89) ......... flavilinea (p. 55)

- Genitalia : saccus with single medial dorsal process, without lateral processes. . 9

9 Genitalia as in Text-figs. 102-104 ....... rectilinea (p. 61)

- Genitalia as in Text-figs. 1 09-1 11 ....... rugosa (p. 62)

Betalbara acuminata (Leech)
(Text-figs. 72-75)

Drepana acuminata Leech, 1890 : 113.

Drepana acuminata Leech ; Strand, 191 1 : 201.

Drepana acuminata Leech ; Gaede, 1931 : 25.

Platypteryx acuminata (Leech) Kirby, 1892 : 731.

Betalbara acuminata (Leech) Inoue, 1959 : 175. [Good fig.]

Betalbara acuminata (Leech) ; Inoue, 1962 : 24. [Good fig.]

Albara ogasawarae Matsumura, 1927 : 47. [Synonymy accepted from Inoue 1959 : 175.]

Albara ogasawarae Matsumura ; Gaede, 1931 : 32.

Albara acuminata ogasawarae Matsumura, Inoue, 1953 : 8.

Albara acuminata ogasawarae Matsumura, Inoue, 1956a : 663.

Drepana ida Bryk, 1942 : 27. [Synonymized with acuminata by Inoue, 1959 : 175.]

Albara ' ogasawarae ' ida (Bryk) Bryk, 1949 : 28.

Distinguished from its close relative manleyi by the male and female genitalia
(Text-figs. 72-75), the bipectinate antennae of the male, the more strongly falcate

50

A. WATSON

fore wing, the paler ground-colour of the wings, and by the presence on the fore
wing of two discocellular spots.

Wing. $ 18-0-21-0 mm. (6) ; $ 24-0 mm. (1).

Distribution. China (Hupeh, S. Shensi) and Japan (see Inoue, 1959, 1962).

Material examined. Types, acuminata. Holotype <$, Ichang [Hupeh], vii. 1888;
Drepanidae genitalia slide No. 735. In the BM(NH).

ogasawarae. Two syntypes, Japan, Honshu ; in the University of Hokkaido,
Japan. [Not seen.]

Figs. 72-75. Betalbara acuminata, genitalia. 72, aedeagus ; 73, $ eighth sternite and
apodemes of eighth tergite ; 74, o* ; 75. ?•

CHINESE DREPANINAE

5i

ida. Holotype <$, Japan, Karinzawa [Karuizawa] ; Drepanidae genitalia slide
No. 736 ; in the Naturhistoriska Riksmuseet, Stockholm.

Other material. Mttseum Koenig, Bonn. China : 2 ex., S. Shensi, Tapaishan
im Tsinling (Hone). BM(NH). China : 1 <?, S. Shensi, Tapaishan im Tsinling,
26. vi. 1935 (Hone).

Figs. 76, 77. Betalbara manleyi manleyi, genitalia. 76, <J ; 77, $.

52

A. WATSON

Betalbara manleyi (Leech)
(PI. 3, fig. 315 ; Text-figs. 76-80)

Drepana manleyi Leech, 1898 : 366.

Separated from the closely allied acuminata by the ciliate and unilamellate
antenna, the less strongly falcate fore wing, the darker ground-colour of the wings,
the presence of only one discocellular spot and by the genitalia of both sexes (Text-
figs. 76-80).

Two subspecies are known : the nominate subspecies (Japan) and prolatior
(China).

Betalbara manleyi manleyi (Leech)
(Text-figs. 76, 77)

Drepana manleyi Leech ; Strand, 191 1 : 201. [Fig.]
Albara manleyi (Leech) Nagano, 191 7 : 38 (English text).
Albara manleyi (Leech) ; Gaede, 1931 : 32.
Betalbara manleyi (Leech) Matsumara, 1927 : 47.
Betalbara manleyi (Leech) ; Inoue, 1953 : 8.

Figs. 78-80. Betalbara manleyi prolatior, q* genitalia. 78, o*

sternite ; 80, aedeagus.

79, eighth tergite and

CHINESE DREPANINAE 53

Betalbara manleyi (Leech) ; Inoue, 1956 : 369.

Betalbara manleyi (Leech) ; Inoue, 1959 : 175. [Good fig.]

Betalbara manleyi (Leech) ; Inoue, 1962 : 23. [Good figs.]

Separable from prolatior by the male genitalia (see prolatior).

Wing. <$ 14-5-17-0 mm. (7) ; $ 20-0 mm. (1).

Distribution. Japan (see papers above by Inoue, 1953-1962).

Type. I select as LECTOTYPE one of the two $ syntypes in the BM(NH)
labelled : Yokohama, Manley Coll ; Leech Coll. 1900-64 ; Drepana manleyi sp. n.,
Type <$ [probably in Leech's handwriting] ; Drepanidae genitalia slide No. 921.

Betalbara manleyi prolatior ssp. n.

(PI. 3, fig. 315 ; Text-figs. 78-80)

o*. Distinguished from the nominate subspecies by the male genitalia. The shape of the
gnathus and valve processes are diagnostic.
?. Unknown.
Wing, o* 16-5-17-0 mm. (6).

Holotype <$. China : Chekiang, West Tien-mu-Shan, 4.vi.i932 (Hone) ;
Drepanidae genitalia slide No. 920. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 3 <$, type-locality, 4.VL1932
(Hone). Daniel Collection, Munich. China: 3 <J, type-locality, 4. vi. 1932 (Hone).

Betalbara prunicolor (Moore) comb. n.

(PI. 2, fig. 312 ; Text-figs. 81-84)

Drepana prunicolor Moore, 1879 : 288.

A Ibara prunicolor (Moore) Warren, 1922 : 468. [Good fig.]

Albara prunicolor flavilinea Leech ; sensn Warren, 1922 : 469. [Good figs.]

A Ibara prunicolor (Moore) Gaede, 193 1 : 33.

Nordstroemia prunicolor (Moore) Bryk, 1943 : 14. [Partim.]

N ordstroemia prunicolor warreni Bryk, 1943 : 14. [Fig- ] syn. n.

This species is illustrated by Warren, (1922) both as prunicolor (49g) and as
flavilinea (49!, g), the latter having been misidentified by Warren. It is readily
separated from its nearest allies flavilinea and rectilinea by the sharply angled
antemedial fascia on the fore wing, by the fact that both R\ and R2 arise from the
areole, and by the genitalia of both sexes (Text-figs. 81-84). I n the male genitalia,
the conspicuous, serrate, lateral process of the aedeagus and the shape of the seventh
sternite are diagnostic.

Wing. <J 12-5-19-5 mm. (11) ; $ 16-5-17-5 mm. (3).

Distribution. N.E. India, Sikkim, N.E. Burma and China.

Material examined. Types, prunicolor. I select as LECTOTYPE a g syntype
from the Staudinger collection in the Zoological Museum, Berlin, labelled :
Darjeeling ; Drepanidae genitalia slide No. 1059.

54

A. WATSON

warreni. Holotype J, N.E. Burma, Kambaiti, 2000 m. Drepanidae genitalia
slide No. 230 ; in the Naturhistoriska Riksmuseet, Stockholm.

Other material. BM(NH). India : 12 ex., Assam, Khasia Hills. Sikkim : 1 <$,
1 $, 1887, 2.x. 1888. Daniel Collection, Munich. China : 1 <$, which may prove
to represent a new subspecies.

Figs. 81-84. Betalbara prunicolor, genitalia. 81, <$ ; 82, aedeagus ; 83, <$ seventh and
eighth sternites, and eighth tergite ; 84, $.

CHINESE DREPANINAE 55

Betalbara leucosticta (Hampson) comb. n.
(PI. 2, fig. 314 ; Text-figs. 85-88)

Drepana leucosticta Hampson, 1895 : 287.

Albara leucosticta (Hampson) Warren, 1922 : 469. [Fig. inaccurate, but useful guide.]

Albara leucosticta (Hampson) ; Gaede, 1931 : 32.

Probably quite closely related to prunicolor but distinguished by the colour-
pattern of both sexes (PI. 2, fig. 314), the bipectinate antennae of the female, the
shape of the male aedeagus and eighth abdominal tergite and sternite and by the
lack of a signum in the female genitalia.

Wing. $ 11 -5-14-0 mm. (8) ; $ 15-0-16-0 mm. (3).

Distribution. Sikkim, N.E. India, China and Malaya.

Material examined. Type. I select as LECTOTYPE a^ syntype in the BM(NH)
labelled : Sikkim, 8.7.91, G. C. Dudgeon, 94-52 ; Drepana leucosticta Hampsn.
type <$ ; Drepanidae genitalia slide No. 1600.

Other material. BM(NH). India : 7 <$, 3 $, Assam, Khasis, xi.1894, x.1895,
1906. Museum Koenig, Bonn. China : 1 <$, [Kwangtung], Canton. Two males
from Malaya in the BM(NH) may prove to represent a new subspecies.

Betalbara flavilinea (Leech) comb. n.
(PI. 2, fig. 313 ; Text-figs. 89-96)
Albara flavilinea Leech, 1890 : 113.

Distinguished from the closely related prunicolor by the dentate (not sharply
angled) postmedial fascia on the fore wing and by the fact that R\ arises from the
distal end of the cell. Separated from both prunicolor and rectilinea by the genitalia,
in particular by the presence of saccular processes in the male and the absence of a
signum in the female.

There are two subspecies, both at present known only from China. There is,
however, a single male, without an abdomen, from Formosa, in the BM(NH) which
will probably prove to represent this species.

Betalbara flavilinea flavilinea (Leech)
(PI. 2, fig. 313 ; Text-figs. 89-91)

Drepana flavilinea (Leech); Strand, 191 1 : 201 [Fig.]
Drepana flavilinea (Leech) ; Gaede, 1931 : 26.
Albara prunicolor flavilinea (Leech) Gaede, 1931 : 33.
Nordstroemia prunicolor flavilinea (Leech) Bryk, 1943 : 14.

In the fore wing of the illustration given by Strand (191 1) the postmedial fascia
should be only slightly arcuate posteriorly and more acutely reflexed costad.

56

A. WATSON

Figs. 85-

87

Betalbara leucosticta, genitalia. 85, $ ; 86, (J ; 87, aedeagus
and eighth sternites, and eighth tergite.

<$ seventh

CHINESE DREPAXINAE

57

Separable from shensiensis by the male genitalia.

Wing. $ ii -5-15-0 mm. (10) ; $ 16-0-18-0 mm. (3).

Bryk (1943) correctly pointed out that Warren, in Seitz, (1922 : 469) misidentified
as ' flavilinea ' specimens of prunicolor.

Distribution. China (Hupeh, Kiangsi, Chekiang).

Material examined. Type. Only one of the two original female syntypes can be
found in the BM(NH) : this specimen is here selected as LECTOTYPE : Chang
Yang, [Hupeh], July 1888 (A. E. Pratt Coll.) ; Drepanidae genitalia slide No. 796.

Other material. BM(NH). China : 2 $, same data as type, 3 <£, Chekiang, West
Tien-mu-Shan, 17.iv.-7.viii.1932 (Hone). Museum Koenig, Bonn. 11 ex., Chekiang,
Tien-mu-Shan (Hone) ; 1 ex., Kiangsi. There is a single male with an incomplete
abdomen, from the province of Fukien, in the Museum Koenig, Bonn, which may
also represent this species.

Figs. 89-91. Betalbara flavilinea flavilinea, <$ genitalia. 89, <J

sternite.

90, aedeagus ; 91, eighth

58 A. WATSON

Betalbara flavilinea shensiensis ssp. n.

(Text-figs. 92-96)

The male genitalia separate shensiensis from the nominate subspecies : nearly all the features
of the genitalia differ in their proportions from those of the latter.
Wing, o* i4-5-i7'5 mm. (20) ; ? 16-0-18-5 mm. (6).

Holotype <$. Sued-Shensi, Tapaishan in Tsinling, c. 1700 m., 12.V.1936 (Hone) ;
Drepanidae genitalia slide No. 789. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 19 $, 4 $, S. Shensi, Tapaishan
im Tsinling, c. 1700 m., 12.v-12.ix.1936 (Hone). BM(NH). China : 1 <$, 1 ?,
S. Shensi, Tapaishan im Tsinling, 31.viii.1935, 20.V.1936 (Hone).

Betalbara cupreogrisea (Hampson) comb. n.

(PI. 3, fig. 316 ; Text-figs. 97 ; and 98-100 of the possible £ of this species)

Problepsidis cupreogrisea Hampson, 1895 : 287.

Microblepsis cupreogrisea (Hampson) Warren, 1922 : 461. [Fig.]

Microblepsis cupreogrisea (Hampson) ; Gaede, 1931 : 16.

$. Head and palps dark brown. Antenna very weakly biserrate.

Colour-pattern of wings as in PI. 3, fig. 316, dark areas brown ; pale areas yellowish brown
on fore wing, brownish white on hind wing ; subterminal fascia of fore wing white ; apex and
anterior half of outer margin of fore wing suffused with yellowish buff. Under surface of
both wings pale brown suffused with buff at apex of fore wing ; fore wing with weakly marked
buff subterminal fascia anteriorly, hind wing with broad diffusely marked antemedial fascia
and postmedial fascia.

Fore wing strongly covex at M3, hind wing angled at M3.

Legs nearly white, but with front surface of fore leg brown.

? genitalia as in Text-fig. 97. Corpus bursae without signum.

o* (tentative identification, see Distribution). As for 9 but with bipectinate antennae.

o* genitalia as in Text-figs. 98-100. Medial diaphragmal sclerotization with dorsal diverti-
culum.

Wing. $ 12-5 mm. (1).

Possibly most closely allied to flavilinea Leech, but separated by a larger taxonomic
gap than that between flavilinea and prunicolor Moore.

Distribution. This species is known with certainty only from the type-locality
in south Burma. Two males from the Khasis (N.E. India) in the BM(NH) do not
exactly match the lectotype in wing shape or colour-pattern and may represent a
new species or a different closely related species. Material of both sexes from both
India and Burma is needed before this problem can be solved.

Material examined. Type. LECTOTYPE $, here selected, labelled : Tenas-
serim Valley, E. of Tavoy, Burmah, Doherty ; Problepsidis cupreogrisea $, Hmpsn.
M.S. type ; Joicey Coll. Brit. Mus. 1925-157 ; Drepanidae genitalia slide No. 72.
IntheBM(NH).

Other material. (See Distribution.)

CHINESE DREPANINAE

59

Figs. 92-96. Betalbara flavilinea shensiensis , genitalia. 92, <J seventh sternite ; 93,
aedeagus ; 94, $ eighth tergite and sternite ; 95, $ ; 96, $.

6o

A. WATSON

Figs. 97-100. Betalbara cupreogrisea, genitalia. 97, $ ; 98, o* ; 99, aedeagus ; 100, o*
seventh and eighth sternites and eighth tergite.

CHINESE DREPANINAE 61

Betalbara rectilinea sp. n.

(Text-figs. 101-104)

<J. Palp and front of head dark brown ; vertex grey-brown. Upper surface of antennae
grey-brown, bipectinate from base to about three-quarters of its length.

Thorax and abdomen as for colour of adjacent surface of wing. Colour-pattern of both
wings as in rugosa (PI. 3, fig. 317), ground-colour grey-brown with very pale brown markings.

Figs. 101-104. Betalbara rectilinea, genitalia. 101, $ ; 102, aedeagus ; 103, <$ ; 104,

g eighth tergite and sternite.

62 A. WATSON

Under surface of both wings pale grey-brown with yellowish costal area apicad on fore wing
and with trace of postmedial fascia on both wings. Legs very pale brown with outer surface
of pro- and meso thoracic legs grey-brown.

o* genitalia as in Text-figs. 102-104.

$. Similar to male, but antenna ciliate and very weakly biserrate.

$ genitalia as in Text-fig. 101.

This species apparently forms a superspecies with rugosa from which it differs in
the male genitalia (see rugosa). It is also closely related to prunicolor and flavilinea
but can be distinguished by the male and female genitalia, and by the shape on the
fore wing of the posterior half of the antemedial fascia which is straight and at right
angles to the anal margin of the wing. It is also separable from prunicolor by the
venation of the fore wing.

Distribution. China (Szechwan).

Wing. <$ 12-0-13-5 mm. (5) ; $ 12-0-14-0 mm. (2).

Holotype <$. China : Kwanhsien, vii.1930 (Franck) ; Drepanidae genitalia
slide No. 792. In the BM(NH).

Paratypes. BM(NH). China : 4 <$, 1 $, Szechwan, Kwanhsien, 15.viii.1925,
10.viii.1926, vii.1930 (Franck) ; 1 $, Szechwan, Mt. Omei, 4000-5000 ft., i.viii.1929
(Franck). U.S. National Museum. China : 1 $, Szechwan, Mt. Omei, Shinkaisi
(Graham).

Betalbara rugosa sp. n.

(PI. 3, fig. 317 ; Text-figs. 109-111)

Separable from rectilinea with which it forms a superspecies, by the male genitalia (particularly
by the shape of the eighth abdominal tergite and sternite, the aedeagus, gnathus, uncus, and
the weakly bifurcate socii).

Wing, o* 120-18-0 mm. (2).

Holotype <$. N.E. India : Naga Hills, 2000 ft., vii,viii.i889 (Doherty) ;
Drepanidae genitalia slide No. 798. In the BM(NH).

Paratypes. BM(NH). N.E.India: 1^, Khasis, x. 1895 (Nat. Coll.). Malaya:
1 <J, Pahang, Cameron Highlands, Ginting Kial, 5000 ft., 23. v. 1939.

Betalbara violacea (Butler) comb. n.

(PI. 3, fig. 318 ; Text-figs. 105-108, 112)

Agnidra violacea Butler, 1889 : 42. [Good fig.]
Drepana violacea (Butler) Strand, 191 1 : 203.
Albara violacea (Butler) Warren, 1922 : 469.
Albara violacea (Butler) ; Gaede, 1931 : 33.

Albara takasago Okano, 1959 : 38. Holotype <£, Central Formosa, Puli-Washe, v. 1958 [not
seen], syn. n.

Distinguished in both sexes from the rest of the genus by the obsolescent sub-
terminal fascia on either fore or hind wing, and in the male by the presence of only
one pair of spurs in the hind tibia. The male genitalia are also diagnostic, especially
by virtue of the presence of eversible sacs and the absence of an uncus.

CHINESE DREPANINAE

Figs. 105-111. Betalbara, <$ genitalia. 105-108, violacea. 105, aedeagus ; 106, seventh
sternite ; 107, eighth tergite, and eighth sternite showing left lateral sac ; 108, <$.
ioq-iii, rugosa. 109, eighth tergite and sternite ; no, aedeagus ; in, o*-

64

A. WATSON

Wings. <J 18 -o-20-o mm. (20) ; $ 18-5-22-0 mm. (9).

I have not seen the type of takasago but I have been able to study a male from
Central Formosa, identified as conspecific with the type by Dr. H. Inoue, who is in
agreement with me concerning the above synonymy.

Distribution. N.W. and N.E. India, Formosa, China (Szechwan, Yunnan,
Kwangtung, Chekiang, Fukien).

Material examined. Type. I select as LECTOTYPE a <$ syntype in the BM(NH)
labelled : Dharmsala 87.59 ; Agnidra violacea Butler type ; Drepanidae genitalia
slide No. 787.

Other material. BM(NH). India : 2 <$, Dharmsala, 1 <$, 1 $, Musuri, ix.1917,
vii-x.1922 (Mackenzie) ; 1 $, Dalhousie (Harford) ; 2 $, Darjeeling, Gopaldhara,
3440-5800 ft. (Stevens). China : 1 <$, Yunnan, 1918 (Forrest) ; 1 <$, [Szechwan],
Tu-pa-kep, 7400 ft., 4JX.1929 ; Formosa: i £, Central Formosa, 1959; 2$,
Fukien, Kuantun, i6.vii ; 9.viii.i938 (Hone) ; 2 <$, 1 $, Chekiang, West Tien-mu-

Fig. 112. Betalbara violacea, 9 genitalia.

CHINESE DREPANINAE 65

shan, 29.ix-26.x.ig32 (Hone). Museum Koenig, Bonn. China : 24 ex., Chekiang,
East and West Tien-mu-shan ; 11 ex., Fukien, Kuatun ; 1 ex., Kwangtung.
Daniel Collection, Munich. Formosa : 2 (J, 1 ?, Washai, vii.1958. China : 2 <$,
Chekiang, West-Tien-mu-shan, 1600 m., 28. iv., 8.vi.i932 (Hone).

Betalbara robusta (Oberthiir) comb. n.
(PI. 3, fig. 319 ; Text-figs. 113-117)

Drepana robusta Oberthiir, 1916 : 372.

Drepana robusta Oberthiir, 191 7 : fig. 3642. [Good fig.]

Drepana robusta Oberthiir, Gaede, 1931 : 27.

Albara robusta (Oberthiir) Gaede, 1933 : 168. [Fig.]

The large size, the colour-pattern, the shape of the fore wing, and the elongate
signum in the female genitalia separate robusta from the rest of the genus. The
affinities of this species are uncertain but it shares sufficient characters with the
remaining species to merit provisional inclusion in Betalbara. I prefer to place it
here rather than in Pseudalbara gen. n. because of the structure of the male eighth
abdominal sternite which, unlike that of Pseudalbara, is not greatly modified.

The specimens of this species from Shensi in the Museum Koenig, Bonn, are the
first known male examples of this species.

Wing, o* 225 mm. (i) ; 9 22-0-260 mm. (11).

Distribution. China (Szechwan, Shensi).

Material examined. Type. I select as LECTOTYPE a female syntype in the
BM(NH), figured by Oberthiir (1917), labelled : Chasseurs Indigenes des Mission-
aires de Ta-tsien-Lou [China, Szechwan], 1906, Drepana robusta Obthr. type ;
3642 ; ex Oberthiir Coll. Brit. Mus. 1927-3 ; Drepanidae genitalia slide No. 81.

Other material. BM(NH), < m\ \ : 5 $, [Szechwan], Ta-tsien-lou, 1910 ; 3 $,
[Szechwan], Tibet, frontiere orientale, 1905 ; 1 $, S. Shensi, Tapai-shan im Tsinling,
20. vi. 1935 (Hone). Museum Koenig, Bonn. China : 11 £, S. Shensi, Tapai-shan
im Tsinling (Hone).

PSEUDALBARA Inoue

(PI. 3, figs. 320, 321 ; Text-figs. 1 18-125)

Pseudalbara Inoue, 1962 : 7. Type-species Drepana parvula Leech, 1890 : 112, by monotypy.

As suggested by Inoue (1962), this genus is probably closest to Betalbara, from
which it can be separated by the presence of a vestigial frenulum in the male, the
colour-pattern of the fore wing, the absence of fasciae on the hind wing, the
anastomosis of 5c + R\ with Rs for a short distance distal to cell in hind wing, the
strongly modified seventh and eighth abdominal tergites and sternites in the male,
and the large corpus bursae in the female genitalia. It can be separated in the male
from all species of Betalbara, except manleyi, by the presence of lamellate antennae.

Two species are known : the type-species parvula (China, S.E. Russia and Japan)
and a new species fuscifascia (China).

66

A. WATSON

Figs. 113-117. Betalbara robusta, genitalia. 113, $ ; 114, $ ; 115, aedeagus ; 116, <J
eighth tergite and sternites ; 117, cj seventh sternite.

CHINESE DREPANINAE 67

Pseudalbara parvula (Leech)
(PI. 3, fig. 320 ; Text-figs. 118-121)

Drepana parvula Leech, 1890 : 112.

Drepana parvula Leech
Drepana parvula Leech
Drepana parvula Leech
Drepana parvula Leech

Leech, 1898 : 368.

Strand, 191 1 : 202.

Gaede, 1931 : 27.

Matsumura, 1931 : 742.
Albara parvula (Leech) Nagano, 1917 : 38.
Betalbara parvula (Leech) Matsumura, 1927 : 47.
Betalbara parvula (Leech) ; Inoue, 1956 : 369.
Betalbara parvula (Leech) ; Inoue, 1959 : 175. [Good fig.]
Pseudalbara parvula (Leech) Inoue, 1962 : 27. [Good figs.]
Drepana musculo Staudinger, 1892 : 335. [Synonymized by Leech (1898).]
Drepana griseola Matsumura, 1908 : 135. [Synonymized by Nagano (1917) ; confirmed by
Dr. H. Inoue in litt]

Distinguished from fascifascia by the short, pale, apical crescent and the absence
of a medial shade on the fore wing, by the venation of the fore wing in which vein Ri
arises from the proximal end of the areole, the bifurcate uncus and symmetric
seventh sternite in the male and by the ovate concave signum in the female genitalia.

Wing. cS n-5-14'0 mm. (2) ; $ 12-0-150 mm. (20).

Distribution. China (Chekiang, Fukien, Hupeh, Hunan, Szechwan, Kwangsi,
Manchuria), south-east U.S.S.R., Japan.

Material examined. Types, parvula. I have selected as LECTOTYPE a <$,
from the five syntypes in the BM(NH), labelled : Ningpoo, April, 1886, Leech ;
Leech Coll. 1900-64 ; Drepana parvula type <$ ; Drepanidae genitalia slide No. 626 ;
B. M. negative No. 29116.

muscula. Holotype <$, China, Szechwan ; Drepanidae genitalia slide No. 956.
In the Zoologisches Museum, Berlin.

griseola. Type material Japan, Kumamato ; presumably in Hokkaido Uni-
versity. [Not seen.]

Other material. BM(NH). China : 3 $, [Szechwan], Kwanhsien, vii.1930
(Franck) ; 1 $, Hupeh, Lui Shin Tze, viii.1912 (Betton) ; 1 $, [Hupeh], Ichang
(Bowring) ; 2 $, Hunan, Hoengshan, 7.vi., 10.viii.1933 ; 2 $, Fukien, Kuatun,
2300 m., 27 40' N, 117 40' E, 20.iv, 17.V.1938 (Klapperich) ; 4 $, Chekiang, West
Tien-mu-shan, 24.lv, 17. v., 30.viii.1932 (Hone) ; 3 $, Yachialing, vii.1922 (Bowring);
1 $, Ruling [? Szechwan, Kulin], vii.1921. Japan : 1 cJ. 5 $. Takao-San, W. of
Tokyo, 7.vii., 15. ix. 1926 (Aigner) ; 1 $, Japanese Alps, vii.1926 (Aigner).

Pseudalbara fuscifascia sp. 11.

(PI. 3, fig. 321 ; Text-figs. 122-125)

o*. Head and outer surface of palp greyish brown. Antenna yellowish brown ; serrate
and ciliate.

Colour of thorax and abdomen as for adjacent surface of wing. In the fore wing i?i arises
from middle of areole and i?2 from near distal end of areole.

68

A. WATSON

Figs. 118-121. Pseudalbara parvula, genitalia. 118, $ ; 119, $ ; 120, aedeagus ; 121,
<J seventh sternite, eighth tergite and sternites.

CHINESE DREPANINAE 69

Colour-pattern of upper surface as in PI. 3, fig. 321 ; pale areas yellowish brown, darker
areas greyish brown Whitish crescentic area at apex of fore wing bordered anteriorly
with dark brown. Upper surface of wings very pale yellowish brown, darker at costa and base
of fore wing. Under surface of fore wing with indistinct postmedial fascia and weakly marked
discocellular spot ; hind wing with two well-marked discocellular spots. Legs similar in
colour to under surface of wings but with outer surface of prothoracic legs greyish brown.

cJ genitalia as in Text-figs. 123-123

9. Similar to male, but antennae with shorter cilia.

$ genitalia as in Text-fig. 122.

Wing. <J 13-0-14-0 mm. (5) ; |. 15-0-16-5 mm. (5).

Separable from parvula by the more elongate, pale, apical crescent and the dark
medial shade on the fore wing, and by the fact that both R^ and R% arise from the
areole in the fore wing. The asymmetric seventh abdominal sternite and the short,
truncate uncus serve to distinguish the male genitalia, and the ribbon-like signum
the female genitalia.

Holotype <$. Chekiang, West Tien-mu-shan, 25.vii.1932 {Hone) ; Drepanidae
genitalia slide No. 929. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 2 $, 2 £, type-locality, 25.vii.-~
10.ix.1932 (Hone). BM(NH). China: 3^, 2 9, Szechwan, Kwansien, 10.viii.1926,
27.vii.1928, vii.1930 (Franck) ; 1 $, Szechwan, Tu-pa-keo, 7400 ft., 5.ix.

NORDSTROEMIA Bryk
(Pis. 3-10, figs. 322-329, 333-35° : Text-figs. 126-164)

Nordstrdmia Bryk, 1043 : 12. Type-species Nordstrdmia amabilis Bryk 1943 : 13, by original

designation.
Nordstrdmia Bryk ; Inoue, 1962 : 26.
Allodrepana Roepke, 1948 : 214. Type-species Allodrepana siccifolia Roepke, 1948 : 214,

by original designation. [Synonymized by Inoue, 1962 : 26.]
Albara Walker ; sensu Gaede, 1931 : 31. [Parti m

cj. Palp upturned to just above labrum. Antenna bipectinate from base to between about
three-fifths and three-quarters of its length ; shaft coated proximally with brilliantly lustrous
scales except in utidata.

Mesothoracic tibia with one pair of spurs, metathoracic tibia with two pairs of spurs. Fore
wing falcate except in some specimens of humerata ; vein /?i arises from proximal half of
areole in vira, but from near distal end of cell in remaining species ; Rz arises from near, or at,
distal end of areole, Sc + R\ in hind wing approximates to Rs for short distance distal to cell.
Ground-colour of upper surface of fore wing one of various shades of buff or grey ; costal area
with two or more dark patches ; antemedial fascia and postmedial fascia well-marked, lunulate
in undata, angled near costa in humerata, straight or nearly so in rest of genus ; two or more
discocellular spots usually present (pale in vira and bicostata, dark in remaining species) or with
line of dark scales along discocellular vein ; subterminal fascia, when present, a row of neural
spots, or in vira a continuous pale line ; fringe of outer margin as for ground-colour of wing,
but dark brown or grey apically. Ground-colour of upper surface of hind wing similar to fore
wing in most species but paler in undata ; in bicostata (q.v.) anal margin of hind wing is similar

7 o

A. WATSON

Figs. 122-125. Pseudalbara fuscifascia, genitalia. 122, $ ; 123, $ seventh sternite,
eighth tergite and sternites ; 124, <J ; 125, aedeagus.

CHINESE DREI'ANINAE 71

to fore wing in coloration but rest of wing is much paler and differently coloured ; antemedial
fascia well-marked and nearly straight in most species, but only present at anal margin in
bicostata, very poorly defined in humerata and either absent or weakly marked and lunulate in
undata ; postmedial fascia arcuate in humerata, present only at anal margin in bicostata, either
absent or weakly marked and lunulate in undata, strongly marked and either straight or slightly
arcuate in remaining species ; subterminal fascia represented by poorly marked neural spots
in at least some specimens of each species, except in bicostata and humerata which have no
subterminal fascia and in vira which has a pale continuous subterminal fascia. Under surface
of wings grey, buff or yellow, variously marked, usually with poorly defined postmedial fasciae.

o* genitalia : valve broad with well-developed sacculus and L- or T-shaped process bearing
long setae at base of costa ; medial sclerotization of diaphragma weakly developed except in
duplicata which bears three short spines ; short spine or lobe at each side arising from lateral
sclerotization of diaphragma, partly concealed by valve, well-developed except in undata
or humerata, spines overlapping in lilacina ; vinculum produced inwards posteriorly in bicostata
to form gnathus ; socius strongly developed, with arcuate apical spine except in undata,
humerata and duplicata ; uncus strongly developed, bifurcate apically except in humerata ;
aedeagus variously ornamented ; seventh abdominal sternite forming part of genitalia, except
in undata and humerata, weakly sclerotized, without apodemes in vira, bicostata and duplicata,
with single medial apodeme in remaining species ; eighth abdominal tergite well-developed,
broad, with lateral apodemes ; eighth abdominal sternite convex posteriorly in duplicata,
emarginate in rest of genus ; long eversible sac on each side of eighth sternite in vira.

9. As for male, but antennae filiform, weakly ciliate.

$ genitalia : ostium with lateral and ventral lips ; eighth segment well-developed, bilobed
and invaginate dorsally except in vira, bicostata and undata ; corpus bursae without signum,
or with single, small, oval, invaginate signum.

Nordstroemia is probably most closely allied to Betalbara Matsumura from which
it can be separated by the colour-pattern of the wings and by the male genitalia.

Except for humerata (and undata which is placed tentatively in Nordstroemia) the
species of this genus are remarkably uniform in colour-pattern and genital structure,
and the interspecific affinities are consequently difficult to assess. For example,
duplicata is externally almost identical to problematica, but similarities in the male
genitalia suggest a much closer relationship between japonica and problematica in
spite of the external differences between the latter two species. There is, however,
sufficient morphological evidence to suggest that particularly close affinities exist
between japonica, grisearia, agna, problematica, simillima, siccifolia, sumatrana,
argenticips, recava, lilacina and ochrozona.

In the following brief revision eight species have been transferred from Albara to
Nordstroemia and one from Drepana, two new species are described and several
names are relegated to synonymy. Sixteen species are now included in Nord-
stroemia. Except for sachalinensis Matsumura, the type of every nominal species
and subspecies has been examined.

Distribution. N. India, Sikkim, N.E. Burma, China, Formosa (undescribed
species), Japan, Malaya, Sumatra, and Java (undescribed species). The following
species are known to occur in China : vira, bicostata, japonica, agna, recava, duplicata
and undata. Twelve of the 16 species of Nordstroemia are endemic to the Indo-
Chinese Subregion, sumatrana and lilacina are endemic to the Malayan Subregion,
grisearia is restricted to the Manchurian Subregion, and japonica is shared by the
Indo-Chinese and Manchurian Subregions. (See Table 1.)

72 A. WATSON

Key to the Species
Males

i Antemedial fascia and postmedial fascia of upper surface of fore wing lunulate, or if

non-lunulate then sharply angled near costa ....... 4

- Antemedial fascia and postmedial fascia of upper surface of fore wing non-lunulate,

not angled near costa ........... 2

2 Subterminal fascia of upper surface of hind wing either absent or represented by row

of dark neural spots ; wing not pale yellow ....... 5

- Subterminal fascia either absent on upper surface of hind wing, in which case wing

is pale yellow except at anal margin, or represented by continuous pale line . 3

3 Subterminal fascia absent on upper surface of hind wing ; fore wing strongly

falcate (PI. 3, fig. 323) ; eighth sternite in male genitalia without eversible sacs
laterally (Text-fig. 129) ....... bicostata (p. 74)

- Subterminal fascia on upper surface of hind wing represented by a continuous pale

line ; fore wing weakly falcate (PI. 3, fig. 322) ; eighth sternite in male genitalia

with long eversible sac on either side ...... vira (p. 73)

4 Antemedial and postmedial fascia of upper surface of fore wing lunulate (PI. 4,

fig. 329) .......... undata (p. 90)

- Antemedial and postmedial fascia of upper surface not lunulate (PI. 4, fig. 328)

humerata (p. 90)

5 Ground-colour of upper surface of fore wing orange-bluff ; anterior half of hind wing

without markings (PI. 4, fig. 326) ...... siccifolia (p. 87)

- Ground-colour of upper surface of fore wing not orange-buff ; anterior third of hind

wing without markings (e.g. PI. 4, fig. 327) ....... 6

6 Outer margin of fore wing angulate at Cwia (PI. 4, fig. 327) . . recava (p. 84)

- Outer margin of fore and hind wing not angulate at Ciii & ..... 7

[The remainder of the Key is based entirely on characters in the male genitalia.']

7 Diaphragma with three short medial spines ; socius with short, non-arcuate, apical

spine ; posterior margin of eighth sternite convex (Text-figs. 158-160) duplicata (p. 88)

- Diaphragma without medial spines ; socius with arcuate apical spine ; posterior

margin of eighth sternite emarginate medially ...... 8

8 Lateral spines of diaphragma extending inwards across medial line (PI. 7, fig. 342)

lilacina (p. 87)

- Lateral spines of diaphragma not extending across medial line .... 9

9 Socius with two arcuate apical spines (PI. 10, fig. 353) . . . ochrozona (p. 88)

- Socius with single apical spine .......... 10

10 Apical spine of socius longer than basal part of socius (Text-fig. 138) problematica (p. 77)

- Apical spine of socius shorter than basal part of socius . . . . . 11

1 1 Anterolateral processes of diaphragma triangular, sharply pointed apically, directed

inwards towards medial line (PI. 6, fig. 339) .... sumatrana (p. 80)

- Anterolateral processes of diaphragma evenly rounded apically and inwardly directed

(e.g. PI. 8, fig. 347), or if not rounded then ventrally directed (PL 5, fig. 336) . 12

12 Anterolateral process of diaphragma evenly rounded apically . . . . 13

- Anterolateral process of diaphragma not rounded (PI. 5, fig. 336) argenticeps (p. 77)

13 Genitalia as in Text-figs. 149-152 ...... grisearia (p. 84)

- Genitalia not as in Text-figs. 149-152 . . . . . . . . 14

14 Genitalia as in Text-figs. 143-146 ...... japonica (p. 80)

- Genitalia not as in Text-figs. 143-146 ........ 15

15 Socius (omitting apical spine) over twice as long as broad; apical processes of uncus

not widely divergent (Text-fig. 133) ...... agna (p. 75)

- Socius (omitting apical spine) less than twice as long as broad ; apical processes of

uncus widely divergent (PI. 8, fig. 347) ..... simillima (p. 87)

CHINESE DREPANINAE 73

Nordstroemia vira (Moore) comb. n.
(PI. 3, fig. 322 ; PI. 5, figs. 333-335 ; Text-fig. 126)

Drepana vira Moore, [1866] : 817.

Drepana vira Moore ; Strand, 191 1 : 201.

Drepana vira Moore ; Gaede, 1931 : 28.

Albara vira (Moore) Warren, 1922 : 470. [Good fig.]

Albara vira (Moore) ; Gaede, 1931 : 28.

Albara erpina Swinhoe, 1894 : 433. [Synonymized by Hampson, 1896 : 476.]

Albara gracilli ma Warren, 1897 : 12. syn. n.

Albara gracillima Warren ; Gaede, 1931 : 31.

Drepana ocellata Oberthur, 1916 : 375. syn. n.

Drepana ocellata Oberthur, 191 7 : pi. 428. [Good fig.]

Albara ocellata (Oberthiir) Gaede, 1933 : 169.

Albara mimetica Warren, 1922 : 470. [Good fig.], syn. n.

Nordstroemia arnabilis Bryk, 1943 : 13. [Good fig.], syn. n.

Nordstroemia mimetica pallidina Bryk, 1943 : 14. [Good fig.], syn. n.

The continuous pale subterminal fascia on both wings and the fact that veins R\
and R2 both arise from the areole in the fore wing distinguish vira from every other
species of Nordstroemia. Similarly diagnostic are the long eversible sacs, placed
one on either side of the eighth abdominal sternite in the male genitalia. Close
affinities between vira and bicostata are suggested by the presence of whitish cell-
spots on the surface of the fore wing, and by the similarities in the shape of the
uncus and seventh abdominal sternite in the male genitalia.

Distribution. N.E. Burma, N. India, Sikkim and China (Szechwan, Fukien).

Material examined. Types, vira. The female type material of this species
(' Darjeeling Coll. A. E. Russell ') has not been traced ; Horn and Kahle (1937 : 380)
state that the Russell collection is lost. I therefore select as NEOTYPE a ^
specimen in the BM(NH) labelled : Darjeeling ; Moore Coll. 94-106.

erpina. LECTOTYPE <J, in the BM(NH), here selected, labelled : Kahsia Hs.
[Assam, Khasia Hills] 94-66 [Swinhoe collection] ; Drepana erpina Swinhoe $ type.

gracillima. Holotype c$, Khasis, Mar. 1895, Nat. Coll. ; Rothschild Bequest
B. M. 1939-1 ; Drepanidae genitalia slide No. 751. In the BM(NH).

ocellata. Holotype $, Siao-Lou [China, Szechwan], 1903, Chasseurs indigenes du
P. Dejean ; Drepanidae genitalia slide No. 753. In the BM(NH).

mimetica. LECTOTYPE <$, in the BM(NH), here selected, labelled : Khasis,
Nat. Coll. ; Albara mimetica Type ^ Warr. [in Warren's handwriting] ; Rothschild
Bequest B. M. 1939-1 ; Drepanidae genitalia slide No. 715.

arnabilis. Holotype c?> N.E. Burma, Kambaiti, 7000 ft., 10-21. iv. ; Drepanidae
genitalia slide No. 714. In the Naturhistoriska Riksmuseet, Stockholm.

pallidina. Holotype <$, N.E. Burma, Kambaiti, 2000 m., 9-17. vi. ; Drepanidae
genitalia slide No. 713. In the Naturhistoriska Riksmuseet, Stockholm.

Other material. BM(NH). India : 3 <£, 1 $, Darjeeling, 7000 ft., 25-31. iii.
1924 ; 1 $, Naini-Tal, 6600 ft., 20.viii.1934 ; 6 <$, 1 $. Khasis ; 1 <£, Assam, Jainta
Hills. Sikkim : 1^,1$, 1887 {M oiler). China : 3 <J, 1 $, Fukien, Kuatun,
2300 m., 27 40' N, 117 40' E, 3.iv.-28.v.i938 (Klapperich) ; 1 <J, [Szechwan]

74

A. WATSON

Kwanhsien, 18.vii.1925 ; 1 $, [Szechwan], Moupin, vi.1890. Museum Koenig,
Bonn. China : 17 ex., Fukien, Kuatun, 2300 m., 27 40' N, 117 40' E (Klapperich).

Nordstroemia bicostata (Hampson) comb. n.
(PI. 3, fig. 323 ; Text-figs. 127-132)
Drepana bicostata Hampson, 1912 : 1272.

This species is probably most closely related to vira but is readily separated from
it by the shape and colour-pattern of the fore wing, the reduced fasciae in the hind

Figs. 126, 127. Nordstroemia, $ genitalia. 126, vira ; 127, bicostata bicostata, $.

CHINESE DREPANINAE 75

wing, and by the genitalia in which the shape of the aedeagus, diaphragmal processes
and valve processes are diagnostic.

Two subspecies are known : the nominate subspecies (India, Sikkim and Burma)
and opalescens (China).

Nordstroemia bicostata bicostata (Hampson)

(PI. 3, fig. 323 ; Text-figs. 127-130)

Albara bicostata (Hampson) ; Warren, 1922 : 470. [PI. 49g as ' bicolorata '.]
Albara bicostata (Hampson) ; Gaede, 1931 : 31.

Separable from opalescens by the shape of the medial part of the gnathus and the
anterior processes of the diaphragma in the male genitalia.
Distribution. N. India, Sikkim and N. Burma.

Type. LECTOTYPE <$, here selected, labelled : Sikkim, ix.1909, F. Moller ;
Drepanidae genitalia slide No. 756. In the BM(NH).

Nordstroemia bicostata opalescens (Oberthiir) comb, n., stat. nov.

(Text-figs. 131, 132)

Drepana opalescens Oberthiir, 1916 : 375.
Drepana opalescens Oberthiir, 1917 : pi. 428. [Good fig]
Drepana opalescens Oberthiir, Gaede, 1931 : 27.
Albara opalescens (Oberthiir) Gaede, 1933 : 169. [Fig.]

Distinguished from the nominate subspecies (q.v.) by the male genitalia (Text-figs.

131. 132).

Distribution. China (Szcchwan).

Material examined. Type. LECTOTYPE q*. here selected, labelled : Tien-
Tsuen, 1897, ex. R. P. Demean ; Drepana opalescens $ Obthr. ex Oberthiir Coll.
Brit. Mus. 1927-3 ; Drepanidae genitalia slide No. 757. In the BM(NH).

Paralectotype. BM(NH). China : <J, [Szechwan], Tien-Tsuen (Dejean).

Other material. BM(NH). China, Szechwan : 1 <$, Tien-tsuen, 1897 (Dejean) ;
1 <J, 1 $, Kwanhsien, vii. 1930 (Franck) ; 1 <J, Kwanhsien, Omei, 10.vii.1929. United
States National Museum. China : 1 C J, Szechwan, Mt. Omei, Shin Kai Si, 4400 ft. ;
1 c$, S. of Suifu.

Nordstroemia agna (Oberthiir) comb. n.

(Text-figs. 133-137)

Drepana agna Oberthiir, 1916 : 373.

Drepana agna Oberthiir ; Oberthiir, 191 7 : pi. 428 (antemedial fascia on fore wing should be

straight in this plate).
Drepana agna Oberthiir ; Gaede, 1931 : 25.
Albara agna (Oberthiir) Gaede, 1933 : 169.

7 6

A. WATSON

Figs. 128-132. Nordstroemia, <$ genitalia. 128-130, bicostata bicostata. 128, 6* ; 129,
eighth tergite and sternite ; 130, aedeagus. 131, 132, bicostata opalescens. 131, aedeagus ;
132, 6*-

CHINESE DREPANINAE 77

The female lectotype and a male specimen in the BM(NH) (possibly a paralecto-
type) differ from the externally similar duplicata, which also occurs in China, in that
the antemedial and postmedial fascia on the upper surface of the wings is nearly
uniformly brown, not edged strongly with buff. The male genitalia also separate
agna and duplicata. The closest ally of agna is possibly problematica (type-locality
N.E. Burma) which has a weakly arcuate antemedial fascia on the fore wing, more
strongly marked antemedial and postmedial fasciae (the former edged proximally
with buff, the latter edged distally with buff, then greyish brown) on both wings,
and well-defined subterminal markings on the hind wing. The male and female
genitalia of agna and problematica are diagnostic.

Distribution. China, Szechwan province.

Material examined. Type. I select as LECTOTYPE the $ syntype figured by
Oberthiir, labelled : Siao-Lou, 1900, Chasseurs indigenes ; Drepana agna $ Obthr.
type ; Ex Oberthiir Coll. Brit. Mus. 1927-3 ; Drepanidae genitalia slide No. 766.
In the BM(NH).

Paralectotype. BM(NH). China, Szechwan : 1 <$, frontiere orientale de
Thibet, 1906 (Dejean).

Nordstroemia problematica (Bryk) comb. n.
(Text-figs. 138-142)

Albara problematica Bryk, 1943 : iy. [Good fig.]

Separable externally from the closely allied agna by the characters already listed
under the latter species (q.v.). The chief diagnostic feature in the male genitalia is
the long arcuate socius spine which is greater in length than the main part of socius.
The colour-pattern of argenticeps (q.v.) differs little from that of problematica but
the male genitalia show that the two species are probably not closely allied.

The type of/, aestivalis Bryk (1943 : 20) has been examined and found to be con-
specific with the type of problematica.

Distribution. Northern Burma.

Material examined. Type. Holotyp e cT» N.E. Burma, Kambaiti, 7000 ft., 5.viii.
(Malaise) ; Drepanidae genitalia slide No. 739 ; in the Naturhistoriska Riksmuseet,
Stockholm.

Other material. BM(NH). Burma : 1 J, N.E. Burma, Kambaiti, 7000 ft.,
4.iv.i934 (Malaise) ; 1 <$, Upper Burma, Htawgaw, 6000 ft. (Swann).

Nordstroemia argenticeps (Warren) comb. n.

(Pis. 5, 6, figs. 336-338)

Albara argenticeps Warren, 1922 : 470. [Fig.]
Albara argenticeps Warren ; Gaede, 1931 : 31.

There is little external difference between this species and sumatrana, except in
the shape of the postmedial fascia which is weakly sigmoid in argenticeps but straight

78

A. WATSON

Figs. 133-137. Nordstroemia agna, genitalia. 133, $ ; 134, <J eighth tergite and sternite ;
J 35. 6* seventh sternite ; 136, aedeagus ; 137, <j> (dorsal view).

CHINESE DREPANINAE

79

Figs. 138-142. Nordstroemia problematica, genitalia. 138, o* ; 139, aedeagus ; 140,
o* eighth tergite and sternite ; 141, <$ seventh sternite ; 142, $ (dorsal view).

80 A. WATSON

or only weakly sigmoid in sumatrana. The male genitalia, which indicate close
affinities with problematica, differ from the latter in the basally constricted socius
and the ventrally directed processes of the lateral sclerites of the diaphragma. In
comparison with agna the male genitalia of argenticeps are characterized chiefly by
the more strongly developed spine at the base of the valve, the longer socius spine,
the more strongly bifurcate uncus, and by the shape of the lateral processes of the
vinculum which are directed ventrally and flattened laterally, not dorsoventrally,
and are minutely spinose anteriorly.
Distribution. N.E. India.

Type. LECTOTYPE $, here selected, labelled: Khasis, Aug. 1895, Nat. Coll. ;
Albara argenticeps Type <$ Warr. ; Rothschild Bequest B.M. 1939-1 ; Drepanidae
genitalia slide No. 777. In the BM(NH).

Nordstroemia sumatrana (Roepke) comb. n.
(PI. 4, fig. 325 ; Pis. 6, 7, figs. 339, 340, 344)
Allodrepana sumatrana Roepke, 1948 : 214. [Figs.]

Probably most closely allied to argenticeps which it resembles externally except
for the more conspicuous subterminal markings and the straight or only slightly
sigmoid antemedial fascia on the fore wing. In the male genitalia the socii are not
constricted proximally and the processes of the lateral sclerites of the diaphragma
are dorsoventrally flattened and inwardly directed.

Distribution. Sumatra, and probably Malaya (ex. in BM(NH)).

Type. Holotype $, S. Sumatra, Mt. Tanggamus, 2100 m., xii. 1934 (Lieftinck
and Toxopeus) ; Drepanidae genitalia slide No. 1882. In the Rijksmuseum van
Natuurlijke Historic Leiden.

Nordstroemia japonica (Moore)
(Text-figs. 143-147)

Drepana japonica Moore, 1877 194.

Drepana japonica Moore ; Strand, 191 1 : 201.

Drepana japonica Moore ; Gaede, 1931 : 26.

Albara japonica (Moore) Warren, 1922 : 469.

Albara japonica (Moore) ; Inoue, 1956a : 663.

Albara japonica (Moore) ; Inoue, 1956 : 368.

Albara japonica (Moore) ; Inoue, 1959 : 175. [Good fig.]

Nordstroemia japonica (Moore) Inoue, 1962 : 27. [Good figs., including genitalia.]

Albara sachalinensis Matsumura, 1921 : 943. [Synonymized by Inoue (1956a).] [A syntype
of each sex of ab. punctifera Strand (191 1 : 201) has been examined and found to be con-
specific with the type of japonica.]

This species is possibly not separable externally from grisearia, to which it is
closely allied, but can be distinguished by the shape of the lateral diaphragmal lobes,
the socii, uncus and aedeagus in the male genitalia, and by the shape of the dorsal

CHINESE DREPANINAE

81

invagination of the eighth tergum and the degree of sclerotization of the ductus
bursae in the female.

Distribution. Japan and China (Hunan, Szechwan).

Material examined. Types, japonica. I select as LECTOTYPE of japonica a
$ [ex Pryer Coll.] in the BM(NH) labelled : Japan, 80-125 ! Drepana japonica
Moore Type [all except ' Type ' probably in Moore's handwriting] ; Drepanidae
genitalia slide No. 762. This specimen agrees exactly in wing-span with the figure
given by Moore (1877). Although it bears no Pryer collection label, the figures
' 80-125 ' [B. M. registration No. 1880-125] provide evidence that the lectotype was
at one time housed in the Pryer collection. Reference to the entry 1880-125 shows
that the specimens registered here are not Lepidoptera and were not taken in Japan.
However, two entries above 1880-125 is the entry 1880-123 ; the specimens regis-
tered here are Lepidoptera from the Pryer collection taken at Shanghai, the source

Figs. 143-146. Nordstroemia japonica, <$ genitalia. 143, <$ ; 144, seventh sternite
145, aedeagus ; 146, eighth tergite and sternite.

82

A. WATSON

of the majority of the material described by Moore in the paper where japonica is
described. It is reasonable to infer from this that the figures 1880-125 which occur
on the first of the lectotype labels should read 1880-123 and that this specimen is
part of the original syntypic material from the Pryer collection.

sachalinensis. [Type material, Japan, Sakhalin ; presumably in Hokkaido
University ; not seen.]

148

Figs. 147-148. Nordstroemia, genitalia. 147, japonica, $ ; 148, grisearia, $.

CHINESE DREPANINAE

83

Other material. BM(NH). Japan : numerous examples. China : 1 cJ,
Hunan, Hoeng-Shan, 900 m. ( 19.V.1935 (Hone) ; 1 $, C. China, Nanjang ; 1 <£,
Szechwan, Ta-tsien-lou, 1897 (Dejean). Museum Koenig, Bonn. China : 2 ex.,
Hunan, Hoeng-Shan, 900 m. (Hone). Japan : 3 ex.

Figs. 149-152. Nordstroemia grisearia, <J genitalia. 149,^; 150, aedeagus ; 151, seventh
sternite ; 152, eighth tergite and sternite.

84 A. WATSON

Nordstroemia grisearia (Staudinger) comb. n.

(Text-figs. 148-152)

Drepana grisearia Staudinger, 1892 : 335. [Poor fig.]
Drepana grisearia Staudinger ; Strand, 191 1 : 202. [Poor fig.]
Drepana grisearia Staudinger ; Gaede, 1931 : 26.

Since its description in 1892, this species has been known only from the female
holotype. Recently examined material from S.E. Russia and from Japan has
been compared with the holotype, and the male identified.

In coloration and colour-pattern there is little difference between this species and
the common Japanese species japonica Moore. The only apparent external differ-
ence in colour-pattern is in the subterminal fascia on the fore wing, which is absent
in all except one of the specimens of grisearia examined but present as faint neural
dots in nearly all the available japonica material. However, several features in the
male and female genitalia serve to distinguish grisearia from japonica (see list of
these under japonica). The extent of these morphological differences are sufficient
to justify specific separation of grisearia and japonica.

Distribution. Russia (Amur region and Vladivostok) and Japan (two localities,
Usui Pass and Chuzenyi, in central Japan, both at an altitude of approximately
1000 metres). The range of grisearia was presumably once much more extensive in
Japan than at the present, permitting genetic interchange with the Russian popula-
tions. Extinction must then have occurred leaving isolated populations of grisearia
in Japan. This is the pattern of distribution for which Kurentzov (1961) postulates
a Pleistocene origin. Possible hybridization with the closely allied and probably
monophyletic japonica may be prevented by the different times of emergence of the
imago which in grisearia appears in June, at least in the Amur region (Staudinger,
1892), and in japonica appears in April-May and July-September (Inoue, 1962).

Material examined. Type. Holotype $, S.E. Russia, Amur R., Raddefka ;
Drepanidae genitalia slide No. 731. In the Zoologisches Museum, Berlin.

Other material. Museum Koenig, Bonn. Japan : 1 $, Japan, Chuzenyi,
13.ix.1928. BM(NH). Japan : 1 <$, Chuzenyi, 10. ix. ; 1 <$, Central Japan,
Kotzuke, Usui-toge, [Central Honshu, Gumma Pref., Usui Pass, 950 m.] mid Aug.
1923 (Sugitani). Zoological Institute, Leningrad. Russia : 2 <J, Vladivostok,
8.iii.i9i4, 13.vi.1916 (Kriger-V oinovsky) .

Nordstroemia recava sp. n.

(PI. 4, fig. 327 ; Text-figs. 153-157)

o\ $. Palp orange-buff ; vertex of head dark brown, front paler brown ; antennae bipec-
tinate to just over half its length in male, filiform in female, proximal half of antennal shaft
very dark brown and brilliantly lustrous dorsally ; collar orange-buff.

Thorax speckled with greyish white and pale grey-brown dorsally, pale buff ventrally.
Colour-pattern of both wings as in PI. 4, fig. 327 ; ground-colour of both wings pale lustrous
grey, densely irrorate with layer of longer scales the latter pale greyish brown at base and
greyish white distally ; costa orange-buff ; apical part of outer margin and costal patches

CHINESE DREPANINAi:

85

dark brown ; fascia yellowish brown with yellow distal border to postmedial fascia, and yellow
proximal border to antemedial fascia ; trace of subterminal spot present on posterior veins
in some specimens. Under surface of both wings moderately lustrous ; ground-colour pale
dull yellow, sometimes slightly greyish basally on hind wing ; both wings with trace of greyish
yellow postmedial fascia ; costa of fore wing orange-buff ; fringe of outer margin dark brown
apically.

Legs pale yellow, but with outer surface of prothoracic and mesothoracic legs greyish brown
(prothoracic legs the darkest).

Dorsal surface of abdomen as for ground-colour of wings anteriorly ; pale yellowish grey
posteriorly ; with yellowish brown band on posterior margin of third segment. Ventral
surface as for ground-colour of hind wing.

<$ genitalia as in Text-figs. 153, 154, 156, 157.

$ genitalia as in Text-fig. 155.

Wing. (J 15-5-19-5 mm - ( 8 ) : ? 18-0-18-5 mm. (3).

Easily distinguished from its close allies, problematica, agna and japonica, by the
distinctive pre-apical emargination of the outer margin of the fore wing. The male
and female genitalia are also diagnostic. In the male, the shape of the uncus and
socii are particularly characteristic.

Distribution. China (Chekiang, Kiangsu).

Figs. 153, 154. Nordstroemia recava, genitalia. 153, <$ ; 154, aedeagus.

86

A. WATSON

Holotype $. China : Chekiang, E. Tien-mu-Shan, 21.V.1931 {Hone) ; Drepan-
idae genitalia slide No. 772. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 2 <$, 1 $, Chekiang, E. Tien-mu-
Shan, 21, 22.V.1931 (Hone) ; 1$, Kiangsu Lungtan b. Nanking, 9,io.ix.i9i8 (Hone) ;
3 (J, 2 $ Ruling, vii.1921. Daniel collection, Munich. China : 1 J, Chekiang,
W. Tien-mu-shan, 1600 m., 21. v. 1932 (Hone).

Figs. 155-157. Nordstroemia recava, genitalia. 155, $ (dorsal view) ; 156, $ eighth
tergite and sternite ; 157, 6* seventh sternite.

CHINESE DREPANINAE 87

Nordstroemia lilacina (Moore) comb. n.

(Pis. 7, 9, figs. 341, 342, 351)

Drepana lilacina Moore, 1888 : 401.
Albara lilacina (Moore) Warren, 1922 : 469.
Albara lilacina (Moore) ; Gaede, 1931 : 32.

The affinities of this species are uncertain. It is externally closest to simillima
(another large pale buff or pale greyish buff species) from which it can be distin-
guished by the dark yellowish brown antemedial and postmedial fasciae (sometimes
edged with buff and greyish brown) on the upper surface of the fore wing, and also
by the fact that the subterminal markings on the fore wing are best developed in the
middle of the wing, not at the apex. The two long, robust, arcuate, overlapping
spines of the lateral diaphragmal sclerites characterize the male genitalia.

An examination of a pair of syntypes of olivacea ab. pallidior Warren (1922 : 469,
good figs.) has shown them to be conspecific with the type of lilacina.

Distribution. N.E. and N.W. India.

Type. I select as LECTOTYPE a <J syntype in the BM(NH), labelled : Dharm-
sala 83.26 ; Drepana lilacina <$ type Moore ; Drepanidae genitalia slide No. 778.

Nordstroemia simillima (Moore) comb. n.
(Pis. 7, 8, figs. 343, 347, 348)

Drepana simillima Moore, 1888 : 402.

Albara simillima (Moore) Warren, 1922 : 469. [Fig.]

Albara simillima (Moore) ; Gaede, 1931 : 33.

This species is externally similar to lilacina with which it is sympatric. It is
separable from lilacina by the brownish white fasciae (edged faintly with pale brown)
on the upper surface of the fore wing, and by the markings of the subterminal fascia
on the fore wing which, when present, are most strongly marked in the anterior half
of the wing. The male genitalia, however, indicate closer relationships with agna,
problematica, japonica, recava but more particularly with siccifolia.

Distribution. N.W. India.

Type. I select as LECTOTYPE a J syntype in the BM(NH), labelled : Dharmsala
83.25 ; Drepana simillima $ type Moore ; Drepanidae genitalia slide No. 779.

Nordstroemia siccifolia (Roepke)

(PI. 4, fig. 326 ; Pis. 8, 9, figs. 345, 346, 352)

Allodrepana siccifolia Roepke, 1948 : 214. [Fig.]
Nordstroemia siccifolia (Roepke) Inoue, 1962 : 27.

The affinities of this species are uncertain, but it is possibly most closely related
to simillima from which it is distinguished by the orange-buff ground-colour of the
upper surface of the wings, by the dark brown proximal edge to the postmedial
fascia on the fore wing, and by the presence of three well-defined spots at the distal

88 A. WATSON

end of the cell. The male genitalia are also diagnostic (particularly the uncus and
the broad lateral lobes of the diaphragma).

Distribution. Sumatra.

Type. Holotype^, S. Sumatra, Mt. Tanggamus, 2100 m., 11.vii.1934 (Toxopeus).

Nordstroemia ochrozona (Bryk) comb, n., stat. n.

(Pis. 9, io, figs. 349, 353, 354)
Albara duplicata ochrozona Bryk, 1943 : 20. [Good fig.]

This species is probably indistinguishable externally from duplicata, but is not
closely allied to it. The affinities of ochrozona are uncertain : it is separable from
the rest of the genus by the hammer-shaped lateral process of the diaphragma and
by the presence of two arcuate spines, curved in places at right-angles to each other
at the apex of the socius.

Distribution. N.E. Burma.

Type. Holotype $, N.E. Burma, Kambaiti, 7000 ft., 10. iv. 1934 (Malaise) ;
Drepanidae genitalia slide No. 717. In Naturhistoriska Riksmuseet, Stockholm.

Nordstroemia duplicata (Warren) comb. n.
(PI. 4, fig. 324 ; Text-figs. 158-160)

Albara duplicata Warren, 1922 : 469. [Published simultaneously with olivacea Warren ; here

selected as the senior synonym.]
Albara duplicata Warren ; Gaede, 193 1 : 31.
Albara olivacea Warren, 1922 : 469. syn. n.
Albara olivacea Warren ; Gaede, 193 1 : 32.

Separable from agna, which also occurs in China and has a similar colour-pattern,
by the buff-edged antemedial and postmedial fascia on the upper surface of the
wings and by the distinctive male genitalia. Although duplicata is almost identical
externally to problematica and ochrozona (both described from N.E. Burmese
material) and not significantly unlike agna, japonica, argenticeps and other species
of this group (see under generic description), the male genitalia of duplicata are
remarkably different from these species (Text-figs. 158-160), especially in the shape
of socii and uncus and the spinose diaphragma.

Warren's description of olivacea was based on specimens differing from the type
material of duplicata in that the ground-colour of the upper surface of the wings was
' olive grey ' [now faded to pale buff], contrasting with the ' lilac grey ' [now pale
grey] of duplicata. The syntypes of olivacea ab. pallidior have been shown to be
conspecific with the type of lilacina (q.v.).

Distribution. N.E. India, China (Chekiang and possibly Yunnan).

Material examined. Types, duplicata. LECTOTYPE $, here selected, labelled :
Khasis, Nat. Coll. ; Coll. H. J. Elwes ; Albara duplicata Type^ Warr. ; Rothschild
Bequest B. M. 1939-1 ; Drepanidae genitalia slide No. 718 ; B. M. negative No.
29113. In the BM(NH).

CHINESE DREPANINAE

89

olivacea. LECTOTYPE $, here selected, labelled : Khasis, Mar. 1895, Nat.
Coll. ; Albara olivacea Type <$ Warr. ; Rothschild Bequest B. M. 1939-1 ;
Drepanidae genitalia slide No. 764. In the BM(NH).

Other material. Museum Koenig, Bonn. China : 1 <$, Chekiang ; 1 $, N.
Yunnan (doubtfully identified).

Figs. 158-160. N ordstroemia duplicata, q" genitalia. 158,0"; 159, aedeagus ; 160, eighth

tergite and sternite.

go A. WATSON

Nordstroemia humerata (Warren) comb. n.

(PI. 4, fig. 328; Pis. 9, 10, figs. 350, 355, 356)

Albara humerata Warren, 1896 : 335.

Albara humerata Warren ; Warren, 1922 : 470. [Fig.]

Albara humerata Warren ; Gaede, 193 1 : 31.

This species is readily separable from the other species of Nordstroemia by the
small size, the weakly falcate fore wings, the distinctive colour-pattern of the wings
(see plate in Warren, 1922) and by the simple unbranched socii and uncus in the
male genitalia. The affinities of humerata within Nordstroemia are doubtful, though
its present generic placement is probably correct.

Distribution. N.E. India.

Type. LECTOTYPE $, here selected, labelled : Khasis, Sept. 1895, Nat. Coll.,
Albara humerata Type g Warr. ; Rothschild Bequest B.M. 1939-1 ; Drepanidae
genitalia slide No. 1670. In the BM(NH).

Nordstroemia undata sp. n.

(PL 4, fig. 329 ; Text-figs. 161-164)

o*. Outer surface of palp, front of head, and pectinations and base of antenna reddish brown ;
rest of antenna and patch posterior to base of antenna buff ; patch between antennae, posterior
margin of vertex and collar very pale buff.

Thorax buff. Colour-pattern of upper surface as in PI. 4, fig. 329. Ground-colour of upper
surface of fore wing buff, slightly darker apically ; medial fasciae and cell-spots darker buff,
subterminal fascia dark brown ; outer margin edged with orange-buff ; fringe dark brown.
Hind wing brownish white on upper surface ; trace of postmedial and subterminal fascia at
anal margin in some specimens, discocellular spot of under surface usually showing through
wing. Under surface of fore wing buff, more brownish apically, at base and along costa ;
with dark brown discocellular spot, trace of spot at posterior angle of cell, and trace of anterior
part of postmedial fascia ; outer margin and fringe as for upper surface. Under surface of
hind wing pale buff irrorate with brownish buff distally and anteriorly ; usually with faintly
marked brownish buff postmedial fascia ; cell-spots as for fore wing. Legs pale buff with
outer surface of prothoracic leg darker buff.

Abdomen brownish white.

o* genitalia as in Text-figs. 162-164. Diaphragma with very weakly developed lateral
sclerites ; seventh abdominal sternum unmodified.

$. As for male but with filiform, sparsely ciliate antennae.

$ genitalia as in Text-fig. 161.

Wing, o* 17-0-19-5 mm. (8) ; $ 17-5-21-0 mm. (17).

Readily distinguished from the rest of the genus by the very distinctive colour-
pattern, undata has been placed in Nordstroemia because of general generic resem-
blances in the male and female genitalia. It is, however, clearly much less closely
allied to the type-species than the remaining species of Nordstroemia.

Holotype J. China : N. Yunnan, Likiang, 22. vi. 1935 (Hone) ; Drepanidae
genitalia slide No. 975. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 1 <J, 5 $ N. Yunnan, A-tun-tse,

CHINESE DREPANINAE

91

7.vii-u.vii.i936, 30.vii.1937 (Hone) ; 9 $, 14 9. N. Yunnan, Likiang, i.vii-30.ix.
1935 (Hone). Daniel Collection, Munich. China : 2 <$, 2$, N. Yunnan, 29. v-
3.vii.i935, 16.vii.1936 (Hone). BM(NH). China : 1 <$, N. Yunnan, A-tun-tse,
7.vii.i936 (Hone) ; 2 <$, 2 $, N. Yunnan, Likiang, 25.vi-26.vii.1935 (Hone).

Figs. 161-164. Nordstroemia undata, genitalia. 161, $ ; 162, aedeagus ; 163, <$ ; 164,

<£ eighth terigte and sternite.

92 A. WATSON

DIDYMANA Bryk

Didymana Bryk, 1943 : 10. Type-species Didymana renei Bryk, 1943 : io, by monotypy.

The affinities of this genus are doubtful, but it is possibly most closely related to
Palaeodrepana Inoue, especially to Palaeodrepana harpagula (Esper). Didymana
is monotypic.

Didymana bidens (Leech) comb. n.

(PI. n, fig. 357 ; Text-figs. 165-169)

Drepana bidens Leech, 1890 : 113.

Drepana bidens Leech ; Strand, 191 1 : 202. [Fig.]

Didymana renei Bryk, 1943 : 10. [Good fig.] syn. n.

Readily distinguished from Palaeodrepana harpagula (Esper) (p. 94), to which it is
possibly most closely related, by the colour-pattern (see Strand, 191 1 and Bryk,
1943) and the genitalia (Text-figs. 65-69). The shape of the seventh sternite and
the eighth tergite in the male are particularly diagnostic.

Distribution. China (Hupeh, Szechwan, Yunnan, Shensi, Fukien) and N.E.
Burma.

Material examined. Types, bidens. Holotype <$, [China, Hupeh], Chang Yang,
vii.1888 {Pratt) ; Drepanidae genitalia slide No. 82. In the BM(NH).

renei. Holotype <$, N.E. Burma, Kambaiti, 2000 m. ; Drepanidae genitalia
slide No. 734. In the Naturhistoriska Riksmuseet, Stockholm.

Other material. BM(NH). China : 6 ex., Szechwan. N.E. Burma : 14 ex.
Museum Koenig, Bonn. China : 29 examples from S. Shensi, Fukien and N.
Yunnan. Naturhistoriska Riksmuseet, Stockholm. N.E. Burma : 11 ^ paratypes
of renei, Kambaiti, 2000 m., 14.iv-14.vi.1934. U.S. National Museum. China :
1 (J, 2 $, Szechwan.

PALAEODREPANA Inoue
(PL 4, figs. 330-332 ; Text-figs. 170-182)

Palaeodrepana Inoue, 1962 : 21. Type-species Bombyx harpagula Esper, [1786] : 373, by
original designation.

Inoue established this genus for three species : harpagula, binaria Hufnagel
(1769 : 516) and cultraria Fabricius (1775 : 621). Of these only harpagula is known
to occur in China — it is also represented in Britain and much of the western part of
the Palaearctic Region, a range matched only by Drepana curvatula Borkhausen.

The closest ally of Palaeodrepana, more particularly of harpagula, is possibly
Didymana Bryk on the evidence of external and genitalic similarities. The species
binaria and cultraria may not be properly placed in Palaeodrepana and merit further
study in this respect.

CHINESE DREPANINAE

93

Figs. 165-169. Didymana bidens, genitalia. 165, <J ; 166, $ seventh sternite ; 167, <J
eighth tergite and sternite ; 168, aedeagus ; 169, $.

94 A. WATSON

Palaeodrepana harpagula (Esper)

(PI. 4, figs. 330-332 ; Text-figs. 170-182)

Bombyx harpagula Esper, [1786] : 373. [Good figs.]

Bombyx harpagula Esper ; Hiibner, [1803] : pi. 11, fig. 41 [labelled ' sicula ' in error] ; [1809] :

113 [including reference to pi. 11, fig. 41 as figuring harpagula].
Drepana harpagula (Esper) Gaede, 1931 : 23.
Palaeodrepana harpagula (Esper) Inoue, 1962 : 22.

Distinguished from the other two species of this genus by the shape and colour-
pattern of the wings and by the male genitalia (see Plate and Text-figs.).

Distribution. Occurs in most of the temperate parts of the Palaearctic Region
and in Oriental China. Four subspecies are known : the nominate subspecies
(Europe, S.E. Russia, China), olivacea (Japan) emarginata (China), and bitorosa
(China).

Palaeodrepana harpagula harpagula (Esper)
(PI. 4, fig. 331 ; Text-figs. 170-174)

Distinguished from the other three subspecies by the shape of the wings and the
genitalia of both sexes.

Distribution. Known from most European countries, and also from S.E. Russia
and from the Chinese province of Manchuria (material in the BM(NH)).

Type. Holotype $, Germany, Frankfurt. Probably lost : there are no specimens
of harpagula in the Esper collection in the Zoologische Sammlung, Munich.

Palaeodrepana harpagula olivacea (Inoue)

Drepana harpagula olivacea Inoue, 1958 : 12.

Palaeodrepana harpagula olivacea (Inoue) Inoue, 1962 : 23. [Good figs.]

A description and comparison of this subspecies is given by Inoue (1958) and
(1962).
Distribution. Known from most parts of Japan (see Inoue, 1962).

Type. Holotype <$, Japan, Takao-san, Toyko, 2.vii.i949 {Inoue). In Inoue
collection.

Palaeodrepana harpagula emarginata ssp. n.

(PI. 4, figs. 332 ; Text-figs. 179-182)

Differs from the nominate subspecies in the following respects : fore wing much more strongly
falcate and outer margin sharply angled outwards at Cwia ; hind wing angled at Cu\a. ', male
genitalia distinctive (see Text-figs. 180-182) ; ostium and eighth segment of female genitalia
differently shaped (Text-fig. 179).

Wing. <$ 16-5-20-5 mm. (14) ; $ 18-0-22-5 mm - (8)-

Distribution. China (Chekiang, Fukien, and possibly Shansi).

CHINESE DREPANINAE

95

HolotypecJ. China : Chekiang, West Tien-Mu-Shan, 1600 m. ( 26.V.1932 {Hone) ;
Drepanidae genitalia slide No. 699. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 4 <$ and 4 9. Chekiang, West
Tien-Mu-Shan, 1600 m., 26.V.-25JX.1932 {Hone) ; 8^ and 1 $, East Tien-Mu-Shan,
I7.v.-i4.vi.i93i {Hone) ; 3^ and 1 $, Fukien, Kuatun, 2300 m., 19.iv.-27-v.1938
{Hone).

Figs. 170-174. Palaeodrepana harpagula harpagula, genitalia. 170, ^ eighth tergite and
sternite ; 171, <J seventh sternite ; 172, aedeagus ; 173, <$ ; 174, $.

A. WATSON

Figs. 175-182. Palaeodrepana, genitalia. 175-178, harpagula bitorosa. 175, $ ostium ;
176, aedeagus ; 177, <$ ; 178, o" eighth sternite. 179-182, harpagula emarginata. 179,
$ ostium ; 180, <J eighth sternite ; 181, aedeagus ; 182, <$.

CHINESE DREPANINAK 97

A female, in the Museum Koenig, from Shansi province, N. China, may also prove
to belong to this subspecies.

Palaeodrepana harpagula bitorosa ssp. n.

(PI. 4, fig. 330 ; Text-figs. 175-178)

Similar externally to emarginata, but with fore wing not so strongly produced in most
specimens. The male genitalia (Text-figs. 176-178) and the shape of the anterior lip of the
ostium in the female genitalia (Text-fig. 175) are diagnostic.

Wing, o* 18-0-21-5 mm (4°) ; ? 18-0-21-5 mm - ( IO )-

Distribution. China (S. Shensi, and possibly Szechwan).

Holotype <J. China : S. Shensi, Tapaishan im Tsinling, c. 1700 m., 22. vi. 1935
{Hone) ; Drcpanidae genitalia slide No. 697. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China: 71^ and 16 $, S. Shensi, Tapaishan
im Tsinling, c. 1700 m., 21.vi.-3.vii.1936 {Hone).

A male from Mt. Omei (Szechwan) in the U.S. National Museum, together with a
female from Ta-tsien-lou (Szechwan) and another from Kwanhsirn (Szechwan), in
the BM(NH), belong either to this subspecies or to a new subspecies, but further
material is needed from Szechwan before either of these alternatives can be con-
firmed.

STREPSIGONIA Warren

Strepsigonia Warren, 1K97 : 17. Type->p«'» ies Strepsigonia nigriwaculata Warren, 1897 : 17,

by original designation.
Strepsigonia Warren ; Gaede, 1931 : 9.
Monurodes Warren, 1923 : 475. Type-species Monurottes trigonoptoa Warren, 1923 : 475,

by monotypy. syn. n.

Distribution. India, Sikkim, Burma, China, Malaysia, Indonesia and the
Philippines. Six species are at present recognized. Only one species, diluta, is
known to occur in China.

A study of the non-Chinese species of Strepsigonia may reveal more precisely
where its affinities lie, but its present placement near Palaeodrepana and Canucha
seems reasonable.

Strepsigonia diluta (Warren)
(PI. 11, fig. 359 ; Text-figs. 183-187)

Tridrepana diluta Warren, 1897 : 18.
Tridrepana diluta Warren ; Warren, 1922 : 467. [Fig.]
Tridrepana diluta Warren ; Gaede, 1931 : 28.
Strepsigonia diluta (Warren) Watson, 1957 : 4 11 -

Distribution. Known from N. India, Sikkim, and from a single Chinese specimen
(Kwangtung, Canton) in the Hone collection, Bonn, which probably represents a
new subspecies of diluta.

9 8

A. WATSON

Figs. 183-187. Strepsigonia diluta, genitalia. 183, $ ; 184, $ ; 185, $ eighth tergite
and sternite ; 186, <$ seventh sternites ; 187, aedeagus.

CHINESE DREPANINAE 99

CANUCHA Walker
(PI. 11, figs. 366-368 ; Text-figs. 188-199)

Canncha Walker, 1866 : 1574. Type-species Canucha curvaria Walker, 1866 : 1574, by mono-

typy.
Canucha Walker ; Gaede, 1931 : 40.
Campylopteryx Warren, 1902 : 340. Type-species Campylopteryx sublignata Warren, 1902 : 340,

by monotypy. syn. n.
Campylopteryx Warren ; Gaede, 1931 : 40.

This is a genus of six described species : curvaria Walker, 1866 : 1574, (Mysol Is.
New Guinea and the Solomons) ; sublignata Warren, 1902 : 340 (Buru, Amboina,
Watubela, New Guinea, Solomons) ; specularis Moore, 1879 : 407 (Ceylon, India,
China, Malaysia, Indonesia) ; miranda Warren, 1923 : 475 (N.E. India, Formosa);
bouvieri Oberthiir, 1916 : 272 (China) and duplexa Moore, 1865 : 816 (N.E. India,
Sikkim, Burma). The distribution is summarized in Table I.

The affinities of Canucha are uncertain, but it is probably not taxonomically
distant from Drepana Schrank.

A short account is given below of the species that occur in Formosa or China
{specularis, bouvieri and miranda) and of duplexa which is very closely allied to
bouvieri.

The name ' depressa Warren ' was first published in 19 16 accompanying a figure in
Die Gross-Schmetterlinge der Erde 10 : pi. 49, but was not then binominal. The
name was used again, this time in a binominal sense with Canucha, Seitz, 1934,
Die Gross-Schmetterlinge der Erde 10 : 859, Druckfchlrr und Berichtigung, but the
name was not accompanied by a diagnosis, thus contravening Article 13 of the
International Code of Zoological Nomenclature, and therefore remains unavailable.
A female ' syntype ' of ' depressa ' from Rendova (Solomons), in the BM(NH), is
probably conspecific with the male holotype of curvaria.

Canucha specularis (Moore)
(PI. 11, fig. 367 ; Text-figs. 196-199)

Drepana specularis Moore, 1879 : 407.

Drepana specularis Moore ; Moore, 1882 : 120. [Fig.]

Canucha specularis (Moore) Warren, 1923 : 475. [Fig.]

Canucha specularis (Moore) ; Gaede, 1931 : 40.

Platypteryx obtruncata Warren, 1900 : 117. [Synonymized by Gaede, 1931.]

The genitalia and the presence of well-developed subterminal spots on the fore
wing and of two closely apposed hyaline patches in the hind wing distinguish this
species from the closely related curvaria Walker.

Distribution. Ceylon, India, China, Java, Sumatra, Borneo and Celebes. A
single male from Lofanshan (S. China) in the Museum Koenig, Bonn, is the only
known Chinese specimen : it may prove to represent a new subspecies of specxdaris.

Types, specxdaris. The female type material (' Ceylon (Sir W. Gregory) ') may
be lost. No statement was given by Moore (1879) concerning the location of the

ioo A. WATSON

type material which may have been deposited in his own collection, now the property
of the BM(NH). The only specimen from the type locality in the BM(NH), a female
presented by W. Lindsay, is slightly too small to fit the measurements given in the
original description though it otherwise matches the description and the figure
subsequently published by Moore (1882).

obtruncata. Holotype $. This bears a printed label ' Bahia ', obviously due to
an error in labelling. Hampson has affixed a label ' Drepana specularis Moore, fr.
Ceylon G.F.H.' The holotype certainly must have been captured in the Oriental
Region.

Canucha duplexa (Moore)
(PI. 11, fig. 368 ; Text-figs. 188-192)

Drepana duplexa Moore, [1866] : 816. [Good fig.]
Canucha duplexa (Moore) Warren, 1923 : 475.

Type. The type material (' Darjeeling ') was deposited in the A. E. Russell
collection, but no trace can be found of this collection, which is lost according to
Horn and Kahle (1937 : 380). I select as NEOTYPE a^ specimen in the BM(NH)

Fig. 188. Canucha duplexa duplexa, $ genitalia.

CHINESE DREPANINAK

Figs. 189-195. Canucha, genitalia. 189, 190 duplexa duplexa. 189, $ ; 190, aedeagus.
191, 192, duplexa birmana. 191, <$ ; 192, aedeagus. 193, bouvieri, $. 194, 195,
miranda. 194, aedeagus ; 195, $.

102 A. WATSON

labelled : Darjeeling (Pilcher) ; Rothschild Bequest B.M. 1939-1 ; Drepanidae
genitalia slide No. 1029.

The colour-pattern and the male genitalia separate duplexa from miranda Warren.
C. bouvieri Oberthiir (q.v.) may later prove to be synonymous with duplexa birmana
when males of the former are available for study.

Two subspecies are at present recognized : the nominate subspecies (India) and
birmana (Burma).

Canucha duplexa duplexa (Moore)

(PI. 11, fig. 368 ; Text-figs. 188-190)

Canucha duplexa (Moore) ; Warren, 1923 : 475. [Good fig.]
Canucha duplexa (Moore) ; Gaede, 1931 : 40.

Distinguished from duplexa birmana by the non-angulate postmedial fascia on the
hind wing and by the male genitalia.
Known only from N.E. India.

Canucha duplexa birmana Bryk
(Text-figs. 191, 192)
Canucha miranda birmana Bryk, 1943 : 27. [Good fig.]

Separable from the nominate subspecies by the male genitalia (Text-figs. 191, 192)
and by the angulate postmedial fascia on the hind wing.

The name birmana may prove to be a junior synonym of bouvieri when male
material of the latter is made available for comparison.

196

Figs. 196-199. Canucha specularis, genitalia. 196, $ ; 197, aedeagus ; 198, <$ eighth
sternite ; 199, $ (unornamented corpus bursae not shown).

CHINESE DREPANINAE 103

Known only from N.E. Burma.

Type. Holotype^, N.E. Burma, Kambaiti, 700 ft., io.v.1934 (Malaise). In the
Naturhistoriska Riksmuseet, Stockholm.

Canucha bouvieri Oberthur
(PI. 11, fig. 366 ; Text-fig. 193)

Canucha bouvieri Oberthur, 1916 : 272.

Canucha bouvieri Oberthur ; Oberthur, 191 7 ; pi. 428. [Good fig.]

Canucha bouvieri Oberthiir ; Gaede, 1932 : 168.

Drepana bouvieri (Oberthur) Gaede, 1931 : 26.

This nominal species may prove to be a subspecies of duplexa (Moore) and its
name a senior synonym of duplexa birmana Bryk, but male specimens from
Szechwan are needed before a satisfactory comparison of material can be made.

Distribution. Known only from two female specimens from the type locality
(China, Szechwan) in the collection of the BM(NH).

Type. Holotype 9 (not <$ as stated by Oberthur), Ta-tsien-lou, 1910 ;
Urepanidae genitalia slide No. 1028. In the BM(NH).

Canucha miranda Warren

(Text-figs. 194, 195)

Canucha miranda Warren, 1923 : 475. [Good coloured fig.]

Canucha miranda Warren, Gaede, 1931 : 40.

Canucha miranda I. formosicola Matsumura, 1931 : 741.

Separable from duplexa by the male genitalia and the colour pattern.

Distribution. Assam, N.E. India and Formosa. Through the kindness of Dr.
H. Inoue I have seen a photograph of the single type specimen of formosicola. It
is doubtless conspecific with the lectotype of miranda although it may prove to
represent a distinct subspecies.

Type, miranda. LECTOTYPE <$, here selected, labelled : Khasis, Oct. 1896,
Nat. Coll. ; Canucha miranda Type <$ Warr. ; Drepanidae genitalia slide No. 1027 ;
Rothschild Bequest B.M. 1939-1. In the BM(NH).

DREPANA Schrank
(PI. 11, figs. 358, 360-365 ; Text-figs. 200-219)

Drepana Schrank, 1802 : 155. Type-species Phalaena falcataria L., 1758 : 519, by subsequent
designation by Westwood, 1840 : 104. [See 1961, Bull. zool. Nom. 18 : 267.]

Drepana Schrank ; Gaede, 1931 : 17.

Drepana Schrank ; Inoue, 1962 : 20.

Drepania Hiibner, [1819 : 140]. An unjustified emendation of Drepana Schrank. [See 1961,
Bull. zool. Nom. 18 : 267.]

Platypteryx Laspeyres, 1803 : 29. Type-species Phalaena falcataria L., 1758 : 519, by sub-
sequent designation by Latreille, 18 10 : 441.

io 4 A. WATSON

' Platypterix ' ; Ochsenheimer, 1816 : 97. An incorrect subsequent spelling of Platypteryx
Laspeyres.

Falcaria Haworth, 1809 : 152. Type-species Phalaena lacertinaria L., 1758 : 519, by sub-
sequent designation by Kirby, 1892 : 733.

Prionia Hubner, [1819] : 150. Type-species Phalaena lacertula Denis and Schiffermuller,
[1776] : 64, by monotypy. syn. n.

Inoue (1962 : 21, 22) recently transferred to a new genus, Palaeodrepana (p. 92),
three species previously classified under Drepana, and it is clear that further re-
classification of Drepana, sensu Gaede (1931), is necessary. I consider, however,
that the four Chinese species dealt with in the following account are congeneric
with the type-species of Drepana. Several species included in Drepana by Gaede
(1931) have been transferred in this paper to other genera (see index).

There are overall similarities between Drepana and Tridrepana Swinhoe (p. 11 1)
which suggest possible close phyletic affinities.

Drepana curvatula (Borkhausen)

(PI. 11, figs. 358, 363)

Phalaena curvatula Borkhausen, 1790 : 460.

Drepana curvatula (Borkhausen) Gaede, 1931 : 20.

Drepana curvatula (Borkhausen) ; Inoue, 1962 : 21. [Good figs, of moth and genitalia.]

This species is closely allied to Drepana falcataria L. (1758 : 519) and is known
to be capable of forming hybrids with the latter (see Strand, 1911 : 199). It can
be distinguished from falcataria by the colour-pattern and by minor differences in
the male genitalia.

Drepana curvatula curvatula (Borkhausen)
(PI. 11, fig. 358)

Drepana curvatula (Borkhausen) ; Gaede, 1931 : 20.
Drepana curvatula (Borkhausen) ; Strand, 191 1 : pi. 23b.

Bombyx sicula Esper ; sensu Hubner, [1803] : pi. 11, figs. 42, 43 ; [1809] : pi. 11, fig. 4 ; [1838] :
pi. 83, fig. 351 ; [1819] : 150 (' sicula Schiffermuller ').

The nominate subspecies has been recorded from most European countries. It
can be distinguished from acuta by the colour-pattern of the wings.

Type. Holotype $, Germany, Frankfurt. [Figured by Ernst and Engramelle
(1786 : pi. 208, figs. 27b, f, g).] In Wiesbaden.

Drepana curvatula acuta Butler

(PI. 11, fig. 363)

Drepana acuta Butler, 1881 : 59.

Drepana curvatula acuta Butler ; Inoue, 1962 : 21. [Good figs, of moth and genitalia.]

Falcaria curvatula acuta (Butler) Inoue, 1959 : 175. [Good fig.]

Drepana curvatula japonibia Strand, 191 1 : 200. [Synonymized with acuta by Inoue, 1962 :

21.]

CHINESE DREPANINAE 105

Drepana curvatula urupitla Bryk, 1942 : 27. [Synonymized with acuta by Inoue, 1962 : 21.]
Drepana curvatula koreula Bryk, 1949 : 27. [Synonymized with acuta by Inoue 1962 : 21.]

Distinguishable from the nominate subspecies only by the colour-pattern (see
references to figs, above).

Distribution. Japan (see Inoue 1959, 1962), S.E. Russia and Kuril Islands,
Korea, China (Manchuria). There is a single male in the Hone collection, Bonn,
from Shansi which also probably represents this subspecies.

Types examined, acuta. I select as LECTOTYPE a male from the original
series in the BM(NH) labelled : Tokei [Tokyo], 89.97 ! Drepana acuta <$ Butler
Type [in Butler's handwriting] ; Drepanidae genitalia slide No. 325.

japonibia. Syntype, Nikko [Japan]. In the Zoologisches Museum, Berlin.

urupitla. Holotype $, [Kuril Is.], L'rup, Kopune. In the Xaturhistoriska
Riksmuseet, Stockholm.

koreula. Holotype $, Korea. In the Xaturhistoriska Riksmuseet, Stockholm.

Drepana rufofasciata Hampson

(PI. 11, fig. 365 ; Text-figs. 200-202)

Drepana rufofasciata Hampson, [1893] : 334.

Drepana rufofasciata Hampson ; Warren, 1922 : 463. [Good figs.]

Drepana rufofasciata Hampson ; Gaede, 193 1 : 27.

This species is externally closest to pallida Moore. It can be distinguished from
the latter by the more reddish medial shade on the fore wing and by the less well
marked postmedial fascia, which is parallel to the subterminal fascia on the fore
wing (unlike pallida). Small differences in the male and female genitalia separate
rufofasciata from pallida and dispilata Warren both of which are closely allied to
rufofasciata.

Distribution. The range of rufofasciata includes Sikkim, and China. The few
Chinese specimens listed below are considerably paler than the Sikkim material but
no significant differences in the genitalia or other characters appear to be present.

Material examined. Type. LECTOTYPE <$, here selected, labelled : Sikkim,
Interior, Moller, Drepana rufofasciata Hmpsn. type J ; Collectio H. J. Elwes ;
Rothschild Bequest B.M. 1939-1 ; Drepanidae genitalia slide No. 687. In the
BM(NH).

Other material. BM(NH). Sikkim: 2 J, 1$ {Moller) ; 1 <$, Tonglo, 10,000 ft.,
vii.1886 (Elwes). China, Tibet : 2 <J, 2 $, Yatung (Hobson) ; 1 <j>, Chumbi valley,
Dopenri.

Drepana pallida Moore

(PI. ii, figs. 360-361 ; Text-figs. 203-214)

Drepana pallida Moore, 1879 : 84.

Drepana pallida Moore ; Warren, 1922 : 463.

Drepana pallida Moore ; Gaede, 1931 : 27.

io6

A. WATSON

Readily distinguished from the closely related dispilata, and rufofasciata by the
colour-pattern and the genitalia.

Four subspecies are known : the nominate subspecies (India, Burma), cretacea
(China, Vietnam), flexuosa (China) and nigromaculata Okano (Formosa). An
account of each of these is given to facilitate comparison between the Chinese and
non-Chinese subspecies.

Figs. 200-202. Drepana rufofasciata, genitalia. 200, $ ; 201, <$ ; 202, aedeagus.

CHINESE DREPANINAE 107

Drepana pallida pallida Moore

(Text-figs. 203-207)

Drepana pallida Moore ; Warren ; 1922 : pi. 48I. [Good figs. <J, ?•]

Distinguished from the other three supspecies by the colour-pattern and by the
male and female genitalia (see plate in Warren (1922) and Text-figs. 203-207).
Distribution. N.E. India and N. Burma.

Type. I select as LECTOTYPE a <$ syntype in the collection of the BM(NH)
labelled : Moore Coll. 94-106 ; Darjiling <$ ; Drepana pallida Moore (Type) <$ [in
Moore's handwriting].

Drepana pallida cretacea Hampson stat. n.
(PI. 11, fig. 360 ; Text-figs. 208, 209)

Drepana cretacea Hampson, 1914 : 107.
Drepana cretacea Hampson ; Gaede, 1932 : 168.

Externally, cretacea differs from the nominate subspecies in the following respects:
fore wing less strongly falcate ; dark spot at posterodistal angle of cell generally
larger ; postmedial fascia nearly straight, not arcuate ; proximal line of postmedial
fascia touching or nearly touching distal end of cell in hind wing.

The male genitalia are similar to those of the nominate subspecies but have a
differently shaped eighth abdominal sternite (Text-fig. 208). The female genitalia
are characterized by the differently shaped ninth abdominal tergum. (Text-fig.
209).

Distribution. China (Szechwan), Vietnam.

Material examined. Type. Holotype $, [China, Szechwan], Huang-Mu-Chang,
7000 ft., vii.1889 {Pratt). In the BM(NH).

Other material. BM(NH). China, Szechwan : 2 $, Ta-tsien-lou, 1897, 1906 ;
3 cS Shin-kai-Si, Mt. Omei, i.viii.1921 (Franck) ; 2 $, Tien-tsuen, 1897 (Dejean) ;
2 $, Tien-tsuen, Yuin-kin, 1899 ; 2 $, Kwanhsien 12.vii.1925, vii.1930 ; 3 <$, 4 $,
Siao-lou, 1899-1903 ; 1 <$, Moupin, 1898 ; 1 $, Pa-tse-fang, 1893. U.S. National
Museum. China, Szechwan : 3 <$, Mt. Omei.

Drepana pallida flexuosa ssp. n.

(PI. 11, fig. 361 ; Text-figs. 210-212)

This is readily distinguished from the other subspecies of pallida by the more strongly arcuate
fore wing, and by the more strongly marked cell-spots and more strongly arcuate postmedial
fascia on the fore wing. It differs from the nominate subspecies by the fact that the proximal
line of the postmedial fascia on the hind wing touches or nearly touches the distal end of the
cell. In the male genitalia flexuosa differs from the nominate subspecies in the shape of the
valves, gnathus and eighth abdominal sternite (Text-figs. 210, 211). The female genitalia
are similar to those of the nominate subspecies but possess a distinctive ninth tergum (Text-fig.
212).

Wing, o* 2 3 - o mm. (1) ; $ 26-5-31-0 mm. (2).

io8

A. WATSON

Figs. 203-209. Drepana, genitalia. 203-207, pallida pallida. 203, $ ; 204, aedeagus ;
205, 6* eighth sternite ; 206, $ ; 207, $ ninth segment (dorsal view). 208, 209, pallida
cretacea. 208, $ eighth sternite ; 209, $ ninth segment, (dorsal view).

CHINESE DREPANINAE 109

Distribution. China (Fukien, Chekiang).

Holotype <$. [China :] Fukien, Kuatun, 230 m., 3JV.1938 (Klapperich). In
the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 1 $, type-locality, 2.vii.ig38
(Hone) ; 1 $, Chekiang, West Tien-mu-shan, 12. iv. 1932 (Hone).

Drepana pallida nigromaculata Okano

(Text-figs. 213, 214)

Drepana pallida nigromaculata Okano, 1959 : 38.

I have examined only two males of this Formosan subspecies. One of these
specimens had been identified and compared with Okano material in Japan by Dr.
H. Inoue. They resemble specimens of cretacea most closely in wing shape and
colour-pattern, but the genitalia are most like those of flexuosa.-

Type. [Not seen.] Holotype <J, central Formosa, Puli-Nushe, v. 1958. In the
Okano collection.

Drepana dispilata Warren

(PI. 11, figs. 362, 364 ; Text-figs. 215-219)
Drepana dispilata Warren, 1922 : 463. [Fig.]

Distinguished from pallida, to which dispilata is probably most closely allied, by
the two large cell-spots on the fore wing and by the genitalia of both sexes (Text-
figs. 215-219).

Three subspecies are known : the nominate subspecies (India, Sikkim, Burma),
rufata (China), and grisearipennis (China).

Drepana dispilata dispilata Warren
(Text-figs. 215-217)

Drepana dispilata Warren ; Gaede, 1931 : 26.

Drepana x-z-nigrum Bryk, 1943 : II. Holotype $, [N.E. Burma], Kambaiti, 7000 ft ; in the
Naturhistoriska Riksmuseet, Stockholm. [Good figs.] syn. n.

The male genitalia (Text-figs. 215-217) and the less strongly falcate fore wing
separate this subspecies from the two Chinese subspecies.
Known from N. India, Sikkim and N. Burma.

Type. LECTOTYPE $, here selected, labelled : Knyvett ; Collectio H. J.
Elwes ; Drepana dispilata Warr. Type $ ; Drepana dispilata Warr. $ ; Rothschild
Bequest B.M. 1939-1 ; Drepanidae genitalia slide No. 684. In the BM(NH).

A. WATSON

Figs. 210-214. Drepana, genitalia. 210-212, pallida flexuosa. 210, <$ ; 211, 0* eighth
sternite ; 212, $ ninth segement (dorsal view). 213, 214, pallida nigromaculata, $. 213,
o* ; 214, eighth sternite.

CHINESE DREPANINAE m

Drepana dispilata rufata ssp. n.

(PI. ii, fig. 362 ; Text-figs. 218, 219)

Distinguished from the nominate subspecies by the more strongly falcate fore wing (PI. 11,
fig. 362) and by the male genitalia (Text-fig. 218). The female genitalia are figured in Text-
fig. 219.

Win^ J 18-5-2 1 -5 mm. (12) ; $ 20-0-21-5 mm - U)-

Distribution. China (Yunnan, Shensi).

Holotype <$. [China:] S. Shensi, Tapaishan im Tsinling, 26. vi. 1935 (Hone) ;
Drepanidae genitalia slide No. 689. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 9 g and 3 $, S. Shensi, Tapaishan
im Tsinling, 22.vi.-2.vii.1935 (Hone) ; 3 <J and 1 $, N. Yunnan, Likiang, 2000 m.,
19-27. iv. 1935 (Hone). Daniel Collection, Munich. China : 1 £, S. Shensi,
Tapaishan im Tsinling, 22. vi. 1935 (Hone).

Drepana dispilata grisearipennis Strand stat. n.
(PI. 11, fig. 364)

Drepana grisearia Leech, 1898 : 365. [A junior homonym.]

Drepana grisearipennis Strand, 191 1 : 201 ; as a replacement name for Drepana grisearia

Leech, preoccupied by Drepana grisearia Staudinger, 1892 : 335.
Drepana grisearipennis Strand ; Gaede, 1931 : 26.

The only available example of this species, the holotype, differs in only minor
respects from the material of rufata, both externally and in the genitalia. Further
material from Szechwan should show whether or not rufata and grisearipennis are
synonymous.

Distribution. China (Szechwan).

Material examined. Type. Holotype $, Pu-tsu-fong, 9820 ft., vi, vii.1890. In
the BM(NH).

TRIDREPANA Swinhoe

Tridrepana Swinhoe, 1895 : 3. Type-species Drepana albonotata Moore, 1879 : 83, by sub-
sequent designation by Warren, 1922 : 464.
Tridrepana Swinhoe ; Watson, 1957 :4ir. [Revision.]
' Iridrepana ' ; Warren, 1922 : 464. [An incorrect subsequent spelling of Tridrepana.]

As stated in an earlier paper (Watson, 1957 : 436) there is some doubt concerning
the whereabouts of the type material of albonotata, the type-species of Tridrepana.
T. albonotata was described from material in the Atkinson collection, now in the
Zoological Museum at Berlin, and I think it is best to select a female specimen from
the Atkinson collection, labelled ' Parisnath ' (the type-locality), as the LECTO-
TYPE, even though it lacks Moore's usual handwritten label indicating its status as
a type. The specimen labelled ' Drepana albonotata Moore, type ', in the BM(NH),
collected in Darjeeling, was presumably labelled by Moore subsequent to the
preparation of the original description of albonotata.

A. WATSON

Figs. 215-219. Drepana, genitalia. 215-217, dispilata dispilata, $. 215, aedeagus
216, $ ; 217, eighth sternite. 218, 219, dispilata rufata. 218, 3 valves and saccus
219, $. Fig. 220. Thymistadopsis undulifera, $ genitalia.

CHINESE DREPANINAE 113

Tridrepana is closely allied to Drepana Schrank. There is a reasonable degree of
concordance externally and in the genitalia.

Distribution. Manchurian Subregion & Oriental Region (see Table 1). With
the exception of fulva, no new material of any significance has been discovered
since the last revision of Tridrepana (Watson, 1957) and reference should be made
to this paper for descriptions and figures of the Chinese species. The following is a
list of the species and subspecies known to occur in China.

Tridrepana fulvata (Snellen), 1876 : 19. N. India, Burma, Malaysia, Indonesia
and China.

Tridrepana fulvata ssp. China (Hainan Is., Kwangtung), Hong Kong.
Tridrepana arikana (Matsumura), 1921 : 949. Bhutan, China, Formosa.

Tridrepana arikana arikana (Matsumura) Formosa, and probably mainland
China.
Tridrepana crocea (Leech), 1888 : 649. Japan, China (Chekiang, Fukien,

Hunan).
Tridrepana unispina Watson, 1957 : 458. Formosa and China (Fukien,

Yunnan).
Tridrepana finita Watson, 1957 : 480. China (Tibet, Szechwan, Yunnan).
Tridrepana rubromarginata (Leech), 1898 : 365. Sikkim, Bhutan, Xepal,
China.

Tridrepana rubromarginata rubromarginata (Leech). China (Szechwan,
Yunnan).
Tridrepana thermopasta (Hampson), 1914 : 106. China (' W. China ', prob-
ably Szechwan).
Tridrepana maculosa Watson, 1957 : 488. China (Szechwan, Yunnan).
Tridrepana marginata Watson, 1957 : 490. China (Szechwan, Yunnan).
Tridrepana fulva (Hampson), [1893] : 342. Sikkim, China (E. Tibet,
Szechwan — 1 $, Tu-pa-keo, in the BM(NH)). Contrary to inference in the
type-citation in Watson (1957 : 489), Hampson did not designate a holotvpe.
I therefore select as LECTOTYPE the J syntype in the BM(XH), labelled :
Sikkim, 1300 ft., Jongri, 1887 ; Drepanidae genitalia slide No. 446 ; B.M.
negative No. 15463.

CALLIDREPANA Felder
(PI. 12, figs. 369-372 ; Text-figs. 221-245)

Callidrepana Felder, 1861 : 30. Type-species Callidrepana saucia Felder, 1861 : 31, by mono-

typy.
Callidrepana Felder ; Gaede, 1931 : 34.
Callidrepana Felder ; Watson, 1965 : 149.
Damna Walker, [1863] : 1570. Type-species Damna gelidata Walker, [1863] : 1570 by mono-

typy- [Synonymized by Warren, 1922 : 471, by transference of type-species.]
Ausaris Walker, [1863] : 1632. Type-species Ansaris scintillata Walker [1863] : 1632, by

monotypy. [Synonymized by Kirby, 1892 : 730.]
Ticilia Walker, 1865 : 394. Type-species Ticilia argentilinea Walker, 1865 : 394, by monotypy.

[Synonymized by Warren, 1922 : 471, by transference of type-species.]

ii 4 A. WATSON

Drepanulides Motschulsky, 1866 : 192. Type-species Drepanulides palleolus Motschulsky,

1866 : 193, by subsequent designation by Inoue, 1962 : 32. [Synonymized by Kirby,

1892 : 730, by transference of type-species.]
Drepanula Gaede, 1914 : 65. Type-species Drepanula argyrobapta Gaede, 1914 : 65, by

monotypy. [A junior homonym of Drepanula Frolich, 1828 : n.]
Drepanulina Gaede, 1927 : 287. Type-species Drepanula argyrobapta Gaede, 1914 : 65, by

monotypy. [A replacement name for Drepanula Gaede, 1914 : 65]. [Synonymized by

Watson, 1965 : 149.]

This genus is distinguished from all other genera of Drepanidae, except Macrocilix
Butler and Tridrepana Swinhoe, by the presence of widely distributed brilliantly
lustrous scales on the upper surface of the wings. (In Tridrepana, one bright yellow
species, melliflua Warren, has similar lustrous scales on the wings. Macrocilix is
a distinctive genus in that the ground-colour of the wings of all the species is white) .

Callidrepana includes over twenty species. It ranges from India to Japan, and
through the Malay Archipelago and New Guinea to the Solomons. Three African
species are known (see Watson, 1965 : 149). Four species occur in China :
hirayamai, pair ana, ovala and gemina (the latter two described here as new). (See
Table 1.)

Key to the Chinese Species of CALLIDREPANA
Both Sexes

Postmedial fascia on upper surface of both wings simple ; ground-colour very pale
yellowish white ; dark marking (where present) at distal end of cell on fore wing
small, ovate (PI. 12, figs. 370, 371). Genitalia as in Text-figs. 232-238

gemina (p. 119)

Postmedial fascia on upper surface of both wing double ; ground-colour buff or
buffish white ; marking (where present) at distal end of cell small and ovate, or large
and rectangular (PI. 12, fig. 372). Genitalia not as in gemina .... 2

Upper surface of fore wing with large, dotted, rectangular marking at distal end of

cell. (PI. 12, fig. 372). Genitalia as in Text-figs. 239-245 . hirayamai (p. 121)

Upper surface of fore wing with small ovate marking at distal end of cell, or fore wing

without cell-marking. Genitalia not as in hirayamai ..... 3

Genitalia as in Text-figs. 221-227 ...... patrana (p. 114)

Genitalia as in Text-figs. 228-231 ....... ovata (p. 117)

Callidrepana patrana (Moore)

(Text-figs. 221-227)

Drepana patrana Moore, [1866] : 816.

This species is separable from ovata, which is closely allied to patrana and forms a
superspecies with it, by the less elongate cell-marking (where present) on the fore
wing, the slightly more strongly falcate fore wing and by small but distinct differ-
ences in the genitalia of both sexes (see ovata). The male genitalia of argenteola
Moore (1858 : 369) (India, Burma, Formosa, Malaysia and Indonesia) indicate that
though this species is externally similar to patrana, except for the more strongly

CHINESE DREPANINAE

Figs. 221-226. Callidrepana, genitalia. 221-224, patrana patrana. 221, $ ; 222,
aedeagus ; 223, $ ; 224, <$ eighth tergite and sternite. 225, 226, patrana palleolus, <J.
225, £ ; 226, eighth sternite.

n6 A. WATSON

falcate fore wing and the narrower cell-marking on the fore wing, it is probably not
closely allied to patrana.

Two subspecies are known : the nominate subspecies, known from India and
China, and palleolus recorded only from Japan.

Callidrepana patrana patrana (Moore)

(Text-figs. 221-224)

Drepana argenteola var. patrana Moore ; Strand, 191 1 : 202.

Callidrepana patrana (Moore) Warren, 1922 : 471.

Callidrepana patrana (Moore) ; Gaede, 1931 : 36.

Callidrepana patrana Inoue, 1962 : 32.

Callidrepana patrana formosana Inoue, ig$5 : 13. [Elevation to subspecific rank of ab. formosana

Matsumura, 1921 : 945] syn. n. [Synonymy anticipated by Inoue, 1962 : 32.]
Callidrepana patrana subbasalis Bryk, 1943 : 21. [Fig.] syn. n. [Synonymy anticipated by

Inoue, 1962 : 32.]

The genitalia of both sexes provide the only reliable diagnostic features in
separating this subspecies from the dark-spotted form of palleolus.

Material examined. Types, patrana. The syntypes (all males) were deposited
in the collections of A. E. Russell and Moore. The Russell collection is almost
certainly lost (see Horn and Kahle, 1937 : 380), and there are no syntypes in the
Moore collection in the BM(NH). I have therefore selected a female specimen in the
BM(NH) as the NEOTYPE, labelled : Darjiling (F. Moller) ; Rothschild Bequest
B.M. 1939-1 ; Drepanidae genitalia slide No. 993.

subbasalis. Holotype <$, N.E. Burma, Kambaiti, 2000 m. ; Drepanidae genitalia
slide No. 1683. In the Naturhistoriska Riksmuseet, Stockholm.

formosana. [Type not seen. Examples of formosana identified by Inoue have
been studied.]

Other material. N.E. India, N. Burma, Formosa and China (Szechwan, Yunnan,
Fukien, Chekiang). A single male from Laos, in the BM(NH), may prove to re-
present this subspecies.

Callidrepana patrana palleolus (Motschulsky)
(Text-figs. 225-227)

Drepanulides palleolus Motschulsky, 1866 : 193.
Drepana palleolus (Motschulsky) Strand, 1911 : 202.
Drepana palleolus (Motschulsky) ; Gaede, 1931 : 27.
Callidrepana palleolus (Motschulsky) Nagano, 1917 : 36.

Callidrepana patrana palleolus (Motschulsky)
Callidrepana patrana palleolus (Motschulsky)
Callidrepana patrana palleolus (Motschulsky)
Callidrepana patrana palleolus (Motschulsky)

Inoue, 1955 : 13.

Inoue, 1956 : 368.

Inoue, 1959 : 175. [Good figs.]

Inoue, 1962 : 32. [Good figs.]

Inoue (1955 : 13) gave the name crassimaculata to the form with an elongate dark
spot at the distal end of the cell on the fore wing. This has been figured by Inoue

CHINESE DREPANINAE 117

(1959, 1962) together with the paler unspotted f. palleolus Motschulsky (= ab.
simplificaria Strand, 191 1 : 202).

The unspotted form of this subspecies is easily distinguished from the nominate
subspecies by the colour-pattern, but the male and female genitalia provide the
only significant diagnostic characters in the dark-spotted form.

Distribution. Japan (see Inoue, 1956).

Type. According to investigations made for me by Dr. K. M. Efron and Dr. A. S.
Danislevskyi in the U.S.S.R., all the Lepidoptera from the Motschulsky collection
have been lost. There seems little doubt, however, from the original description
that Nagano (1917) and Inoue (1959) have correctly identified this subspecies.

Callidrepana ovata sp. n.

(PI. 12, fig. 369 ; Text-figs. 228-231)

d. Front of head dark reddish brown ; vertex, palp and bipectinate antenna dull yellowish
orange ; collar yellowish white.

Thorax buff or yellowish white. Colour-pattern of upper surface of wings as in PI. 12,
fig. 369 ; colours similar to the pale form of C. patrana palleolus but usually paler ; scattering
of brilliantly lustrous scales present along veins proximal to postmedial fascia, along distal
edge of postmedial fascia and in streak in basal half of costal area. In fore wing R\ arises
from near distal end of cell, Ro from near distal end of areole ; Rs and Afj from a point or on
short stalk. Sc + R\ approximates to Rs for short distance distal to end of cell in hind wing.
Under surface of both wings pale yellowish buff, but fore wing more brownish basally ; fore
wing usually with trace of reddish brown postmedial fascia anteriorly, corresponding in position
to proximal part of this fascia on upper surface ; hind wing usually with trace of reddish
brown postmedial fascia, corresponding in position to distal part of this fascia on upperside.
Legs buff, but with outer surface of prothoracic tibia and tarsus brownish buff.

Abdomen similar in colour to adjacent surface of hind wing.

o" genitalia as in Text-figs. 228, 230, 231.

$. As for male but with shorter antennal pectinations.

Wing. 6* 15-5-19-0 mm. (45) ; 9 17-5-22-0 mm. (15).

The cell-spot on the fore wing in most specimens of this species is less elongate
than that in the closely allied patrana and the fore wing is slightly more strongly
falcate. The male and female genitalia also distinguish ovata from patrana : in the
male the aedeagus is more slender apically and the socii less strongly arcuate
apically, while in the female the signum is over twice as long as in patrana.

There is some individual variation in the postmedial fascia of the upper surface
of the wings, which in a few males and one female is much broader than in the
illustrated specimen, as in patrana palleolus (q.v.).

Distribution. The sympatry which exists between this species and patrana at
Kwanhsien (Szechwan province) indicates that they are probably specifically
distinct. The distribution and morphological similarity between ovata and patrana
suggest a superspecific relationship (Artenkreis). Known from the following
Chinese provinces : Shensi, Hupeh and Szechwan.

Type. Holotype <$. China : S. Shensi, Tapaishan im Tsinling, 13.vii.1936
(Hone) ; Drepanidae genitalia slide No. 1009. In the Museum Koenig, Bonn.

n8

A. WATSON

Figs. 227-231. Callidrepana, genitalia. 227, patrana palleolus, <j>. 228-231, ovata.
228, aedeagus ; 229, £ ; 230, $ ; 231, $ eighth tergite and sternite.

CHINESE DREPANINAE 119

Paratypes. Museum Koenig, Bonn. China : 50 <$, 14 $, S. Shensi, Tapaishan
im Tsinling, 16.v-28.viii.1935, 1936 {Hone). BM(NH). China : 6 <$, 1 $, S.
Shensi, Tapaishan im Tsinling, 23.vi-25.vii.1935, 17.v-29.vii.1936 (Hone). 1 $,
Ichang, viii.1888 ; 1 <J, 1 $, Kwanhsien, 8.viii.i926, viii.1930 (Franck) ; 1 <J, 2 $,
Chang Yang, vi-vii.i

Callidrepana gemina sp. n.

(PI. 12, figs. 370, 371 ; Text-figs. 232-238)

£, $. Front of head dark reddish brown dorsally, dull yellowish orange ventrally ; vertex
and antenna dull yellowish orange ; outer surface of palp yellowish brown. Antenna bipec-
tinate. Collar whitish.

Thorax very pale yellow or yellowish white. Ground-colour of upper surface yellowish
white ; colour-pattern as in PI. 12, figs. 370, 372 ; apical markings, cell-spot and subterminal
spots dark reddish brown, other markings yellowish brown ; costa dull yellowish orange at
base ; both wings with bands of brilliant l\ lustrous scales along veins proximal to postmedial
fascia, along distal edge of postmedial fascia and along basal half of costa. Cell-spot sometimes
reduced in size, completely absent in one male examined. Venation of both wings as for
ovata (q.v). Under surface of both wings slightly paler than upper surface, but fore wing
yellowish brown antero-proximally and with dark reddish brown crescent near apex at outer
margin ; trace of postmedial fascia on hind wing ; well-marked reddish brown discocellular
spot. Legs pale yellow but with front surface of tibia and tarsus of prothoracic leg greyish
brown and femur yellowish orange.

Abdomen similar in colour to adjacent surface of hind wing.

Genitalia as in Text-figs. 232-238. Of special diagnostic importance in the male are the
heavily sclerotized valves, the broad eighth tergite and the shape of the aedeagus.

The single postmedial fascia on the upper surface of both wings and the more
proximal position of the postmedial fascia on the hind wing distinguish gemina from
its closest relatives patrana and ovata. The genitalia of both sexes are also character-
istic. The chief diagnostic characters in the male genitalia can usually be seen
without dissection.

Two subspecies are known : the nominate subspecies (India) and curta (China).

Callidrepana gemina gemina ssp. n.

(PI. 12, fig. 371 ; Text-figs. 232-235)

Distinguished from curta by the broader, reddish brown crescentic marking at the outer
margin near apex of fore wing and by the male genitalia, especially the eighth tergite and
sternite, aedeagus, valves and socii.

The female genitalia is figured in Text-fig. 235.

Wing, o* 150-17-0 mm. (6) ; ? 17-5-20-0 mm. (5).

Distribution. Only known from the type locality (N.E. India). A single male
in the BM(NH) from Szechwan (China), which lacks a cell-spot on the fore wing
may prove to represent this subspecies.

A. WATSON

Holotype <J. Darjeeling, Gopaldhara, 4720 ft. (Stevens) ; Drepanidae genitalia
slide No. 994. In the BM(NH).

Paratypes. BM(NH). N.E. India : 5 J, 5 $. Darjeeling, Gopaldhara, 4720 ft.,
ix.1916-ix.1918 (Stevens).

Figs. 232-235. Callidrepana gemina gemina, genitalia. 232, $ ; 233, aedeagus ; 234,
$ eighth tergite and sternite ; 235, $.

CHINESE DREPANINAE 121

Callidrepana gemina curta ssp. n.

(PI. 12, fig. 370 ; Text-figs. 236-238)

<J. Distinguished from the nominate subspecies by the narrower, dull yellowish orange
apical marking at the outer margin of the fore wing and by the male genitalia (Text-figs. 236-

238).

$. Not known.

Wing. <$ 160-170 mm. (5).

Distribution. China (Kwangtung, Fukien, Chekiang).

Holotype <$. China : Fukien, Kuatun, 16. v. 1938 (Klapperich) ; Drepanidae
genitalia slide No. 987. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 2 <$, Kwangtung, Linping,
iv.viii.1922 {Hone) ; 1 <$, Chekiang, W. Tien-mu-shan, 27.vii.1932 {Hone). BM{NH).
China : 1 <$, Hupeh, Chang Yang, vi.1888 {Pratt) ; 1 <$, Kwangtung, Linping,
viii.1922 {Hone). Zool. Museum, Berlin. China : 2 <$ {Mell) ; 1 £, Canton,
Gao Fung {Mell).

Callidrepana hirayamai Nagano
(PI. 12, fig. 372 ; Text-figs. 239-245)
Callidrepana hirayamai Nagano, 191 7 : 492. [Fig]

The identity of this species was revealed by Inoue (1961). Through the kindness
of Dr. H. Inoue I was able to borrow a Japanese male of this species and to compare
it with the Chinese material at Bonn and in the collection of the BM(NH).

The colour-pattern separates this species from ovata and patrana which it resembles
to some extent externally : the dotted rectangular marking at the end of the cell
on the fore wing is particularly diagnostic. The coloration is similar to patrana,
but in many specimens the ground-colour is a dull greyish yellow as in the holotype
of hirayamai forcipulata. The presence of an uncus, and the shape of the valves
and socii are characteristic features of the male genitalia.

Two subspecies are known : the nominate subspecies (Japan) and forcipidata
(China).

Callidrepana hirayamai hirayamai Nagano

(Text-figs. 239-241)

Callidrepana hirayamai Nagano, 1917 : 492. [Fig.]

Callidrepana hirayamai Nagano ; Inoue, 1956 : 370.

Callidrepana hirayamai Nagano ; Inoue, 1961 : 9. [Good fig.] [Translation into English of

original description. Distribution.]
Callidrepana hirayamai Nagano ; Inoue, 1962 : 33. [Good fig.]
Callidrepana yaknshimalis Yamamoto, i960 : 334. [Good figs.] [Synonymized by Inoue,

1961.]

Separable from forcipulata by the male genitalia. As only one specimen of the

A. WATSON

Figs. 236-241. Callidrepana, <$ genitalia. 236-238, gemina curta. 236, $ ; 237,
aedeagus ; 238, eighth tergite and sternite. 239-241. hirayamai hirayamai. 239,
eighth tergite and sternite ; 240, <$ ; 241, aedeagus.

CHINESE DREPANINAE

123

nominate subspecies has been seen (a male in the Inoue collection), no reliable
comparison of external characters could be made but there appears to be little to
distinguish the two subspecies.

Distribution. Japan (see Inoue 1956, 1961, 1962).

Types, hirayamai. Holotype <J, [Japan], Kiso-yama, Shinano, 3.VU.1915
(Hirayama). Stated by Inoue (1956) to be probably lost.

Figs. 242-245. Callidrepana hirayamai forcipulata, genitalia. 242, $ eighth tergite and
sternite ; 243, <J ; 244, aedeagus ; 245, $.

I2 4 A. WATSON

yakushimalis. Holotype $, Japan, Kosugidani, Yakushima, 5.viii.ig48 (Yama-
moto). In Yamamoto collection. [Not seen.]

Callidrepana hirayamai forcipulata ssp. n.

(PI. 12, fig. 372 ; Text-figs. 242-245)

Distinguished from the nominate subspecies by the male genitalia (in particular the shape of
the eighth tergite and sternite, valves and uncus.

Wing. cJ 13-5-18-0 mm. (15) ; ? 16-5-19-0 mm. (5).

Distribution. China (Fukien, Hunan). A single male from Ta-tsien-lou (Szech-
wan), in the BM(NH), and a male from Wenchow (Chekiang) and four females from
West Tien-mu-Shan (Chekiang) in the Museum Koenig, Bonn, may also prove to
represent this subspecies.

Holotype $. China : Hunan, Hoeng-Shan, 17.vii.1933 {Hone) ; Drepanidae
genitalia slide No. 997. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 10 (J, 5 ?, Hunan, Koeng-Shan
900 m., 24.iv-11.xi.1933 (Hone) ; 4$, 1 $, Fukien, Kuatun, 2300 m., 5.iv-2.vii.i938
(Hone, Klapperich).

DRAPETODES Guenee

(PI. 12, fig. 373)

Drapetodes Guenee, 1857 : 424. Type-species Drapetodes mitaria Guenee, 1857 : 424, pi. 18,

fig. 6, by monotypy. Type-locality : central India.
Drapetodes Guenee ; Gaede, 1931 : 14.

Drapetodes is characterized chiefly by the long labial palps, the non-falcate fore wing, the
colour-pattern, the absence of an areole on the fore wing, and by the broad bifid uncus in the
male genitalia.

This small, chiefly Indo-Malayan genus, is represented in Formosa and the Chinese
provinces of Szechwan and Kwangtung. The specific identity of the material in the
BM(NH) is doubtful. For example, the male from Chung-king (Szechwan) probably
represents mitaria Guenee, while the female from Canton (the only other specimen
seen from the mainland of China) could prove to be a specimen of either mitaria, or
circumscripta Warren (1922 : 460, pi. 481) (type-locality : Sumatra), or possibly
deumbrata Warren (1922 : 459, pi. 481) (type-locality : Bali).

THYMISTADOPSIS Warren

(PI. 12, fig. 374 ; Text-figs. 220, 246-250)

Thymistadopsis Warren, 1922 : 461. Type-species Problepsidis albidescens Hampson, 1895 : 288

[Good fig.], by monotypy.
Thymistadopsis Warren ; Gaede, 1931 : 16.

No species other than the type-species has previously been placed in this genus,
the nearest relative of which is possibly Leucoblepsis Warren (1922 : 462).

Distribution, albidescens is known only from the type-locality, in Sikkim, and

CHINESE DREPAXIXAE

125

Assam (N.E. India) ; trilinearia has a similar range but occurs in China ; undulifera
is known only from Tibet at an altitude of 10,500 ft.

Thymistadopsis trilinearia (Moore) comb. n.

(PI. 12, fig. 374 ; Text-figs. 246-250)

Drepanodes trilinearia Moore, 1867 : 618.

Externally trilinearia is readily separable from the type-species by the absence
of an outer marginal process on the fore wing, by the straight or weakly lunulate

Figs. 246-250. Thymistadopsis , genitalia. 246-249, trilinearia trilinearia. 246, o* ; 247,
aedeagus ; 248, o* eighth tergite and sternite ; 249, 9 (ostium and anal papillae). 250,
trilinearia pulvis, $ (ostium and anal papillae).

126 A. WATSON

postmedial fascia on the fore wing, and by the pale postmedial fascia and the
anastomosis oi Sc -{- R\ with Rs for a short distance distal to the cell in the hind
wing. The genitalia of both sexes are closely similar to those of albidescens but are
distinguishable by differences in the proportions of the various parts.

Two subspecies are known : the nominate subspecies (Sikkim and N.E. India),
and pulvis (China) .

Thymistadopsis trilinearia trilinearia (Moore)
(Text-figs. 246-249)

Drepanodes trilinearia Moore, 1867 : 618.

Drepana trilinearia (Moore) Hampson, [1893] : 33^-

Drepana trilinearia (Moore) ; Hampson, 1897 : 288.

Drepana trilinearia (Moore) ; Warren, 1922 : 467. [Good fig., as ' bilinearia '.]

Drepana trilinearia (Moore) ; Gaede, 1931 : 30.

Tridrepana trisulcata Warren, 1896 : 340. [Synonymized by Hampson, 1897 : 288.]

Tridrepana trisulcata Warren ; Gaede, 1931 : 31.

In some females the area between the antemedial fascia and the postmedial fascia
on the fore wing is heavily suffused with dark brown. In the remaining females
and all the males studied this area is greyish white.

Separated from pulvis by the ostium in the female genitalia. (The male of pulvis
is not known.)

Distribution. Sikkim and N.E. India.

Type. LECTOTYPE <$, here selected, labelled : Darjeeling ; Moore Coll.
94-106 ; Drepanidae genitalia slide No. 745. In the BM(NH).

Thymistadopsis trilinearia pulvis (Oberthiir) stat. n., comb. n.

(PI. 12, fig. 374 ; Text-fig. 250)

Drepana pulvis Oberthiir, 1916 : 375.

Drepana pulvis Oberthiir, 1917 : pi. 428. [Good fig.]

Albara pulvis (Oberthiir) Gaede, 1933 : 169.

The type is the only known specimen apart from a doubtfully identified female
from Tu-pa-keo (Szechwan) in the BM(NH).

Distinguished from the nominate subspecies by the female genitalia.

Type. Holotype $, China, Szechwan, Siao-Lou, 1903 (Chasseurs indigenes du
P. Dejean) ; Drepanidae genitalia slide No. 746. In the BM(NH).

Thymistadopsis undulifera (Hampson) comb. n.

(Text-fig. 220)

Drepana undulifera Hampson, 1900 : 228. [Good fig.]
Drepana undulifera Hampson ; Strand, 191 1 : 202.
Drepana undulifera Hampson ; Gaede, 1931 : 28.

Until male specimens become available the affinities of undulifera will probably

CHINESE DREPANINAE 127

remain doubtful. However, an examination of the female genitalia and of external
characters such as the shape of the antennae, the wing shape, the colour-pattern and
the coloration has shown no great discordance between undulifera and the other
two species of Thymistadopsis except for the rather abbreviated colour-pattern of
the upper surface.

Distribution. China (Tibet).

Material examined. Type. LECTOTYPE $ [not $ as stated by Hampson,
1900], here selected, labelled : Yatung [10,500'], Tibet, A. E. Hobson, 98-201 ;
Drepanidae genitalia slide No. 1707. In the BM(NH).

Paralectotypes. BM(NH). China : 2 9. type locality.

DEROCA Walker

(PI. 12, fig. 375)

Deroca Walker, 1855 : 822. Type-species Deroca hyalina Walker, 1855 : 823, by monotypy.
Deroca Walker ; Watson, 1957a : 129. [Revision.]

Probably most closely allied to Callicilix Butler but distinguished from it by the

colour-pattern and the lack of yellow coloration on the wings, differences in the

male genitalia, and the presence of well-developed mid tibial spurs on the hind leg.

Distribution. N. India, N. Burma, China, Formosa, Korea and Japan. (See

Table I.)

This genus has been revised by the present author (Watson, 1957a) and no
further material has been made available since this revision. The following list
includes the species and subspecies known to occur in China together with their
known distribution. (All four described species of Deroca occur in China.)
Deroca hyalina Walker, 1855 : 823. India, Burma, China.

D. hyalina latizona Watson, 1957a : 134. China (Kwangtung, Szechwan,
Hunan, Kiangsi, Fukien, Chekiang). See Plate 12, fig. 375.
Deroca hidda Swinhoe, 1900 : 306. Sikkim, N. India, Burma, China.
D. hidda bifida Watson, 1957a : 137. X. India, China (N. Yunnan).
Deroca pulla Watson, 1957a : 139. China (Szechwan, Hupeh).
Deroca inconclusa (Walker), 1856 : 1727. N. India, Burma, China, Formosa,
Korea, Japan.
D. inconclusa inconclusa (Walker). N. India, Burma, China (Szechwan,

Yunnan).
D. inconclusa carinata Watson, 1957a : 143. China (Shensi).

CALLICILIX Butler

(PI. 12, figs. 376, 377 ; Text-figs. 251-257)

Callicilix Butler, 1885 : 124. Type-species Callicilix abraxata Butler, 1885 : 124, by monotypy.

Platypteryx nguldoe Oberthur, 1893, the only species subsequently added to
Callicilix, has been relegated to subspecific rank.

128 A. WATSON

The genus is distinguished, in both sexes, from the closely related Auzata Walker
by the bipectinate antennae and, as already indicated by Inoue (1962), by the fact
that the outer spur of the middle pair on the hind tibia is very small and partly
hidden by scales.

Distribution. Japan, China and N.W. India.

Callicilix abraxata Butler
(PI. 12, figs. 376, 377 ; Text-figs. 251-257)
Callicilix abraxata Butler, 1885 : 124.

C. abraxata is known to have two subspecies. A single male from ' Masuri '
[N.W. India, Mussoorie] in the BM(NH) will probably prove to represent a third
subspecies.

Callicilix abraxata abraxata Butler

(PI. 12, fig. 376 ; Text-figs. 251-254)

Callicilix abraxata abraxata Butler ; Strand, 191 1 : 198.
Callicilix abraxata Butler ; Gaede, 1931 : 4. [Partim.]

Callicilix abraxata abraxata Butler ; Inoue, 1962 : 17, pi. 2, figs. 21, 22, figs. 45-60, (antenna,
venation, hind tibia, $ and $ genitalia).

This subspecies is distinguished from nguldoe by the colour-pattern (PI. 12, fig.
376, and colour-plate in Inoue (1962)) and in the male genitalia by the shape of the
eighth sternite and uncus.

Distribution. Japan. The localities have been listed by Inoue (1956) (in
English) and (1962) (in Japanese).

Type. I select as LECTOTYPE a $ in the collection of the BM(NH), labelled
' Callicilix abraxata Butler, type ; Yezo [Japan] ; Drepanidae genitalia slide No.
27 '. The locality of this specimen corresponds with that given by Butler in the
original description and it is doubtless one of the specimens, or the only specimen,
before Butler when he described this species.

Callicilix abraxata nguldoe (Oberthiir)

(PI. 12, fig. 377 ; Text-figs. 255-257)

Platypteryx nguldoe Oberthiir, 1893 : 22 - [Good fig.]
Callicilix abraxata nguldoe (Oberthiir) Strand, 191 1 : 198. [Poor fig.]
Callicilix abraxata var. nguldoe (Oberthiir) ; Gaede, 1931 14.
Callicilix abraxata formosana Okano, i960 : 11. syn. n.

This is distinguished from the nominate subspecies by the colour-pattern (PI. 12,
fig. 377, and colour-plate in Oberthiir (1893)), and in the male genitalia by the shape
of the eighth abdominal sternite and uncus. The female genitalia are illustrated in
Text-fig. 257.

Distribution. There are Chinese specimens from Oberthiir localities (Szechwan

CHINESE DREPANINAE

129

Figs. 251-257. Callicilix, genitalia. 251-254, abraxata abraxata, £. 251, eighth tergite
252, aedeagus ; 253, eighth sternite ; 254, $. 255-257, abraxata ngnldoe, 255, $ (part)
256, <$ eighth sternite ; 257, $.

i 3 o A. WATSON

and Tibet), and others from Kweichow and Hunan in the BM(NH). There are no
records of this subspecies from the eastern coastal provinces of China in spite of
the intensive collecting carried out there by Hone, although its presence in Formosa
suggests that it may yet be found on the adjacent mainland of China.

Material examined. Type of nguldoe. LECTOTYPE $, here selected, labelled :
de Ta-tsien-lou a Moupin, Mai, Juin 1892 (Chasseurs Thibetains). In the BM(NH).
This specimen was illustrated by Oberthur (1893).

Type of formosana. Holotype $, Central Formosa, Lushan, Nantowhsien,
ix.1959 ; in the Okano collection.

Other material. BM(NH). China, Tibet : i<J, 1 $, Tibet, ' frontiere orientale ',
1905 (Dejean). Szechwan : 6 J 1 , 1 $, Mt. Omei, vi-vii.1890, 1892, 28-31. vii.1921 ;
3 <J, Kwanhsien, 14.viii.1925, n.viii.1926 (Franck), viii.1930 (Franck) ; 1 <J, 2 $,
Tien-tsuen, 1897 (Dejean), 1899, 1903 (Dejean) ; 1 <$, 2 $, Siao-lou, 1900, 1908 ;
1 <$, 1 9, Siao-lou, Tchang-chau-pin, 1899 ; 1 <$, Moupin, 1898 ; 1 $, Ta-tsien-lou,
1910. Kweichow : 1^,1$ vi-vii.1890. Hunan : 3 <$ (Pratt). U.S. National
Museum. China, Szechwan : 1 <$, Mt. Omei (Graham) ; 1 $, Lingi Si, 3500 ft.
(Graham).

AUZAT A Walker

Auzata Walker, [1863] : 1620. Type-species Auzata semipavonaria Walker, [1863] : 1620, by

monotypy.
Auzata Walker ; Watson, 1959 : 232. [Revision.]
Gonocilix Warren, 1896 : 337. Type-species Gonocilix ocellata Warren, 1896 : 337 by monotypy.

[Synonymized by Gaede, 193 1 : 4.]

Distinguished from its nearest relative Callicilix Butler by the lamellate antennae,
and the presence of well-developed mid-tibial spurs on the hind leg.

Distribution. N. India, China, Korea, Japan, and S.E. Russia. (See Table 1.)
Auzata was revised by the present author in 1959 and no additional material has
been seen since then. The following list contains the species of this genus known
to occur in China, together with their distribution.
Auzata chinensis Leech, 1898 : 362. China.

A. chinensis Leech. China (Hunan, Szechwan).
A. chinensis prolixa Watson, 1959 : 238. China (Chekiang).
A. chinensis arcuata Watson, 1959 : 242. China (S. Shensi, Szechwan).
Auzata simpliciata Warren, 1897 : 13. N. India, China (N. Yunnan).
Auzata superba Butler, 1878 : 52. China, Japan, Korea, Quelpart Is., S.E.
Russia.

A. superba cristata Watson, 1959 : 248. China (Chekiang, Shansi, and prob-
ably Shensi).
Auzata minuta Leech, 1898 : 362. China.

A. minuta minuta Leech. China (Hupeh, Szechwan, Kweichow, Yunnan).
A. minuta spiculata Watson, 1959 : 253. China (Chekiang, and probably
Shansi).
Auzata ocellata (Warren), 1896 : 337. N. India, N. Burma, China, (Fukien).

CHINESE DREPANINAE 131

MACROCILIX Butler
(PI. 13, figs. 379-382 ; Text-figs. 258-270)

Macrocilix Butler, 1886 : 18. Type-species Argyris mysticata Walker, [1863] : 1617, by

monotypy.
Macrocilix Butler ; Gaede, 1931 : 5.

Macrocilix Butler ; Inoue, 1962 : 8. [Key to, and figs, of, Japanese species.]
Dipriodonta Warren, 1897 : 14. Type-species Dipriodonta sericea Warren, 1897 : 14, by

monotypy. [Synonymized with Macrocilix by Gaede, 1931 : 5.]
Sewa Swinhoe, 1900 : 591. Type-species Abraxas orbiferata Walker, 1862 : 1126, by monotypy.

[Synonymized with Macrocilix by Gaede, 1931 : 5.]

This genus comprises five species : mysticata Walker (see below), (India, Burma,
China, Formosa, Japan) ; maia Leech (see p. 133) (India, China, Japan, Malaya,
Sumatra) ; taiwana Wileman (see p. 135) (Formosa) ; orbiferata Walker (see p. 134)
(India, Sikkim, Burma, China, Malaya, Java, Borneo) ; and sericea Warren,
1897 : 14 (N. India). (See Table I.) The latter species is doubtless not congeneric
with the type-species of Macrocilix and will have to be transferred from Macrocilix
when its affinities can be more closely studied.

Macrocilix is probably most closely allied to Aitzala Walker from which it can be
distinguished by the yellow and grey, or yellow and black area at the anal angle of
the upper surface of the hind wing, and by the genitalia of both sexes.

Macrocilix mysticata (Walker)

(PL 13. figs. 381, 382 ; Text-figs. 258-264)

Argyris mysticata Walker, [1863] : 161 7.

Easily distinguished from the closely allied maia Leech by the colour-pattern and
the genitalia of both sexes. There are three subspecies, each of which is reviewed
below.

Distribution. N. India, Sikkim, N. Burma, China, Formosa and Japan. The
type of mysticata ab. flavotincta Wileman, 1915 : 19 will probably prove to represent
a new Formosan subspecies when further material becomes available ; the hind
wing pattern is similar to that of mysticata watsoni, whereas the fore wing pattern
most closely resembles that of the nominate subspecies.

Macrocilix mysticata mysticata (Walker)
(Text-figs. 261-264)

Macrocilix mysticata (Walker) ; Butler, 1886 : 19. [Good fig.]
Macrocilix mysticata (Walker) ; Strand, 1911 : 197. [Good fig.]
Macrocilix mysticata (Walker) ; Gaede, 1931 : 5.
Macrocilix mysticata mysticata (Walker) ; Inoue, 1958 : 12.

Macrocilix mysticata bidentata Bryk, 1943 : 5. [Good fig.] [Synonymized with mysticata
by Inoue, 1958 : 12.]

132

A. WATSON

Figs. 258-264. Macrocilix, genitalia. 258, 259, mysticata watsoni. 258, $ ; 259, <$ eighth
sternite. 260, mysticata brevinotata, $. 261-264, mysticata mysticata. 261, aedeagus ;
262, <J eighth sternite ; 263, <J ; 264, $.

CHINESE DREPANINAE 133

Distinguished from subspecies watsoni Inoue by the male and female genitalia, from brevinotata
ssp. n. by the female genitalia, and from both watsoni and brevinotata by the colour-pattern of
the wings. (See Text-figs.)

Type. Holotype $ [No locality data ; ' North Hindostan ' according to original
description] ; (Col. Buckley) ; Drepanidae genitalia slide No. 31. In the BM(NH).

Distribution. N. India, Sikkim ; N. Burma.

Macrocilix mysticata watsoni Inoue
(PI. 13, fig. 382 ; Text-figs. 258, 259)

Macrocilix mysticata watsoni Inoue, 1958 : II,

Macrocilix mysticata watsoni Inoue, 1962 : 10. [Good figs]

Separated from the nominate subspecies and brevinotata by the colour-pattern of the wings
and the female genitalia (Text-fig. 258), and from the nominate subspecies also by the eighth
sternum of the male genitalia (Text-fig. 259).

Distribution. Japan and China. There are specimens from the Chinese provinces
of Fukien, Chekiang, Kwangtung and Yunnan, in the Hone collection at Bonn, and
from Szechwan and Yunnan in the BM(NH).

Macrocilix mysticata brevinotata ssp. n.

(PI. 13, fig. 381 ; Text-fig. 260)

$. Distinguished from watsoni and the nominate subspecies by the female genitalia and the
colour-pattern. The abbreviated postmedial fascia on the hind wing is particularly diagnostic.
Wing. 190-230 mm. (5).
cj. Not known.

Distribution. This subspecies is probably restricted to the higher regions of
Szechwan. The subspecies watsoni also occurs in this province (1 $, Kwanhsien,
in BM(NH)), at a much lower altitude, but no intermediates between these two
subspecies have been captured.

Holotype $. China : Szechwan, Tu-pa-keo, 7400 ft., 4.ix.ig29 (Kelley-
Roosevelt Expedition) ; Drepanidae genitalia slide No. 249. In the BM(NH).

Paratypes. BM(NH). 5 $, type locality, 3-7. ix. 1929 (Kelley-Roosevelt Ex-
pedition).

Macrocilix maia (Leech)

Argyris maia Leech, 1888 : 647. [Good fig.]

Macrocilix maia (Leech) Leech, 1898 : 360.

Macrocilix maia (Leech) ; Gaede, 1931 : 5.

Macrocilix maia (Leech) ; Inoue, 1962 : 10. [Good figs. : including genitalia.]

Readily distinguished from mysticata Walker by the distinctive colour-pattern
and genitalia of both sexes (see Inoue, 1962).

134 A. WATSON

Distribution. India, China (i $ from Shantung, Tsingtau ; in the Museum
Koenig, Bonn), Korea, Japan, Malaya, Sumatra. A study of some of the available
material of this species has shown the presence of two new subspecies ; one in India,
and the other in Sumatra (and probably also Malaya) , The single female from China
mentioned above almost certainly belongs to the nominate subspecies, which occurs
in Japan and Korea.

Type. Holotype $, Korea, Gensan, vii.1887 {Leech) ; Drepanidae genitalia slide
No. 1612. In the BM(NH).

Macrocilix orbiferata (Walker)
(PI. 13, fig. 379 ; Text-figs. 265-270)
Abraxas orbiferata Walker, 1862 : 1126.

This species can be separated easily from maia Leech and mysticata Walker by
the colour-pattern of the wings, the lamellate antennae, and the genitalia of both
sexes. From taiwana Wileman (PI. 13, fig. 380), orbiferata is distinguished by the
colour-pattern and by the female genitalia (the male of taiwana is unknown).

Two subspecies are known : the nominate subspecies (Indo-Chinese and Malayan
Subregions), and cilicoides Snellen (1889 : 9, pi. 1, fig. 3) (Java) (see Text-figs.
269, 270).

Macrocilix orbiferata orbiferata (Walker)

(PI. 13, fig. 379)

Macrocilix orbiferata (Walker) ; Hampson, [1893] : 330.

Macrocilix orbiferata (Walker) ; Strand, 191 1 : 197.

Macrocilix orbiferata (Walker) ; Gaede, 1931 : 6.

Sewa orbiferata (Walker) Swinhoe, 1900 : 591.

Sewa orbiferata (Walker) ; Warren, 1922 : 447. [Good fig.]

Argyris insignata Moore, 1867 : 645. [Synonymy accepted from Hampson [1893] : 330.]

Distinguished from orbiferata cilicoides Snellen (1889) by the colour-pattern of the
fore wing. No reliable diagnostic features have been found in the female genitalia,
though little cilicoides material was available for study.

Distribution. N. India, N. Burma, China (Szechwan, Chekiang, Fukien), Malaya
and Borneo. Two females from the Chinese province of Hunan (in the BM(NH))
also probably belong to this subspecies.

Types, orbiferata. Holotype $, Borneo, Sarawak. In the Hope Department
Museum, Oxford.

insignata. The type material originated from ' Bengal ' according to the original
description. It formed part of the A. E. Russell collection which is apparently
lost (see Horn and Kahle, 1937 : 380).

CHINESE DREPANINAE 135

Macrocilix taiivana Wileman
(PI. 13, fig. 380)
Macrocilix taiwana Wileman, 1911 : 148.

Distinguished from its closest apparent relative, orbiferata, by the colour-pattern
and by the female genitalia (the male is unknown). The chief distinguishing
characters in the genitalia are the signum which extends for almost the whole length
of the corpus bursae, the presence of a deep invagination of the posterior margin of
the eighth abdominal sternite into which the ostium opens, and the ductus bursae
which is more heavily sclerotized than in orbiferata and is strongly constricted a
short distance before the ostium.

Distribution. Unknown outside Formosa.

Type. LECTOTYPE $, here selected, labelled : $, Arizan, Formosa, 7300 ft.,
22.viii.1908 (A. E. Wileman) ; 1225 [?] ; Wileman Coll., B.M. 1929-261 ; Macro-

Figs. 265-270. Macrocilix, genitalia. 265-268, orbiferata orbiferata. 265, $ ; 266, <£ ;
267, aedeagus ; 268, 3* eighth sternite. 269, 270, orbiferata cilicoides, $. 269, eighth
sternite ; 270, aedeagus.

136

A. WATSON

cilix taiwana sp., Type $ ; Drepanidae genitalia slide No. 1615. In the BM(NH).

Paralectotype. $, Formosa, Rantaizan, 7500 ft., n.v.1909 {A. E. Wileman).
In the BM(NH).

Other material. Formosa : 3 $, type-locality, 16.iii.-20.ix {Wileman).

HYALOSPECTRA Warren

(PI. 13, fig. 378)

Hyalospectra Warren, 1906 : 62. Type-species Hyalospectra grisea Warren, 1906 : 62, by
monotypy.

A comparison of hyalinata Moore, the only known Chinese species of Hyalospectra,

Figs. 271-274. Hyalospectra hyalinata, genitalia. 271, $ ; 272, aedeagus ; 273, $ eighth

sternite ; 274, <$.

CHINESE DREPANINAE 137

with the type-species from New Guinea has shown that these two species are certainly
not congeneric and that hyalinata, together with arizana Wileman (191 1 : 149), will
have to be removed from Hyalospectra when further revisionary work has shown
where the affinities of hyalinata and arizana lie.

Hyalospectra hyalinata (Moore)

(PI. 13, fig. 378 ; Text-figs. 271-274)

Comibaena hyalinata Moore, 1867 : 638.

Hyalospectra hyalinata (Moore) ; Warren, 1022 : 451. [PI. 48c (as hyalicosta).]

Hyalospectra hyalinata (Moore) ; Gaede, 1931 : 9.

Hyalospectra postfasciata Bryk, 1043 : 7. syn.n.

Distinguished from arizana Wileman (1911 : 149) by the broader subterminal
fasciae of the wings, and the presence of a brown medial shade on the fore wing.
No comparison of the genitalia could be made because of the badly damaged
abdomen of the holotype of arizana, the only specimen of this species. The genitalia
of hyalinata are figured in Text-figs. 271-274.

Distribution. N. India, Sikkini, N. Burma, China (N. Yunnan : ex. in BM(NH)
and Museum Koenig, Bonn). Specimens from Malaya, Thailand and Ceylon, in the
BM(NH), probably also represent this species.

Types, hyalinata. LECTOTYPE <$, here selected, labelled : Bengal ; N.E.
Bengal ; Russell ; Comibaena hyalinata type Moore [two labels] ; Moore Coll.
94-106. In the BM(NH).

postfasciata. Holotype $ [not <$ as stated by Bryk, 1943 : 7], N.E. Burma,
Kambaiti, 200 m., 9^.1934 {Malaise) ; in the Naturhistoriska Riksmuseet,
Stockholm.

CILIX Leach

(PI. 13, figs. 383, 386-388 ; Text-figs. 275-290)

Cilix Leach, [1815] : 135. Type-species Bombyx compressa Fabricius, 1777 : 279, by monotypy.

[B. compressa is a junior subjective synonym of Phalaena glaucata Scopoli, 1763 : 221.]
Cilix Leach ; Strand, 191 1 : 204.
Cilix Leach ; Gaede, 1931 : 37.

This is an easily recognizable genus of small white species distinguished from other
white Drepanidae by the irregularly shaped medial marking on the fore wing, and,
except for depalpata, by the anastomosis in the fore wing of 5c -f- R\ with the cell
near its base (see Inoue, 1962).

Seven species are known : glaucata Scopoli (1763 : 221) (Western Europe and
North Africa) ; asiatica Bang-Haas (1907 : 70), stat. n. (Middle-East) ; depalpata
Strand (191 1 : 204), sp. rev. (N.W. India, Afghanistan) ; filipjevi Kardakoff (Russia,
China, Korea, Japan) ; patula sp. n. (China) ; danieli sp. n. (China), and tatsienluica
Oberthiir (China). The four species represented in China are dealt with below.
(See Table 1.)

138

A. WATSON

Figs. 275-279. Cilix filipjevi, genitalia. 275-278, filipjevi filipjevi. 275, $ ; 276,
aedeagus ; 277, $ eighth sternite and posterior margin of eighth tergite ; 278, $ ; 279,
filipjevi malivora, $.

CHINESE DREl'AXIXAE 139

Cilix filipjevi Kardakoff
(PI. 13, figs. 386, 387 ; Text-figs. 275-279)
Cilix glancata filipjevi Kardakoff, 1928 : 417. [Fig.]

This species is readily separable from the west European glancata Scopoli (1763 :
221) by small differences in the colour-pattern (see plate) and by significant differ-
ences in the genitalia of both sexes. The species patnla sp. n. and danieli sp. n.,
which are characterized by a broader medial band on the fore wing and by the male
and female genitalia, are probably the closest relatives of filipjevi.

Two subspecies are known : the nominate subspecies (U.S.S.R., Korea and China)
and malivora Inoue (Japan).

Cilix filipjevi filipjevi Kardakoff
(PL 13, fig. 386 ; Text-figs. 275-278)

Cilix glaucata filipjt 11 Kardakoff ; Gaede, 1931 : 38.
Cilix filipjevi Kardakoff ; Inoue, 1958 : 13.

Distinguished from the Japanese subspecies malivora by the male and female
genitalia and by the shape of the medial marking on the fore wing.

Distribution. South-east U.S.S.R. (type-locality), Korea and north-east China
(several examples from Manchuria in the Naturhistorisches Museum, Vienna, two
examples in the BM(NH), and one in the Museum Koenig, Bonn).

Type. Holotype <$, S. Ussurigebiet, Russ Inseln [8 Km. S.W. of Vladivostock] ;
Zool. Mus. Berlin genitalia slide No. 101. In the Zoological Museum, Berlin.

Cilix filipjevi malivora Inoue

(PI. 13, fig. 387 ; Text-figs. 279)

Cilix filipjevi malivora Inoue, 1958 : 13. [Good figs, o* genitalia.]

Separable from the nominate subspecies by the male and female genitalia and the
shape of the medial marking on the fore wing.

Type. Holotype (J, [Japan, Honshu], Hida-niigawa, 3-4. viii. 1954 (S. Issiki).
In the Inoue Collection.

Cilix patnla sp. n.

(PI. 13, fig. 383 ; Text-figs. 280-283)

This species is probably most closely allied to danieli. It can be separated with certainty
from the latter by the male genitalia (particularly the elongate saccus, broad valves, spatulate
gnathus, and longer socii and valve processes). [See note on possible colour-pattern difference
under danieli.] It can be distinguished from filipjevi, which occurs in northern China
(Manchuria), by the broader medial marking of the fore wing and by the genitalia of both
sexes, particularly the anellus, socii and uncus in the male, and the length of the ductus bursae
and the shape of the ninth segment in the female.

Wing, <J 10-0-12-5 mm. (4) ; $ 11-0-12-5 mm - (5)-

140

A. WATSON

Type. Holotype^, China, N. Yunnan, Likiang, 30.viii.1935 {Hone) ; Drepanidae
genitalia slide No. 1017. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 62 <J, 5 $, N. Yunnan, Likiang,
15.iii-8.iv.1935, 24.V.1937 (Hone) ; 1 <?, N. Yunnan, A-tun-tse, c. 3000 m. /18.V.1937

Figs. 280-283. Cilix patula, genitalia. 280,^; 281, aedeagus ; 282, <$ eighth tergite and

sternite ; 283, $.

CHINESE DREPANINAE 141

(Hone). BM(NH). China : 5 $, 1 $, N. Yunnan, Likiang, i.vii.1934, 23.iii-
3JX.1935 (Hone), Daniel Collection, Munich. China : 2 $, N. Yunnan, Likiang,
i.vii, 8.ix.i935 (Hone).

Cilix danieli sp. n.

(Text-figs. 284-286)

Separable from its close ally pahtla by the male genitalia (see patula), and by the postmedial
fascia on the fore wing which in the few specimens available for study is much less strongly
marked than in patula.

Wing. £ 11 -0-12-0 mm. (3) ; $ 11-5 (1).

Holotype <$. China : Shansi, Mien-Shan, c. 2000 m., i.viii.1937 (Hone) ;
Drepanidae genitalia slide No. 1671. In the Museum Koenig, Bonn.

Paratypes. Museum Koenig, Bonn. China : 1 <J, 1 ?, Shansi, Mien-shan,
c. 2000 m. 3-9. viii. 1937 (Hone). BM(NH). China : 1 <£, Shansi, Mien-shan
c. 2000 m., 31.vii.1937 (Hone). Daniel Collection, Munich. China : 1 $ Shansi,
Mien-shan, c. 2000 m., 14.vii.1937 (Hone).

Figs. 284-286. Cilix danieli, <$ genitalia. 284, ^ ; 285, aedeagus ; 286, eighth tergite

and sternite.

i 4 2 A. WATSON

Cilix tatsienluica Oberthur
(Pl. 13, fig. 388 ; Text-figs. 287-290)

Cilix spinula tatsienluica Oberthur, 1916 : 371.

Cilix tatsienluica Oberthur ; Oberthur, 1917 : 41. [Good fig.]

Cilix glaucata var. tatsienluica Oberthur ; Gaede, 1931 : 38.

This species is readily distinguished from the rest of the genus by the absence on
the fore wing of a terminal fascia, the poorly marked postmedial fascia and, except
at the apex, by the white fringe. The male and female genitalia are also diagnostic
(in particular, the presence of a single signum in the female and the structure of the
aedeagus, anellus, gnathus, valves and uncus in the male).

Distribution. China (Szechwan, Hupeh, Yunnan, Shansi, Shensi).

Material examined. Type. I select as LECTOTYPE a $ syntype in the BM(NH),
labelled : Ta-tsien-Lou, Chasseurs indigenes, 1893 ; Drepanidae genitalia slide No.
1021.

Other material. BM(NH). China : 1 <J, [Szechwan), Ta-tsien-Lou, v-vi.1866 ;
1 <$, [Szechwan), Tsekou, 1898 (Dubernard) ; 1 $, [Hupeh], Chang Yang, vi.1888
(Pratt). Museum Koenig, Bonn. China : numerous examples from Shansi, Mien-
shan, and S. Shensi, Tapaishan ; 4 ex., N. Yunnan. Daniel Collection, Munich.
China : 3 ex., Shansi, Mien-Shan ; c. 2000 m. (Hone) 3 ex., S. Shensi, Tapaishan im
Tsinling, c. 1700 m. (Hone) ; 2 ex., N. Yunnan, A-tun-tse, c. 3000 m. (Hone).

MACRAUZATA Butler

(PI. 13, fig. 385)

Macrauzata Butler, 1889 : 43. Type-species, Comibaena fenestraria Moore, 1867 : 639, by

monotypy.

Macrauzata Butler ; Gaede, 1931 : 4.

Macrauzata Butler ; Inoue, i960 : 314. [Good figs.]

This genus has been recently reviewed by Inoue (i960 : 314) who added a new
polytypic species maxima [Japan and China] to the genus. The colour-pattern (in
particular the large hyaline patches in both wings) and the genitalia distinguish
Macrauzata from all other Drepaninae. Its affinities are doubtful.

Distribution. Three species are known : fenestraria Moore (1867 : 639) [India,
Sikkim], minor Okano (1959 : 37) [Formosa] and maxima. M. maxima Inoue
(i960 : 314) comprises two subspecies ; the nominate subspecies [Japan] and
maxima chinensis Inoue (i960 : 314) [China] (see PI. 13, fig. 385). In addition to the
type from Szechwan and the paratype from Chekiang listed by Inoue (i960) for
chinensis, the provinces of Yunnan, Hunan, Fukien and Kiangsu are represented by
specimens in the Museum Koenig, Bonn. A specimen labelled ' Mongolia Whitely '
in the BM(NH) may also prove to represent this subspecies. (See Table 1.)

The genus Macrauzata occurs also in the Philippines, Celebes, Sumatra and Java
but the specific identity of the available material, in the BM(NH), has yet to be
investigated.

CHINESE DREl'ANINAE 143

PHALACRA Walker
(PI. 13, fig. 384 ; Text-figs. 291-293)

Phalacra Walker, 1866 : 1638. Type-species Phalacra metagonaria Walker, 1866 : 1639, by
monotypy. Holotype <$, ' Hindostan ' [India] ; in the BM(NH). [P. metagonaria Walker
is a junior subjective synonym of Hemerophila vidfiisara Walker, i860 : 319.]

Phalacra Walker ; Gaede, 193 1 : 13.

287

Figs. 287-290. Cilix tatsienluica, genitalia. 287, $ ; 288, <J ; 289, <£ eighth tergite and

sternite ; 290, aedeagus.

i 4 4

A. WATSON

The validity of the classification of the species at present included in Phalacra
needs to be investigated. An examination of the type-species of Phalacropsis
Swinhoe (1895 : 5) and Pseuderosia Snellen (1889 : 15) has shown that these genera
have close affinities with Phalacra.

The species of Phalacra are Indian, Chinese, Malaysian or Indonesian in distribu-
tion. Two species occur in China : strigata Warren (see below), and an undescribed
species from Kwantung represented by a single male in the Museum Koenig, Bonn.

Phalacra strigata Warren

(PI. 13, fig. 384 ; Text-figs. 291-293)

Phalacra strigata Warren, 1896 : 338.
Phalacra strigata Warren ; Warren, 1922 : 458.
Phalacra strigata Warren ; Gaede, 1931 : 14.
Phalacra multilineata Warren, 1897 : 16. syn. n.
Phalacra multilineata Warren ; Warren, 1922 : 458.
of strigata.]

[Fig. 48h closely matches the holotype

The Chinese examples in the Hone collection at Bonn from Lingping (Kwangsi
Province) and Canton (Kwangtung Province), and in the BM(NH) from localities

Figs. 291-293. Phalacra strigata, $ genitalia. 291, o* ; 292, posterior margin of eighth

sternite ; 293, aedeagus.

CHINESE DREPANINAE 145

in Szechwan probably represent a new subspecies of strigata, but there is insufficient
material available from the type-locality to allow a satisfactory comparison to be
made.

Distribution. N. India and China.

Types, strigata. Holotype £, Cherrapunji, xii.1893. In the BM(NH).

multilineata. LECTOTYPE $ here selected, labelled : Khasis, July, 1896, Nat.
Coll. ; Phalacra multilineata Type $ Warren ; Rothschild Bequest B.M. 1939-1 ;
Drepanidae genitalia slide No. 642. In the BM(NH).

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Drepanidae : Drepaninae). Trans. R. ent. Soc. Lond. 119 : 303-362, 4 pis.
Yamamoto, Y. i960. A new species of the Drepanidae from the Island of Yakushima, South
Japan. Tinea, Tokyo 5 : 334-335, 3 text-figs.

INDEX

abraxata Butler, 128
acuminata Leech, 49
acuta Butler, 104
agna Oberthiir, 75
Agnidra Moore, 29
Albara Walker, 16
" Albaria "; Walker 16,
albidescens Hampson, 124
albinotata Moore, 1 1 1
Allodrepana Roepke, 69
amabilis Bryk, 73
arcuata Walker, 8
arcuata Watson, 130
argenticeps Warren, 77
argentilinea Walker, 113
argyrobapta Gaede, 114
arikana Matsumura, 113
asiatica Bang-Haas, 137
Ausaris Walker, 113
Auzata Walker, 130

Betalbara Matsumura, 46
bicostata Hampson, 74
bidens Leech, 92
bidentata Bryk, 131
bifida Watson, 127
bilineata Packard, 8
binaria Hufnagel, 92
birmana Bryk, 102
birmanica Bryk, 40
bitorosa ssp. n., 97
bouvieri Oberthiir, 103
brevinotata ssp. n., 133

Callicilix Butler, 127
Callidrepana Felder, 113

Campylopteryx Warreri, 99
Canucha Walker, 99
carinata Watson, 127
cervinaria Moore, 27
chinensis Inoue, 142
chinensis Leech, 130
cilicoides Snellen, 134
Cilix Leech, 137
circumscripta Warren, 124
compressa Fabricius, 137
corticata Warren, 39
crassimaculata Inoue, 116
cretacea Hampson, 107
cristata Watson, 130
crocea Leech, 113
cultraria Fabricius, 92
cupreogrisea Hampson, 58
curta ssp. n., 121
curvaria Walker, 99
curvatula Borkhausen, 104
Cyclura Warren, 7

Damna Walker, 113
danieli sp. n., 141
depalpata Strand, 137
" depressa " Warren, 99
Deroca Walker, 127
deumbrata Warren, 124
Didymana Bryk, 92
diluta Warren, 97
Dipriodonta Warren, 131
" discipilaria "; Moore, 44
discispilaria Moore, 44
dispilata Warren, 109
Ditrigona Moore, 4
divisa Bryk, 27

150

A. WATSON

Drapetodes Guenee, 124
Drepana Schrank, 103
Drepania Hiibner, 103
Drepanula Gaede, 114
Drepanulides Motschulsky, 114
Drepanulina Gaede, 114
duplexa Moore, 100
duplicata Warren, 88

emarginata ssp. n., 94
erpina Swinhoe, 73

Falcaria Haworth, 104
falcataria L., 103
fenestra Leech, 34
fenestraria Moore, 142
filipjevi Kardakoff, 139
finita Watson, 113
fixseni Bryk, 42
flavilinea Leech, 55
flavotincta Wileman, 131
flexuosa ssp. n., 107
forcipulata ssp. n., 124
formosana Inoue, 116
formosana Okano, 128
formosicola Matsumura, 103
francki ssp. n., 40
fulva Hampson, 113
fulvata Snellen, 113
fulvior sp. n., 31
furva sp. n., 33
fuscifascia sp. n., 67
fuscilinea Watson, 44

gelidata Walker, 113
gemina sp. n., 119
glaucata Scopoli, 137
Gonocilix Warren, 130
gracillima Warren, 73
grisea Warren, 136
grisearia Leech, in
grisearia Staudinger, 84
grisearipennis Strand, in
griseola Matsumura, 67
griseotincta Wileman, 17
grotesca Bryk, 27

harpagula Esper, 94
hidda Swinhoe, 127
hirayamai Nagano, 121
hoenei sp. n., 31
horishana Matsumura, 17
humerata Warren, 90
hyalina Walker, 127

hyalinata Moore, 137
Hyalospectra Warren, 136
Hybodrepana Bryk, 24

ida Bryk, 49

inaequiscata Warren, 20

inconclusa Walker, 127

insignata Moore, 134

" Iridrepana " ; Warren, in

japonibia Strand, 104
japonica Moore, 80

koreula Bryk, 105

lacertinaria L., 104

lacertula Denis and Schiffermuller, 104

latizona Watson, 127

Leucodrepana Hampson, 4

leucosticta Hampson, 55

lilacina Moore, 87

Macrauzata Butler, 142
Macrocilix Butler, 131
macularis Bryk, 46
maculosa Watson, 113
" magnadiscata "; Warren, 46
magnidiscata Warren, 44
maia Leech, 133
malivora Inoue, 139
manleyi Leech, 52
marginata Watson, 113
maxima Inoue, 142
metagonaria Walker, 143
Microblepsis Warren, 46
mimetica Warren, 73
minor Okano, 142
minuta Leech, 130
miranda Warren, 103
mitaria Guenee, 124
Monurodes Warren, 97
multilineata Warren, 144
muscula Staudinger, 67
muscularia Walker, 20
mysticata Walker, 131

nguldoe Oberthiir, 128
nigrimaculata Warren, 97
nigritincta Warren, 27
nigromaculata Okano, 109
Nordstroemia Bryk, 69

obtruncata Warren, 99
ocellata Oberthiir, 73
ocellata Warren, 130

INDEX

151

ochracina Bryk, 36
ochrozona Bryk, 88
ogasawarae Matsumura, 49
olivacea Inoue, 94
olivacea Warren, 88
opalescens Oberthiir, 75
opalescens Warren, 17
orbiferata Walker, 134
Oreta Walker, 7
orphnina Hampson, 20
ovata sp. n., 117

Palaeodrepana Inoue, 92

palleolus Motschulsky, 116

pallida Moore, 105

pallidina Bryk, 73

pallidinota sp. n., 24

Paralbara gen. n., 19

parvula Leech, 67

patrana Moore, 114

patula sp. n., 139

perhamata Hampson, 22

Peridrepana Butler, 4

Phalacra Walker, 143

Phalacropsis Swinhoe, 144

" Platypterix "; Ochsenheimer, 104

Platypteryx Laspeyres, 103

postfasciata Bryk, 137

Prionia Hiibner, 104

problematica Bryk, 77

prolatior ssp. n, 53

prolixa Watson, 130

prunicolor Moore, 53

Pseudalbara Inoue, 65

Pseuderosia Snellen, 144

pulchripes Butler, 8

pulla Watson, 127

pulvis Oberthiir, 126

puctifera Strand, 80

recava sp. n., 84
rectilinea sp. n., 61
renei Bryk, 92
reversaria Walker, 17
robusta Oberthiir, 65
rosea Walker, 8
rubromarginata Leech, 113
rufa Warren, 27
rufata ssp. n., 111
rufofasciata Hampson, 105
rugosa sp. n., 62

sachalinensis Matsumura, 80
saucia Felder, 113
scabiosa Butler, 42

scintillata Walker, 113
semipavonaria Walker, 130
sericea Warren, 131
Sewa Swinhoe, 131
shensiensis ssp. n., 58
siccifolia Roepke, 87
sicula Esper, 104
simillima Moore, 87
simpliciata Warren, 130
simplificaria Strand, 117
specularia Walker, 36
specularis Moore, 99
spicula sp. n., 22
spiculata Watson, 13
Strepsigonia Warren, 97
strigata Warren, 144
subbasalis Inoue, 116
sublignata Warren, 99
subpallida Warren, 20
sumatrana Roepke, 80
superba Butler, 130

taiwana Wileman, 135
takasago Okano, 62
tatsienluica Oberthiir, 142
Teldenia Moore, 4
thermopasta Hampson, 113
Thymistida Walker, 24
Thymistadopsis Warren, 124
Ticilia Walker, 113
Tridrepana Swinhoe, 1 1 1
trigonoptera Warren, 97
trilinearia Moore, 125
tripunctata Walker, 27
trisulcata Hampson, 126

undata sp. n., 90
undilineata Warren, 24
undulifera Hampson, 126
unispina Watson, 113
urupula Bryk, 105
usta Butler, 44

vidhisara Walker, 143
vinacea Moore, 40
violacea Butler, 62
vira Moore, 73

warreni Bryk, 53
watsoni Inoue, 133

x-z-nigrum Bryk, 109

yakushimalis Yamamato, 121

Zanclalbara Inoue, 29

PLATE i

A Ibara

Figs. 296, 297, reversaria opalescens (xif). Fig. 296, <$ (holotype of griseotincta) ; Fig.
297 $.

Paralbara ( x 2)

Fig. 294, muscularia, neotype <$. Fig. 295, pallidinota, holotype o*- Fig. 298, spicula,
holotype $. Fig. 299, perhamata, q*.

Thymistida
Fig. 300, nigritincta, <J ( X2).

Agnidra ( x 2)

Fig. 301 scabiosa scabiosa, o*- Fig. 302, scabiosa fixseni, o*- Fig. 303, corticata francki, $.
Fig. 304, corticata corticata, o*. Fig. 305, vinacea, $.

Hull. Br. Mus. a, it. Hist. (Ent.) Suppl. 12

PLATE i

294

295

296

297

298

299

m

300

301

302

303

304

305

PLATE 2

Agnidra

Fig. 306, specularia, <$ (X2). Fio. 307, fenestra, $ (X3). Fig. 308, discispilaria, <$ (xi£).
Fig. 309, hoenei, holotype $ (X2). Fig. 310, fulvior, holotype £ (X2). Fig. 311, furva,
holotype <$ ( X2).

Betalbara
Fig. 312, prunicolor, $ (X3). Fig. 313, flavilinea flavilinea, $ (X2). Fig. 314, leucosticta
(?(X 3 ).

Hull. lir. Mus. not. Hist. (Knt.) Suppl. [2

PLATE 2

306

307

308

Wi t ' *

309

310

312

313

314

PLATE 3
Betalbara
Fig. 315, manleyi prolatiov, holotype $ (X2). Fig. 316, cupreogrisea, $ (X2). Fig. 317,
rugosa, holotype 5" ( X2). Fig. 318, violacea, $ ( x if). Fig. 319, robusta, 6* ( x 2).

Pseudalbara
Fig. 320, parvula, <J ( X3J). Fig. 321, fusci fascia, $ ( X3).

Nordstroemia
Fig. 322, vira, $ ( X3). Fig. 323, bicostata bicostata, holotype o* ( X3).

Hull. lir. Mus. nat. Hist. (Ent.) Suppl. 12

PLATE 3

\

315

316

317

318

319

320

321

322

323

PLATE 4
Nordstroemia
Fig. 324, duplicata, lectotype rj (X2). Fig. 325, samatrana, holotype ? ( x 1, approx.).
Fig. 326, siccifolia, <$ ( X2). Fig. 327, recava, holotype <$ ( X2). Fig. 328, humerata, $ ( X3).
Fig. 329, undata, <$ paratype ( x 2).

Palaeodrepana ( x 2)

Fig. 330, harpagula bitorosa, holotype J. Fig. 331, harpagula harpagula, $. Fig. 332,
harpagula emarginata, holotype $.

Bull. Br. Mus. not. Hist (Km.) Suppl. n

PLATE 4

324

325

326

^T

^>

H

327

f

328

329

330

33

332

PLATE 5

Nordstroemia

Figs. 333~335. vira, 6* genitalia, Drepanidae genitalia slide No. 752. Fig. 333, genitalia.
FlG - 334- aedeagus. Fig. 335, 8th abdominal segment with tergite, sternite and lateral sacs.
Fig. 336, argenticeps , holotype <$ genitalia, Drepanidae genitalia slide No. 777.

Bull. Br. Mus. nat. Hist. (Ent.) Suppl. 12

PLATE 5

333

PLATE 6

Nordstroemia

Figs. 337, 338, argenticeps, holotype <J genitalia, Drepanidae genitalia slide No. 777. Fig.
337, aedeagus. Fig. 338, posterior end of abdomen showing 7th sternite, 8th tergite and 8th
sternite. Figs. 339, 340, sumatrana, holotype o* genitalia, Drepanidae genitalia slide No. 1665.
Fig. 339, genitalia. Fig. 340, posterior end of abdomen showing 7th sternite, 8th tergite and
8th sternite.

Hull. Br. Mus. nut. Hist. (Eat.) Suppl. [2

PLATE

PLATE 7
Nordstroemia

Figs. 341, 342, lilacina, lectotype $ genitalia, Drepanidae genitalia slide No. 780. Fig. 341,
aedeagus. Fig. 342, genitalia. Fig. 343, simillima, lectotype <J, Drepanidae genitalia slide
No. 779, posterior end of abdomen showing 7th sternite, 8th tergite, and 8th sternite. Fig.
344, sumatrana, $, Drepanidae genitalia slide No. 1665, aedeagus.

Bull. By. Mus. nat. Hist. (Ent.) Suppl. 12

PLATE 7

PLATE 8

Nordstroemia
Figs. 345, 346, siccifolia, q* genitalia, Drepanidae genitalia slide No. 1666. Fig. 345, aedeagus.
Fig. 346, genitalia. Figs. 347, 348, simillima, lectotype <$ genitalia, Drepanidae genitalia
slide No. 779. Fig. 347, genitalia. Fig. 348, aedeagus.

Bull. Br. Mus. nat. Hist. (Ent.) Suppl. i:

PLATE 8

PLATE 9
Nordstroemia
Fig. 349, ochrozona, posterior end of abdomen, showing 7th sternite, 8th tergite and 8th
sternite, Drepanidae genitalia slide No. 938. Fig. 350, humerata, posterior end of abdomen,
showing 7th sternite, 8th tergite and 8th sternite, Drepanidae genitalia slide No. 1670. Fig.
351, lilacina, posterior end of abdomen, showing 7th sternite, 8th tergite and 8th sternite,
Drepanidae genitalia slide No. 780. Fig. 352, siccifolia, posterior end of abdomen, showing
7th sternite, 8th tergite, and 8th sternite, Drepanidae genitalia slide No. 1666.

Bull. Br. Mus. nat. Hist. (Ent.) Suppl. 12

PLATE 9

PLATE 10

Nordstroemia

Figs. 353, 354, ochrozona, <$ genitalia, Drepanidae gentialia slide No. 938. Fig. 353, genitalia.
F IG - 354. aedeagus. Figs, 355, 356, humerata $ genitalia, Drepanidae genitalia slide No. 1670.
Fig. 355, genitalia. Fig. 356, aedeagus.

Bull, lit Mus. not. Hist (Ent.) Suppl. n

PLATE 10

t

4

355

PLATE u
Didymana
Fig. 357, bidens, <$ ( x 2) .

Drepana ( x 2)
Fig. 358, curvatula curvatula, $. Fig. 360, pallida cretacea, <J. Fig. 361, pallida flexuosa,
holotype <J. Fig. 362, dispilata rufata, holotype $. Fig. 363, curvatula acuta, $. Fig. 364,'
dispilata grisearipennis Strand, holotype $. Fig. 365, ritfofasciata, lectotype $.

Strepsigonia
Fig. 359, diluta, $ (X3).

Canucha ( x 1)
Fig. 366, bouvieri, holotype $. Fig. 367, specularis, $. Fig. 368, duplexa duplexa, <J.

Hull. Br. Mus. not. Hist. (Ent.) Suppl. i.

PLATE ii

357

/

360

362

\7 ^* *^y

PLATE 12
Callidrepana ( X 2)

Fig. 369, ovata, holotype (J. Fig. 370, gemina curta, holotype $. Fig. 371, gemina gemina,
holotype <$. Fig. 372, hirayamai forcipulata, holotype 5.

Drapetodcs
Fig. 373, mitaria, $ (X3).

Thymistadopsis
Fig. 374, trilinearia, $ ( X2).

Deroca
Fig. 375, hyalina latizona, $ (X2).

Callicillix ( x 2)
Fig. 376, abraxata abraxata, J. Fig. 377, abraxata nguldoe, <$.

Bull. Br. Mus. nat. Hist. (Ent.) Suppl. n

PLATE 12

369

370

371

T

372

373

374

f ' '■

375

PLATE 13
Hyalospectra
Fig. 378, hyalinata, lectotype $ of postfasciata (synonym) ( x 2).

Macrocilix ( X 2)

Fig. 379, ovbiferata orbiferata, $. Fig. 380, taiwana, $. Fig. 381, mysticata brevinotata,
holotype $. Fig. 382, mysticata watsoni, paratype <J.

Cilix ( x 3£)

Fig. 383, patula, $. Fig. 386, filipjevi filipjevi, $. Fig. 38-/, filipjevi malivora, £. Fig. 388,

tatsienluica, $.

Phalacra

Fig. 384, strigata, <$ ( X2).

Macrauzata

Fig. 385, maxima chinensis, g ( X 2).

Bull. Br. Mus. nat. Hist. (Ent.) Suppl. i.

PLATE 13

PLATE 14

Thymistida

Figs. 389-392, tripunctata genitalia, Drepanidae genitalia slides Nos. i22(cJ) and 123(9)
Fig. 389, $ ; Fig. 390, aedeagus ; Fig. 391, posterior end of abdomen showing modified seventh
and eighth sternites and eighth tergite ; Fig. 392, $.

Bull. Br. Mus. nut. Hist. (Ent.) Suppl. 12

PLATE 14

'

\ i

1

iT — i "-

/' __.

391

A LIST OF SUPPLEMENTS
TO THE ENTOMOLOGICAL SERIES
OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

i. 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. Sep-
tember, 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.

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the
world species of Cleora (Lepidoptera : Geometridae). Pp. 119 : 14 plates, 146
text -figures, 9 maps. February, 1967. £3 10s.

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species
(Lepidoptera : Rhopalocera) . Pp. 509. £& 10s.

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera : Rho-
palocera). Pp. 322 : 348 text-figures. August, 1967. £8.

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172 :
82 text-figures. May, 1968. £4.

■$&£§? Printed in England by Staples Printers Limited at their Kettering, Northants, establishment

I l0 jan m

MORPHOLOGY AND TAXONOMY OF

THE ADULT MALES OF THE FAMILIES

PSEUDOCOCCIDAE AND

ERIOCOCCIDAE

(HOMOPTERA: COCCOIDEA)

S. A. AFIFI

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Supplement 13

LONDON: 1968

MORPHOLOGY AND TAXONOMY OF

THE ADULT MALES OF THE FAMILIES

PSEUDOCOCCIDAE AND ERIOCOCCIDAE

(HOMOPTERA: COCCOIDEA)

BY

S. A. AFIFI

Dept. of Entomology, Ministry of Agriculture, Egypt (U.A.R.)

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Supplement 13

LONDON: 1968

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. 13 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) 1968

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 31 December, 1968 Price £5

MORPHOLOGY AND TAXONOMY OF THE

ADULT MALES OF THE FAMILIES

PSEUDOCOCCIDAE AND ERIOCOCCIDAE

(HOMOPTERA : COCCOIDEA)

By S. A. AFIFI*

* The information here provided was submitted in the form of a thesis for the degree of Doctor of
Philosophy in the University of London, January, 1967.

10 JAN196<

CONTENTS

Acknowledgments
Introduction
Review of literature
Material, technique and illustrations
General morphology
pseudococcidae

General Characteristics

MACROPTEROUS MALES

Head ....

Head Capsule

Antennae .

Thorax

Prothorax .

Mesothorax

Metathorax

Wings and Hamulohalterae

Legs ....

Abdomen .

Pregenital Segments

Genital Segment and External Genitalia
Apterous males

Head .

Thorax

Abdomen .
Eriococcidae .

General Characteristics
Macropterous males

Head ....

Head Capsule

Antennae .

Thorax

Prothorax .

Mesothorax

Metathorax

Wings and Hamulohalterae

Legs ....

Abdomen .

Pregenital Segments

Genital Segment and External Genitalia

Page

4

5

6

6

10

11

11

12

12

12

15

!7
*7

18

21
22
22
23
23
24
25
25
26
26
26
26
27
27
27
27
28
28
28
28
29

29
29

29
29

4

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Brachypterous males
Head .
Thorax
Abdomen .
Apterous males
Discussion

Status of Pseudococcidae and Eriococcidae
Taxonomic Significance of the Characters
Classification of Family Pseudococcidae
Groups of Genera
Genera .
Species .
Family Eriococcidae
Relationships of Pseudococcidae and Eriococcidae with other Coccoidea
Analysis .....
Description of species
Pseudococcidae
planococcus Group
pseudococcus Group
saccharicoccus group .
octococcus Group,
ceroputo Group

NAIROBIA Group

Eriococcidae .
Keys .....

Lettering used in text-figures
List of abbreviations
References ....
Index .....

30
30
30
30
30
31
3i
54
58
63
69
72
74
74
79
92

93
93
131
143
151
155
163
167
196
203
204
206
210

SYNOPSIS

The adult males of 17 pseudococcid species (belonging to 13 genera, and represented by
macropterous and apterous forms) and 7 eriococcid species (belonging to 4 genera, and repre-
sented by macropterous, brachypterous and apterous forms) were described and illustrated
in detail. The study indicated that the two groups, whose taxonomic status was yet contro-
versial, deserved the rank of separate families, and the characters differentiating the males of the
two families were given. The general morphology of the males was discussed and a number
of new terms introduced. The affinities of the studied species (and 3 others described in detail
by Giliomee, 1961) were determined and statistically analysed, and a classification of Pseudo-
coccidae was suggested. The relationships of the lecanoid types of male (Pseudococcidae,
Eriococcidae and Coccidae), between each other and with other families of Coccoidea were
discussed. The results of this work confirmed earlier conclusions by Ghauri (1962) and
Giliomee (1961 & 1967) that the males afford significant characters at all the taxonomic levels
including the specific. Detailed keys to the families downwards to the species were constructed.

ACKNOWLEDGMENTS

I am greatly indebted to Dr. K. L. Boratynski, who suggested and supervised this
study, and also placed some material from his own collection at my disposal. I am
also grateful to Prof. O. W. Richards, F.R.S., Head of the Department of Zoology
and Applied Entomology, Imperial College, for the facilities granted, and to Mr. R. G.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 5

Davies for reading part of the manuscript, and highly valued criticism. The
technical remarks received from Dr. J. H. Giliomee, the earlier Ph.D. student in the
department, are thankfully acknowledged. My deep appreciation is also due to
the Trustees of the British Museum (Nat. Hist.), and also to Dr. D. J. Williams,
of the Commonwealth Institute of Entomology, for providing specimens of four
species ; a large number of other workers have also kindly furnished material from
their collections, or even took the trouble to collect and breed the males specially
for the purpose of this study, namely : the late Prof. N. S. Borchsenius, of the
Zoological Institute, Academy of Sciences, Leningrad ; Dr. G. M. Das, of the
Tocklai Experiment Station, Cinnamara, Assam, India ; Mr. G. De Lotto, of
the Research Institute for Plant Protection, Pretoria ; Mgr. A. Dziedzicka, of the
Pedagogical Higher School, Krakow, Poland ; Mr. A. I. Ezz, of the Department of
Entomology, Ministry of Agriculture, Cairo, U.A.R. ; Mr. J. M. Hoy, of the
Entomology Division, Dept. of Scientific and Industrial Research, New Zealand ;
Prof. M. Kosztarab, Virginia Polytechnic Institute, Blacksburg, Virginia ; Dr. G.
Matesova, of the Institute of Zoology, Academy of Sciences, Alma-Ata, Kazakhstan ;
Mr. D. N. McNutt, of the Agricultural Department, Mbale, Uganda, East Africa ;
Ing. N. Mitic-Muzina, of the Institute for Plant Protection, Belgrade ; Mr. J.
Munting, of the Plant Protection Research Institute, Pretoria.

In this opportunity, I would like also to express my gratitude to Prof. Dr. Y. M.
Ezzat, Head of the Department of Zoology, University of Assiout, U.A.R., for his
constant and unfailing encouragement, which has always given me a great deal
of confidence.

The Ministry of Agriculture of the United Arab Republic (Egypt) has the credit
of originally awarding me a study leave, which was later elevated to a full mission ;
a financial support without which this work would have not been accomplished.

INTRODUCTION

This work is the fourth in a series of detailed morphological and taxonomical
studies on the males of Coccoidea (Theron, 1958, studied the general morphology of
7 species representing 4 families ; Ghauri, 1962, described in detail 24 species of the
family Diaspididae ; Giliomee, 1967, similarly treated 23 species of the family
Coccidae) carried out by the postgraduate students in the Department of Zoology
and Applied Entomology, Imperial College, suggested and supervised by Dr. K.
Boratynski. The present publication deals with 24 species belonging to two groups,
the pseudococcids (17 species) and the eriococcids (7 species), whose taxonomic
status is still controversial ; they have been considered as (1) merely tribes of one
subfamily, Pseudococcidae (Cockerell, 1899 ; Balachowsky, 1948) ; (2) subfamilies
of one family (Balachowsky, 1942 ; Borchsenius, 1949), or (3) separate families,
with rather remote connection between them (Ferris, 1937, 1957 ; Borchsenius,
1963). Apart from the grouping of certain pseudococcid genera by some authors
(the Rhizoecus group including 8 genera by Hambleton, 1946, and the tribe
Planococcini by Ezzat & McConnel, 1956), no comprehensive classification of the
pseudococcids or the eriococcids has been hitherto proposed. The purpose of the

6 MORPHOLOGY AND TAXONOMY OF ADULT MALES

present study therefore was (i) to carry out a detailed morphological study of
the available male representative of the two groups ; (2) to evaluate the various
characters for classificatory purposes ; (3) to determine the status and relationships
of the two groups with each other and with other families of Coccoidea ; and (4)
to suggest a classification for each of these groups.

REVIEW OF THE LITERATURE

The literature on the males of Coccoidea has been well covered by Ghauri (1962)
and Giliomee (1967) ; therefore only the papers either omitted by these authors,
or published subsequent to their works, will be discussed here.

One of the early papers not mentioned by them is Comstock's (1881) First Report
on Scale Insects, in which the main features of the males of a number of species of
the subfamilies Diaspinae (=Diaspididae) and one eriococcid, Rhizococcus
araucariae (Mask.) are briefly discussed. In many other early papers, however, the
reference to the males are restricted to brief accounts mainly on the shape, size and
colour of their bodies, or on the available puparia, apparently the empty ones
(e.g. Rutherford, 1914 ; Brain, 1920 ; Hall, 1928 ; Takahashi, 193 1). A paper
by Kuwana (1923) (mentioned by neither Ghauri nor Giliomee) includes descrip-
tions and illustrations of four scale insects, one mealybug, and one margaroid.
MacDougall (1926) provided generalized descriptions of the males of Pseudococcus
comstocki Kuwana. Rao (1943) drew attention to the existence of the apterous
males of Trionymus sacchari Ckll. but made no description of any morphological
importance. Beardsley published several papers dealing with a large number of
pseudococcid males occurring in Hawaii (i960, 62, 63, 64 and 65) ; his illustrations
of almost all the species only included ventral aspects of the penial sheath. In his
paper of 1964 he suggested close relationship between Phenacoleachia and Pseudo-
coccidae, which was also recognized by Giliomee (1967). In 1965, Beardsley
showed that the males of Antonina crawii Ckll., unlike their counterpart females,
do not exhibit features departing from the usual pseudococcid male type.
Dziedzicka (1961), while studying the developmental stages of Gossyparia spuria
(Modeer), described two forms of the adult males (the brachypterous and the
macropterous) ; her descriptions are concise and her interpretation of the abdominal
segments is inaccurate. Lellakova-Duskova (1965) briefly described the different
male stages of Quadraspidiotus marani Zahradnik, including the adult. Jakubski
(1965) provided short accounts on the available males of the family Margarodidae.
Pesson & Bielenin (1966) redescribed the males of Icerya corticalis Vayssiere ;
they sank it as a synonym of /. maxima Newstead, for which they erected the new
genus Gigantococcus. Giliomee (1967) studied in great detail the males of 23 species
of the family Coccidae ; he recorded few structures not previously observed,
suggested a classification of the family based on the males, and discussed inter and
intra-family relationships within Coccoidea.

MATERIAL, TECHNIQUE AND ILLUSTRATIONS

Material.

The males of 24 species, belonging to 17 genera and two coccid groups were

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 7

studied. The author came to the conclusion that these groups represent two
closely related, though well separated families, Pseudococcidae and Eriococcidae.
Most of the species were represented by the macropterous males only, but some
were either brachypterous (G. salicicola) or apterous (P. fraxini) or with more than
one form available (macropterous and apterous in S. sacchari, and macropterous and
brachypterous in G. spuria). As already mentioned, comprehensive classification
of the families Pseudococcidae and Eriococcidae does not exist ; thus, the species
studied are given in the following list with those of Pseudococcidae arranged in
groups suggested by the results of the present work.

PSEUDOCOCCIDAE

PLANOCOCCUS Group

Planococcus Ferris, 1950.
P. citri (Risso, 1813).
P. kenyae (Le Pelley, 1935).
P. dioscoreae Williams, i960.

Planococcoides Ezzat & McConnell, 1956.
P. ireneus De Lotto, 1963.

Nipaecoccus Sulc, 1945.

N. vastator (Maskell, 1894).
N. nipae (Maskell, 1892).

Maconellicoccus Ezzat, 1958.
M. hirsutus (Green, 1908).

Ferrisiana Takahashi, 1929.

F. virgata (Cockerell, 1893).

Trionymus Berg, 1899.

T. newsteadi (Green, 1917).

PSEUDOCOCCUS Group

Pseudococcus Westwood, 1840.
P. obscurus (Essig, 1909).
P. citriculus Green, 1922.

Dysmicoccus Ferris, 1950.

D. alazon Williams, i960.

SACCHARICOCCUS Group

Saccharicoccus Ferris, 1950.

S. sacchari (Cockerell, 1895).

8 MORPHOLOGY AND TAXONOMY OF ADULT MALES

OCTOCOCCUS Group

Octococcus Hall, 1939.

0. africanus (Brain, 1915).

CEROPUTO Group

Ceroputo Sulc, 1898.

C. pilosellae Sulc, 1898.

Centrococcus Borchsenius, 1948.
C. insolitus (Green, 1908).

Nairobia De Lotto, 1964.

N. bifrons De Lotto, 1964.

NAIROBIA Group

ERIOCOCCIDAE

Eriococcus Targioni-Tozzetti, 1868.
E. araucariae Maskell, 1879.
E. orariensis Hoy, 1954.
E. buxi (Fonscolombe, 1834).

Ovaticoccus Kloet, 1944.

0. agavium Douglas, 1888.

Gossyparia Signoret, 1875.
G. spuria (Modeer, 1778).
G. salicicola Borchsenius, 1949.

Pseudochermes Nitsche, 1895.

P.fraxini (Kaltenbach, i860).

The material was secured from the following three sources :

Material received from other workers.

Specimens of most of the studied species were acquired through the kind co-opera-
tion of many workers from various parts of the world, at the request of Dr. K.
Boratynski (see acknowledgements). Alcohol-preserved material of 15 species,
slide preparations mounted in " Berlese fluid " of one species (E. orariensis), and
dry material of another (E. buxi), were received. The " Berlese fluid " slides were
soaked in warm water until the mountant was dissolved and the specimens released.
The dry material was treated with 10% KOH for 24 hours, and the males were
gently dissected from their puparia with a finely pointed quill.

Material collected in pupal stages.

The writer was able to collect and breed the prepupae and pupae of 10 species ;
these were kept in glass tubes under laboratory conditions and produced adult

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 9

males. Of these, 5 were collected in England and 4 in Egypt, U.A.R. Among
those collected in England were the males of D. alazon, whose pupae were accidentally
found on a bunch of bananas, bought in a London fruit shop and imported from the
Canary Islands. Two others (Pseudococcus fragilis and P. adonidum) presented the
problem of correct identity, which frequently occurs in coccids ; they were found
in a mixed colony, and since the identification still depends largely on the females,
the actual copulation had to be observed as an evidence to identify the males.
Descriptions of these two species are not given here, since they have already been
treated in detail by Giliomee (1961) ; his data, however, were included in my tables
and used in the discussion.

Material obtained from colonies.

Colonies of Pseudococcus obscurus and Planococcus citri on potatoes, and of
Chorizococcus lounsburyi on potted bulbs (Amaryllis sp.) were bred in the laboratory.

It should be noticed that the results of Beardsley's studies (i960, '62, '63, '64,
and '65), on a wide variety on pseudococcid species were also utilized, although not
entered in the tables due to his different style of description.

Preparation and technique.

Theron (1958) introduced a method for mounting the coccid males, in which the
specimens were stained by Chlorazol Black E (saturated solution in absolute methyl
alcohol). His method was basically followed by Ghauri (1962) and Giliomee (1961
and 1967), and also here with slight modifications regarding the clearing (in KOH)
and staining length of time. For details of the method adopted and illustration
of the tools, see Afifi and Kosztarab (1967).

The data, in most species, were taken from 10 specimens each. The length of
the thorax was taken from the postoccipital ridge to the posterior margin of the
mesopostphragma. The length of the prescutum was topographically measured
from its anterior margin to the prescutal suture ; the width of the prescutum, and
also the lengths and widths of other structures (e.g. scutellum, basisternum) included
the boundary ridges. The measurements of the leg segments were taken at their
maximum ; the length of the tarsus only indicates the length of the distal tarsomere.
The length of the abdomen was taken from the mesopostphragma to the anterior
margin of the basal ridge of the penial sheath, or the antero-ventral margin of the
genital capsule.

Part of the material of this study has been deposited in the collections of the
British Museum (Nat. Hist.), London, and in the Department of Entomology,
Ministry of Agriculture, Cairo, Egypt, U.A.R.

Illustrations.

The drawings were made to scale on graph paper, using a square graticule fitted
into the microscope eye-piece. The front view of the head was particularly con-
sidered, to illustrate the relationship between all arms of the midcranial ridge.

io MORPHOLOGY AND TAXONOMY OF ADULT MALES

In the general views of the body, the dorsal and the ventral arms of the midcranial
ridge were drawn just outside the median line, so that the degree of their develop-
ment could be shown.

GENERAL MORPHOLOGY

Newstead (1903) described, in general terms, the males of some species of the
subfamily Dactylopinae (=Pseudococcidae and Eriococcidae), giving some particulars
about the antennae, legs and the genital armature. Brain (1915) gave even shorter
accounts of the males of some species of the subfamily Pseudococcinae. Makel
(1942) was the first to carry out combined morphological and anatomical studies of
3 species of the genus Pseudococcus. Sulc (1943, 1944, 1945) gave comparatively
detailed descriptions and illustrations of the males of Phenacoccus aceris Signoret,
Peukinococcus piceae (Loew), and Nipaecoccus nipae (Mask.), respectively.
Morrison (1945) described the macropterous males of Heterococcus graminicola,
with generalized illustration of the body and more detailed drawings of antenna,
hind leg, abdominal penultimate and genital segments. Reyne (1954) similarly
treated the males of Puto antennatus Signoret, giving detailed accounts of the
structure of the head and the genital segment ; he concluded that the males of
Puto and Macrocerococcus are closely related, and included certain characters (e.g.
number of eyes, structure of penis, number of caudal filaments) widely separating
their type-species (P. antennatus Sign, and M. snperbus Leon.) from that of Phena-
coccus (P. aceris Sign., described by Sulc, 1943). He also supported the conception
of Borchsenius (1948) regarding the re-establishment of the genus Macrocerococcus.
Giliomee (1961) studied the males of Pseudococcus fragilis Brain, P. adonidutn
(Linn), and P. maritimus (Ehrhorn) comparatively ; he recognized two " types "
of the last species, although later (1967) decided that these " types " represent two
distinct species, as was confirmed by the present writer (see description of Pseudo-
coccus obscurus).

In his excellent study on the subject, Theron (1958) recognized 10 general
morphological characters identifying the lecanoid type of male, based on studies of
two genera, Eulecanium (=Parthenolecanium-Coccida.e) and Pseudococcus (=Plano-
cocois-Pseudococcidae) ; he also indicated that the Pseudococcidae differ from
Coccidae by more generalized condition of the genitalia (primitive character), the
condition of the anterior tentorial arms and the ocular ridges (specialized characters).
Giliomee (1961) found that two of Theron's generalized characters do not apply to
Pseudococcidae and this was confirmed by the present study. Later, Giliomee
(1967) discussed the relationships between the males of Pseudococcidae and
Coccidae, listing 10 morphological characters (5 primitive and 5 specialized) separat-
ing the males of the two families. This again was confirmed by the present author.

It was here found that the pseudococcid and the eriococcid males differ by a
number of morphological characters (listed later in the discussion), the importance
of which is comparable with those separating Pseudococcidae and Coccidae. There-
fore, it has been concluded that they deserve the status of separate families and
will be here treated as such.

OF I'SEl'DOCOCCIDAE & ERIOCOCCIDAE n

The homologies and terminology of the various structures introduced by Theron
(1958), and later supplemented by Ghauri (1962) and Giliomee (1961, 1967) are here
adopted ; a few additional terms and abbreviations have also been employed.

PSEUDOCOCCIDAE

General Characteristics

All the studied species were represented by ruacropterous males, and one species
(S. sacchari) was also represented by apterous specimens. Brachypterous forms
were reported by earlier authors (Pahnicola palmarum, described by Beardsley,
i960), but were not available for the present study.

Pupation of the males (at least of the species obtained from pupae by the writer
in the laboratory) takes place inside fluffy puparia of waxy threads ; the adult
male emerges backing through the loosely felted posterior extremity of the
puparium.

Appearance. The body, as in all other Coccoidea (Theron, 1958, and Ghauri,
1962), consists of the well defined head (with non-functional mouth parts), thorax
and abdomen. Usually the males are narrow and slender, although those of some
of the species studied are moderately robust (0. ajricanus) ; broadest at the thoracic
region and usually round throughout, but sometimes dorso-ventrally flattened
(S. sacchari). The head subtriangular in dorsal view with the genae bulging
posteriorly. The neck region is neither entirely absent as in Diaspididae (Ghauri,
1962), nor distinctly pronounced as in many Coccidae (Giliomee, 1967), but merely
indicated by a distinct constriction. The lrj^s arc well developed, usually long and
slender. The alate forms have the anterior pair of wings well developed, and the
posterior wings modified into hamulohalterae, each normally with one apically
hooked seta. The abdomen is largely membranous, becoming gradually narrower
posteriorly ; with eight well separated pregenital and the terminal genital segments.

Size. The coccid males generally are very small insects and although the actual
size varies considerably, they are comparatively large in some families and small
in others. The males of Pseudococcidae occupy an intermediate position in that
respect, i.e. smaller than Margarodidae and some Coccidae, and larger than
Diaspididae, which are apparently the smallest of all. Among the species studied,
the smallest was N. nipae (840-980, average 896[x long), and the largest F. virgata
(1274-1596, av. 1386(1 long).

Colour. Examination of the available living material of a few species indicates
that the colour of the males, although somewhat variable, is apparently character-
istic of each species. The basic colour is always brownish, light (P. citri), or dark
(T. newsteadi), or with yellowish or greenish tinge (M. hirsutus and D. alazon,
respectively) ; the eyes usually are also brown but sometimes dark red (D. alazon).
In alcohol-preserved material, the colour gradually fades away, becoming indefinable
and the differences disappear ; unfortunately most of the species studied were
alcohol-preserved.

12 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Derm Vestiture. The body setae : These are of two main types : The fleshy
(Giliomee, 1961) or " digitiform " (Beardsley, i960), which are comparatively thick
and apically obtuse ; and the hair-like (Giliomee, I.e.) or " filamentous " (Beardsley,
I.e.), which are much thinner, with very acute tips.

The fleshy setae may occur on any part of the body itself (i.e. on the head, thorax
or abdomen), as well as on the antennae and legs (the Pseudococcus group), or their
presence may be limited to the appendages only (most species of other groups),
occurring both on the antennae and on the legs (P. citri), or on the antennae only
(N. vastator) ; or are absent altogether (C. pilosellae). The hair-like setae always
occur on the body and on the appendages. Other types of more specialized setae
will be described later, together with the other details of the parts on which they
occur.

The disc pores. With the exception of Nairobia group, the males of Pseudo-
coccidae carry a number of disc pores, sometimes on the thorax and the abdomen
only (e.g. Octococcus group), or also on the head (e.g. Pseudococcus group). In the
material studied, the pores were usually quadrilocular, occasionally trilocular,
quinquelocular or 6-locular ; in some species, Beardsley (i960) observed pores with
more than 6 peripheral loculi.

MACROPTEROUS MALES

Head

The head capsule of the coccid male, as discussed by Theron (1958), is almost
entirely made up of the epicranium ; the absence of the functional mouth parts
has resulted in a considerable reduction of the " vorderkopf " and absence of the
labium.

The head has the form of a somewhat irregular tetrahedron ; subtriangular in
dorsal, lateral and frontal views ; broadest posteriorly across the genae, and
becoming gradually narrower anteriorly towards the truncate apex and antero-
ventrally towards the cone on which the ventral eyes are situated. Between the
apex of the head and the ventral cone, the surface of the head is often depressed ;
this ventral preocular depression (vprd) may be deep and conspicuous (e.g. P.
dioscoreae) , or shallow and ill-defined (e.g. P. citri) ; in the Saccharicoccus group, the
depression is absent.

The head capsule is reinforced by a number of ridges, of variable shape, degree of
development, sclerotization, etc. The midcranial ridge (mcr) with its longitudinal
dorsal (dmcr), ventral (vmcr), and the apical transverse lateral arms (lmcr), gives
support to the anterior part of the head. The dorsal arm is usually distinct although
slender, but sometimes reduced and its position only marked by heavy sclerotization
(S. sacchari), or absent altogether (F. virgata). This arm, when present, posteriorly
reaches at least to the level of the dorsal eyes and then fades away (e.g. P. citri),
or extends further back to meet or almost meet the postoccipital ridge (e.g. P.
obscuras). Anteriorly, the dorsal arm is usually separated from the other arms
by a very short distance. The ventral arm anteriorly gives off two branches, the
lateral arms, forming a Y, or occasionally T-shaped ridge (N. vastator, Text-fig.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 13

11B) at the apex of the head ; posteriorly, the ventral arm disappears before
reaching the level of the ventral eyes. In Ceroputo and Nairobia groups, all arms
of the midcranial ridge meet at the apex of the head, and a cruciform structure is
formed (Text-figs. 33B, 35B, 37B). The lateral arms are usually well developed,
but sometimes reduced (N. nipae). The postoccipital ridge (por) is slender and
usually distinct although sometimes only marked by a sclerotized stripe (C. pilo-
sellae) ; the ridge is U-shaped and medially continuous in almost all the species,
but in S. sacchari it is V-shaped and medially interrupted. According to Makel
(1942), the postoccipital ridge serves for the attachment of the cephalothoracic
muscles. The area bounded posteriorly by this ridge is the dorsomedial part of
the epicranium (dmep), which corresponds to the median crest in Diaspididae (Theron,
1958 and Ghauri, 1962) ; this area is slightly raised, well sclerotized, without
polygonal reticulation.

The preocular (procr) and postocular (pocr) ridges were described as being fused
below the lateral ocellus to form the characteristic Y-shaped structure on each side
of the head (Theron, 1958 ; Giliomee, 1961), and this condition was considered by
Theron as " a minor specialization " in Pseudococcidea. The present study showed
that indeed this condition obtains in most species studied, but certain differences
in details in a few of them, as well as the conditions described by Giliomee (1967)
in some species of Coccidae, indicate that the interpretation of this " fusion " is
rather inaccurate. Giliomee (I.e.) found, in Eriopeltis spp. for example, that the
pre- and postocular ridges are connected just below the ocellus by a longitudinal
inter ocular ridge (ior), giving support to the preocular ridge and its articulation with
the antennae ; he suggested that this condition may be a forerunner of the situation
in Pseudococcidae where the two ridges are fused. The present author found that
most Pseudococcidae have the connecting interocular ridge well developed, and in
some species (e.g. P. citri, Text-fig. 5 and T. newsteadi, Text-fig. 20) a short, though
well marked lower part of the preocular ridge extends below the point of junction.
It appears, therefore, that the Y-shaped ridge on the side of the head is a complex
structure of : (1) the upper part of the preocular ridge, intimately fused with (2)
the interocular ridge, which joins (3) the postocular ridge below the ocellus, with
simultaneous more or less considerable reduction of the lower part of the preocular
ridge. This condition, although reminiscent of that found in some species of
Coccidae, is not necessarily evolved from it, but probably developed independently
within Pseudococcidae and represents a specialization of this family. It also seems
that this Y-shaped fusion of the ocular ridges serves to render support to the weakly-
developed preocular ridge and its articulation with the antennae.

In Ceroputo and Nairobia groups the pre- and postocular ridges are well separated,
but a longer (C. pilosellae, Text-fig. 34), or a shorter (N. bifrons, Text-fig. 38)
rudiment of the connecting ridge is present, arising from the anterior edge of the
postocular ridge just below the ocellus and extending anteriorly towards the
preocular ridge. An aberrant condition was found in C. insolitus (Text-fig. 36),
where a ridge anteriorly arising from the postocular ridge and directed towards

i 4 MORPHOLOGY AND TAXONOMY OF ADULT MALES

the preocular ridge is present but above the ocellus, thus it is not strictly homologous
with the interocular ridge ; nevertheless, it probably serves the same purpose and
may represent another independent morphological means to the same functional end.

The ocular sclerites (ocs) are weakly sclerotized plates on each side of the head ;
rather large in Ceropnto and Nairobia groups (where the ocular ridges are separated),
but are comparatively smaller and traversed on each side by the interocular ridge,
in the remaining groups.

The preoral ridge (pror) is extremely slender and latero-posteriorly connected to
both the postocular ridge and to the proepisternum + cervical sclerite, by means
of a small triangular and weakly sclerotized plate.

Two pairs of simple eyes (accessory eyes, Berlese, 1893 ; ocelli, Green, 1922)
surrounded by a narrow area of polygonal reticulation, and one pair of transparent
lateral ocelli (o) (primary eyes, Krecker, 1909 ; rudimentary eyes, Green, 1922) are
present in the species studied. Beardsley (1962) recorded 7 pairs of simple eyes
and a pair of ocelli in one species, Puto yuccae, and the complete absence of the
ocelli in another, Rhizoecus falcifer. The widely separated dorsal eyes (dse) are
borne on the dorsal part of the ocular sclerites, between pre- and postocular ridges.
The ventral eyes (vse) are much approximated and placed on the ventral protrusion
of the head ; these, although usually somewhat larger than the dorsal eyes, are
sometimes smaller (N. vastator), or both subequal (e.g. C. pilosellae). The ocelli
are usually large and well developed ; in the Planococcus, Pseudococcus, Sacchari-
coccus and Octococcus groups the ocelli are situated at the base of the fork of the
Y-shaped complex of ocular ridges, and are dorsally supported by a slender ocellar
ridge (see lateral views) ; in Ceroputo group, where the Y-shaped complex is absent,
they are supported by the postocular ridge and either a rudimentary sclerotized
projection (Text-fig. 34), or a ridge-like arm (Text-fig. 36). In Nairobia group, the
ocelli are vestigial and merely represented by atrophied spots.

The genae (g) are membranous, laterally bulging behind the postocular ridges and
without any reticulation ; they form the latero-posterior margins of the head.

The ventral cavity (vc) is a longitudinal, narrow, slit-like invagination in the
median line of the head. From the roof of this invagination arises the internal
cranial apophysis (ca) which is always apically truncate ; according to Theron
(1958), the cranial apophysis serves for the attachment of the antennal muscles.

The non-functional month opening (mo) is small, situated behind the preoral
ridge. The " tendon-like apodeme " described in other families (Theron, I.e.) is
absent in the species here studied. The posterior tentorial pits (ptp), from which
the internal posterior tentorial arms (pta) originate, are minute and placed on the
membrane on each side of the mouth opening. The posterior arms are connected
with each other by means of the transverse slender tentorial bridge (tb). The
anterior tentorial arms (ata) usually fuse just before meeting the cranial apophysis,
but in Ceroputo and Nairobia groups they are well separated ; the anterior tentorial
pits, therefore, are not visible externally.

The hair-like setae are always present on the head ; in the Pseudococcus group

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 15

the fleshy setae are also present. The head setae are generally arranged in the
following groups :

(1) The dorsal head setae (dhs) : These occur anterior to the postoccipital ridge.

(2) Thegenal setae (gs), are present dorso-laterally on the genae ; sometimes there
are 3 setae or less (e.g. P. citri), 7 or more (Pseudoccus group), or they occur in
intermediate numbers (T. newsteadi).

(3) The setae of the ocular sclerites (ocse) : These are present in the Psendococcus
group only, and occur in two conditions relative to their topography : (a) the
dorsal ocular setae (dos) occurring on the dorsal part of the ocular sclerites, were
found only in one species, D. alazon ; and (b) the ventral ocular setae (vos), which
occur on the ventral part of the ocular sclerites.

(4) The ventral head setae (vhs) are arranged in 3 distinct groups : (a) a group
along the longitudinal median line between the ventral eyes ; these setae may be
always present (Planococcus, Pseudococcus and Saccharicoccus groups), present or
absent (Nairobia group), or entirely absent (Oclococcus and Ceroputo groups) ; (b)
a group forming a transverse band across the area of the ventral preocular depression,
just anterior to the ventral eyes ; (c) a group of usually a few setae, arranged in a
single line on each side of the ventral arm of the midcranial ridge.

One to three dorsal head pores (dhp) are usually present on each side of the mid-
cranial ridge, near the base of the antennae (A and B in the figures), but in some
species (P. dioscoreae and S. sacchari) 4 or more pores are present. In Ceroputo
group, however, there is a ventral pore (vhp) occurring at least on one side of the
head, and the dorsal pore may be absent (C. pilosellae, Text-fig. 33B), or also present
(C. insolitus, Text-fig. 35B) ; in some other species (F. virgata) the head pores are
absent altogether.

Antennae

The scape is inserted laterally at the anterior apex of the head and articulates
with the antennal process of the preocular ridge ; the lateral arms of the midcranial
ridge, when well developed, also extend towards the base of the scape but with
which no articulation takes place. The antennae are typical filiform, normally
10-segmented, and vary considerably in length within the family ; in Octococcus
and Ceroputo groups they are comparatively long (the body usually less than i-6
times longer) ; in Pseudococcus and many species of Planococcus groups they are
rather short (the body 1-6-2 -5 times longer) ; in Nairobia and some species of
Planococcus groups (Nipaecoccus spp.) the antennae are intermediate in length ;
in Saccharicoccus group they are very short (the body more than 2-5 times longer).
The flagellar segments of the pseudococcid males are cylindrical and usually con-
spicuously longer than wide.

The scape (scp) is the shortest and the widest (at the base) of the antennal segments.
It becomes narrower distally, with the dorsal margin usually longer than the ventral ;
the basal part is sclerotized and laterally articulates with the articular process of
the preocular ridge. Distally, the scape articulates with the pedicel by means of a

16 MORPHOLOGY AND TAXONOMY OF ADULT MALES

process, into which the ventral sclerotization is produced. Usually 4 hair-like setae
are present on the scape, but in Pseudococcus, Saccharicoccus and some species of
Planococcus groups (e.g. P. citri), the scape carries more than 4 setae ; there are
no other types of setae on the scape.

The pedicel (pdc) is narrow at the base and widest near the distal end, thus having
a club-shaped appearance. At the base, the pedicel is ventrally supported by a
ridge, which provides a minute process articulating with the scape. At the distal
end of the pedicel, a narrow area of polygonal reticulation is present, where a small
circular plate, presumably a sensillum placodeum (spl), is dorsally borne. The
pedicel carries fleshy and hair-like setae.

The flagellum includes the intermediate (III to IX) and the terminal segments.
The surface of these segments is somewhat irregular, and they are usually well
separated, although partial fusion between two or more adjacent segments is
frequent in some species (e.g. P. ireneus). Segment III is club-shaped, with a short
constricted peduncle inserted into the pedicel ; this segment is usually the longest ;
at least subequal in length to the terminal segment (Saccharicoccus, Nairobia and
some species of Planococcus groups), or often up to i| times longer (Pseudococcus,
Octococcus and other species of Planococcus groups), or even more (Ceroputo group).
The width of the 3rd segment in relation to its length is also variable ; sometimes
the length is 2-3-3 times the width (Saccharicoccus and many species of Planococcus
groups, e.g. P. ireneus and N. vastator), 3-3-5-3 times (Pseudococcus, Octococcus,
Nairobia and other species of Planococcus groups), or more than 5-3 times (Ceroputo
group). Segments IV to X are relatively long in comparison to their width ; the
terminal segment elongate, with a constricted, rounded or pointed apex.

Antenna! setae : The following types of setae occur on the antennal segments :

(a) The fleshy setae (is), usually present on segments II to X and represent the
majority of the antennal setae, but sometimes the fleshy setae are entirely absent

(C. pilosellae).

(b) The hair-like setae (hs) , which always occur at least on the first three antennal
segments.

(c) The subapical sensory setae (set. scla) are apically knobbed and occur on the
terminal segment only, but sometimes are absent altogether (Ceroputo group).

(d) The capitate sensory setae (set. ca) : This group includes setae similar to the
latter type but present on the antennal segments III to X ; these setae were found
in Octococcus and Nairobia groups only.

(e) The antennal bristles (ab) are the stoutest bristle-like setae, usually longer
than the fleshy setae and present on the last three terminal segments ; segments
VIII and IX always with one ventral bristle, at about half the length of these
segments. The terminal segment (Xth) with at least 3 such preapical bristles,
one dorsal and one on each side ; in Ceroputo and some species of Planococcus
groups (N. vastator), two additional and conspicuously smaller bristles are also
present latero-ventrally, at a greater distance from the apex of the segment.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 17

Thorax

Makel (1942) was the first to give adequate attention to the pleural region of the
thorax ; Ezzat (1956) gave more detailed illustrations, but Theron's (1958)
identification of the thoracic structures reached an incomparably higher standard.

Prothorax

The prothorax is distinctly separated from the head by the constricted neck ;
in this respect, Pseudococcidae and Eriococcidae occupy an intermediate position
between Diaspididae, where the neck region is not differentiated, and most Coccidae
where the neck is very well pronounced. The pronotum is represented by the
following structures : (1) the collar-like transverse pronotal ridges (prnr), which are
medially interrupted and extend laterally on each side ; (2) two small lateral
pronotal sclerites (prn), one on each side, anteriorly bounded by the pronotal ridge ;
(3) a postero-dorsal pair of small sclerites, the post-tergites (pt).

The proepisternum is a distinct, triangular and well sclerotized pleural sclerite,
on each side of the prothorax ; anteriorly it fuses with the rudiment of the cervical
sclerite at a weak point to form one structure, the proepisternum -\- cervical sclerite
(pepcv), which is dorsally approached near its anterior end by the pronotal ridge,
and anteriorly articulates with the postocular ridge (see lateral views). Posteriorly,
this sclerite (pepcv) is fused with a short pleural ridge (plr^, which carries a small
invaginated pleural apophysis (plai) ; the pleural ridge extends downwards and
articulates with the basal process of the coxa. The proepimeron, which is usually
apparent in certain families (Margarodidae and Diaspididae), is entirely indis-
tinguishable in Pseudococcidae (and Eriococcidae).

The dorsal margin of the proepisternum is always heavily sclerotized and ridge-
like ; in Ceroputo group, the ventral margin also is similarly sclerotized.

The prosternum (stni) is reduced to a small and usually triangular plate ; in some
species (N. bifrons) the plate is very short and wide, and in others (T. newsteadi)
comparatively long and narrow ; in C. insolitus (Text-fig. 35) it is represented by
two small separate plates. Posteriorly, the prosternum is usually bounded by a
narrow transverse prosternal ridge (stnlr) , which however is absent in Saccharicoccus
group and replaced by heavier sclerotization of the posterior margin (Text-fig. 27) ;
the median longitudinal ridge of the prosternum, which occurs in many members of
the other families (Margarodidae, Coccidae and Diaspididae), is absent in Pseudo-
coccidae (and Eriococcidae).

Dermal structures of the prothorax. The hair-like setae are always present on the
prothorax ; in Pseudococcus group, the fleshy setae also occur. These setae are
arranged in the following groups :

(1) The medial pronotal setae (mpns), which are present on the median and sub-
median areas between the pronotal ridge and the post-tergites ; 3 or more of these
setae are usually present on each side of the median line (P. citriculus, Text-fig. 23),
but in other species they are less than 3 (e.g. P. citri, Text-fig. 4), or absent altogether
(Octococcus, Ceroputo and Nairobia groups).

18 MORPHOLOGY AND TAXONOMY OF ADULT MALES

(2) The post-tergital setae (pts) occur on, or immediately behind, the post-tergites ;
this group is present in F. virgata among the Planococcus group, and in all species
of the Pseudococcus and Saccharicoccus group, and may be also present in Ceroputo
group.

(3) The lateral pronotal setae (lps) are situated on, or lateral to the lateral pronotal
sclerites.

(4) The antespiracular dorsal setae (asds) occur laterally at about the level of the
post-tergites ; 2-4 setae are usually present on each side, but in Saccharicoccus
group there are 5 or more setae, and in Octococcus and Ceroputo groups they are
usually absent.

(5) The antespiracular ventral setae (asvs) are present latero-ventrally, just behind
the front coxa ; one seta almost always occurs on each side, but in S. sacchari two
setae were found.

(6) The prosternal setae (stnis) are present medially, on the prosternum or on the
membranous part immediately anterior to it ; these setae may be less than 4 on
each side (Planococcus and Nairobia groups), 4 or more (Pseudococcus group), or
usually absent (Octococcus and Ceroputo groups).

The disc pores. In the groups of genera other than Nairobia, the disc pores are
present, usually occurring in association with the prothoracic setae, and are arranged
in the following groups : the medial pronotal pores (mpnp) ; the post-tergital pores
(ptdp) ; the lateral pronotal pores (lpp) ; the antespiracular dorsal pores (asdp) ;
and the prosternal pores (stnip). The number of these pores is variable, but the
post-tergital pores were found in P. dioscoreae only (Text-figs. 7, 8).

Mesothorax

The mesothorax is strongly developed. The shape and conditions of the sclero-
tized areas vary considerably within the family, providing a number of taxonomic-
ally important characters.

The mesotergum is divided into a mesonotum (or alinotum) and a mesopostnotum,
which are widely separated by a large membranous area (postscutellum of Berlese,
1893) ; the mesonotum is further subdivided into the distinct prescutum, scutum
and scutellum.

The prescutum (prsc), the antero-median area of the mesonotum, is dome-shaped ;
in the dorsal view it is either transversely rectangular (Planococcus, Pseudococcus,
Saccharicoccus and Octococcus groups), or triangular (Ceruputo and Nairobia groups).
The anterior margin of the prescutum is invaginated, forming the mesoprephragma
(phri) with its inner margin slightly notched in the middle. The prescutum is
bounded laterally and posteriorly by the prescutal ridges (pscr) and the prescutal
suture (pscs), respectively ; the suture is sometimes absent (Saccharicoccus group),
or strongly developed, ridge-like and continuous with the prescutal ridges (Ceroputo
group, Text-figs. 33 & 35).

The scutum (set) is large, uniformly sclerotized throughout (Ceroputo and Nairobia
groups), or with a median longitudinal narrow membranous area (the remaining

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 19

groups), and with the anterolateral extensions surrounding the prescutum latero-
posteriorly. These lateral extensions are produced into the prealares (pra), from
which they are separated by the secondary prealar ridges (prar) ; the latter are
anteriorly invaginated into small finger-like apodemes (a), and posteriorly support
small anterior notal wing processes (anp). Each prealare is laterally differentiated
into a strongly sclerotized triangular plate (tp) which forms the antero-dorsal
boundary of the episternum ; this plate was called " prealar wing process " by
Ezzat (1956). Behind the anterior notal wing processes, the postero-lateral exten-
sions of the scutum are inflected downwards and then upwards to form the posterior
notal wing processes (pnp) which are attached to the postalare.

The scutellum (scl) is pentagonal in dorsal view ; antero-laterally bounded by the
scutoscutellar suture (sets), and posteriorly by the inward fold of the posterior margin
of the notum (rd) ; the semi-cylindrical structure of the scutellum is due to deep
inward inflection of its anterior and posterior margins ; these invaginated margins
do not meet internally, thus the scutellar foramen which occurs in other families,
e.g. Margarodidae, Diaspididae and Coccidae (Theron, 1958 •; Ghauri, 1962 and
Giliomee, 1967) is absent in Pseudococcidae (and Eriococcidae).

The relative lengths and widths of the various mesonotal structures differ con-
siderably within the family and the following ratios were found of taxonomic
importance : length to width of prescutum ; length of prescutum to length of
scutum ; length to width of scutellum ; length of scutellum to length of scutum.

The posteriormost part of the mesotergum, the mesopostnotum, is overlapped by
the metanotum and thus externally invisible. This invaginated part constitutes
the mesopostphragma (phr2) which is slightly larger than the mesoprephragma,
and also has a small median notch. Laterally, the postnotum gives rise to a pair
of strong finger-like postnotal apophyses (pna) within the mesothoracic cavity.
Antero-lateral of the apophysis, the postnotum is produced into a postalare (pa)
which reaches the pleural area and articulates with the pleural ridge of the meso-
thorax. The postalare is reinforced by the distinctly separated anterior and
posterior postalar ridges (apar & ppar) ; dorsally, it bears two small processes
associated with the posterior margin of the wing and the posterior marginal fold
of the notum, respectively.

The mesopleuron : The mesopleural ridge (plr2) posteriorly articulates with the
base of the coxa, and extends antero-dorsally towards the base of the wing. At
about half length, the ridge makes the characteristic sharp double bend and carries
at this point the internal mesopleural apophysis (pla2) ; externally, this part of the
ridge is overlapped by the postalare. The ridge is interrupted at a short distance
above the coxal articulation ; this interruption was overlooked by Theron (1958)
on describing P. citri, and according to Ghauri (1962) and Giliomee (1967) does not
occur in Diaspididae and many Coccidae. The dorsal part of the (plr2) is relatively
strong and terminates in a semi-circular pleural wing process (pwp2) ; the anterior
margin of the latter is connected with the episternum by means of a ridge-like
basalare (bas). Behind the pleural wing process, there is a small sclerite, the
subalare (sa), whose articulation with the alary sclerites is rather obscure. The

20 MORPHOLOGY AND TAXONOMY OF ADULT MALES

subepisternal ridge (ser) articulates dorsally with the triangular plate and extends
obliquely to approach the marginal ridge of the basisternum, thus bounding the
mesepisternum (eps2) anteriorly. The mesepisternum is divided into a larger
dorsal part, the supraepisternum or anepisternum, and the small ventral part, the
infraepisternum or katepisternum (Snodgrass, 1935), both of which are separated
by a membranous area. The katepisternum anteriorly fuses with the later opleurite
(lpl) which is attached to the lateral arms of the marginal ridge of the basisternum ;
the lateropleurite is narrow (e.g. Planococcus group), but comparatively wide (e.g.
Nairobia group). A small and well sclerotized mesepimeron (epm2) is present
immediately behind the pleural ridge, just above its articulation with the coxa.
A slender sclerite, presumably representing the trochantin (tn) occurs just anterior
to the coxal articulation, and was found in all the species studied except N. bifrons.
This sclerite was first described and illustrated by Giliomee (1961).

The mesothoracic spiracle (sp2) with its atrium and its supporting bar, the peri-
treme, is placed latero-ventrally anterior to the subepisternal ridge.

The mesosternum is represented by a large, slightly convex and hexagonal plate,
the basisternum (stn2) ; this plate is framed antero-laterally and latero-posteriorly
by the marginal and the precoxal ridges (mr & pcr2), respectively. In Saccharicoccus
group the median part of the marginal ridge is completely absent (Text-figs. 27-30).
The precoxal ridge posteriorly fades away before reaching the median line, where
the posterior margin of the basisternum becomes slightly inflected to form a furcal
pit (fp) from which the strong, two-armed furca (f) originates. The longitudinal
median ridge of the basisternum, which occurs in many species of Margarodidae,
Diaspididae and Coccidae (Theron, 1958 ; Ghauri, 1962 and Giliomee, 1967), is
absent in Pseudococcidae (and Eriococcidae).

The wing articulation : The articular system of the fore wings is apparently the
same in all the coccids, and only slight differences are shown in the size of the alar
sclerites, the pter alia, which include : the tegula (teg), the axillary (first, second and
third) (axi, ax2 and ax3) and the additional sclerites (asc). The costal complex of
veins (ccx) and the axillary cord (axe) give support to the anterior and posterior
margins of the wings at their bases. Apart from these structures, the articulation
also involves the anterior notal wing process, the pleural wing process, the basalare
and probably the subalare.

The dermal structures of the mesothorax : The fleshy setae occur in the Pseudo-
coccus group only, in the postmesostigmatal area, but the hair-like setae are always
present, arranged in the following topographical groups : the prescutal setae (pscse)
the scutal setae (sctse) ; the scutellar setae (scls) ; the tegular setae (tegs) ; the
postmesostigmatal setae (pms) ; the basisternal setae (stn2s). The postmesostigmatal
setae may be absent (Nairobia group) or present only laterally behind the meso-
thoracic spiracles (e.g. P. kenyae, Text-fig. 3), or also in the median and the sub-
median areas in a transverse band (e.g. P. citriculus, Text-fig. 23).

The numbers of the mesothoracic setae vary in the different species and con-
stitute useful key characters.

With the exception of Nairobia group, the mesothorax usually carries a number

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 21

of disc pores, namely the mesospiracular disc pores (sp2p) and the median or sub-
median postmesostigmatal pores (pmp).

Metathorax

The metathorax is largely membranous and the metathoracic sclerites are con-
siderably reduced, as a result of the modification of the hind pair of wings into
hamulohalterae. The metanotum overlaps the inflected mesopostnotum and is
indicated externally by a pair of small lateral suspensorial sclerites (ss), attached
to the base of the hamulohalterae by means of a fine sclerotized tendon. The
metapostnotum (pn3) is represented at the posterior extremity of the metathorax
by small subtriangular sclerites, one on each side of the median line ; these sclerites
are usually connected by a distinct transverse metapostnotal ridge (pn3r) (Piano-
coccus, Pseudococcus and Saccharicoccus groups) ; this ridge is weakly developed
in Octococcus and Ceroputo groups (Text-figs. 31-36), and entirely absent in Nairobia
group (Text-fig. 37). The metapostnotal ridge, which is apparently absent in all
other studied families, was overlooked by Theron (1958) and illustrated but not
discussed by Giliomee (1961).

The metapleuron : The metapleural ridge (plr3) is well developed and articulates
ventrally with the hind coxa ; as the ridge obliquely extends antero-dorsally, it
attenuates near the middle where a small metapleural apophysis (pla3) is inwardly
invaginated ; at the lower part, the ridge separates two irregular, sclerotized areas,
the anterior metepisternutn (eps3), and the posterior metepimeron (epm3) ; dorsally
it supports a minute metapleural wing process (pwp3). The precoxal ridge of the
metathorax (pcr3) originates below the episternum and extends ventro-medially ;
this ridge, while being well developed in Planococcus, Pseudococcus, Saccharicoccus
and Octococcus groups, is comparatively weak in Ceroputo and Nairobia groups.
The metathoracic spiracle (sp3) is identical with that of the mesothorax, and lies
ventro-laterally in the membrane anterior to the metepisternum.

The metasternum is represented by a pair of small submedian metasternal apophyses
(sta) ; according to Ghauri (1962) and Giliomee (1967) these apophyses are absent
in Diaspididae and Coccidae. The metasternal plates are absent in Pseudoccidae
(and Eriococcidae).

The dermal structures of the metathorax : Fleshy setae occur in Pseudococcus
group only, but the hair-like setae are present in all species. The metathoracic
setae include the following groups : The metatergal setae (mts), anterior to the
metapostnotal sclerites ; the metapleural setae (mps), just behind the metathoracic
spiracles ; the anterior metasternal setae (amss), in the median and the submedian
areas on the membrane between the basisternum and the metasternal apophyses ;
the postmetastigmatal setae (eps3s), on the metepisternum along its antero- ventral
margin, or just below it on the membranous area (this group of setae was found in
P. citriculus only) ; and the posterior metasternal setae (pmss), medially on the
metasternal membrane behind the metasternal apophyses.

The following groups of metathoracic derm pores, associated with the corres-
ponding groups of setae, are also recognized : The metatergal pores (mtp) ; the

22 MORPHOLOGY AND TAXONOMY OF ADULT MALES

metaspiracular pores (sp3p) ; the anterior metasternal pores (amsp) ; and the
posterior metasternal pores (pmsp).

Wings and Hamulohalterae

The fore wings are membranous, large and elongate, rounded distally and narrow
basally ; the posterior margin of the wing is produced near its base to form a small
alar lobe (al) which is adapted to receive the apically hooked seta of the hamulo-
haltera. The entire surface of the wing (except the area of the alar lobe) is evenly
covered with minute hairs, the microtrichia. The venation is reduced to two veins,
radius (rad) and media (med). Near their bases, the wings usually carry a variable
number of alar setae (als), and a compact dorsal row of few circular sensor ia (sens.),
just anterior to the radial vein.

The metathoracic wings are modified into small and elongate hamulohalterae (h).
This term (singular hamulohaltera), was recently proposed for the coccid males by
Kawecki (1965) as a substitute to what was referred to as " pseudohalteres " in his
earlier paper (1958b), or as " halteres " by other authors. On explaining the
original definition of the word " haltere ", Kawecki showed how it could be applied
correctly to the second pair of wings in Diptera and male Strepsiptera, but emphas-
ized that their different appearance, structure and function in the males of
Coccoidea demanded the adoption of another term. He also referred to the term
" retinacoli ", used by earlier authors, maintaining that it is equally erroneous
because it denotes an apparatus for fastening the wings, located on the anterior
pair. The writer feels therefore that the proposed term seems to be more acceptable.

The hamulohaltera is membranous, supported along its anterior margin by a
weak and slender hamulohalteral ridge (hr). This ridge was overlooked by many
writers, but Jancke (1955) referred to a strengthening of the anterior margin of the
hamulohaltera, and Giliomee (1967) described it as resembling a wing vein.

Legs

Three pairs of well developed legs, of variable lengths and stoutness, are present.
The fore legs are usually the shortest and the hind legs the longest, but sometimes
the middle legs are the shortest (Ceroputo group), or the fore and the middle legs
subequal (e.g. Nipaecoccus spp.). The legs consist of the usual segments, all of
which (except the claw) are beset with numerous hair-like, and usually fleshy
setae also.

The coxa (ex) is supported at its broad base by a well developed ridge ; this ridge
provides a small process dorsally, which articulates with the pleural ridge. The
distal end of the coxa is narrower, and also supported by a ridge, which provides
two processes, one on each side, serving for the articulation with the trochanter.

The trochanter (tr) with sclerotized basal and distal margins ; this segment is
divided into a relatively long proximal part and a short distal one, by means of a
distinct constriction supported by a weakly developed short ridge. One compara-
tively long hair-like apical seta (ase) may occur on the outer side of the distal part
(e.g. N. vastator). Three circular sensilla are usually arranged in a triangle on each
side, but in few individuals two or four on one side were observed.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 23

The femur (fern) is relatively long and stout, broadest near the distal end, where
a ridge with two processes is well developed.

The tibia (tib) is usually the longest segment of the leg, but in few species (P.
irenens and 5. sacchari) the tibia of the fore leg is somewhat shorter than the femur ;
at its base, the tibia carries a ridge with two processes which articulate with the
corresponding processes of the femur. The distal end of the tibia carries a pair
of setae modified into strong spurs, and a varying number of smaller spines.

The tarsus (tar) is elongate, broad at the basal half and tapered slightly distally ;
composed of two tarsomeres, the first of which is very short, and the second much
longer. A campaniform sensillum (cam.s) always present on the dorsal surface,
at the base of the second tarsomere. Two long and apically-knobbed tarsal
digitules (tdgt) are usually present near the apex of the tarsus ; these digitules are
absent in both Ceroputo and Nairobia groups.

The claw (cl) is slightly curved, uniformly tapering to a sharply pointed tip, and
dorsally articulates with the tarsus ; the ungual digitules (udgt), one on each side
of the claw, are fine, with acute tips.

The ratios of several measurements of the legs were found to be taxonomically
useful and the following are here employed : the length of the hind leg to the total
body length ; the width of the hind femur to its length ; and the length of the
femur to the length of the tibia in the front leg.

Abdomen

The abdomen consists of the largely membranous pregenital segments and the
sclerotized genital segment. The segmental boundaries are rather obscure, but
the segmentation could be readily determined by the segmental arrangement of the
setae and derm pores.

Pregenital segments

Eight pregenital abdominal segments are recognized, and considered to represent
either I to VIII abdominal segments by some authors (Theron, 1958 ; Ghauri,
1962 and Giliomee, 1967), and also here adopted), or II to IX by the others
(Beardsley, i960) ; Beardsley explained that his interpretation was based on the
position of the posterior dorsal ostioles, equating their position with that established
by Ferris (1950) for the females ; Ferris regarded the first abdominal segment
(in the females) as completely absent, and that the posterior ostioles lie on what is
morphologically the 7th abdominal segment. In a subsequent paper, Beardsley
(1962) discussed Theron and Giliomee's views, and considered the abdominal
segmentation in Pseudococcidae as an unsettled matter.

The first abdominal segment is developed dorsally and laterally only, and is
indiscernible ventrally ; this is indicated by a pair of small tergites, one on each
side of the median line ; similar, and even smaller tergites are usually also present
on segment II and sometimes III. No tergites occur on segments IV to VII, but
segment VIII carries a large, median tergal plate (at8). The ventral sclerotization

24 MORPHOLOGY AND TAXONOMY OF ADULT MALES

is much reduced and only a pair of small, usually triangular plates represent the
sternites of segment VIII.

The abdominal segment VI usually bears a pair of dorsal ostioles (ost) situated
sublaterally near the posterior margin of the segment ; these ostioles are sometimes
well developed, prominent and with a slit-like orifice (e.g. P. citri), or ill-defined
(e.g. P. obscurus), or absent altogether (e.g. N. nipae). According to Beardsley
(1964), two pairs of ostioles, though poorly defined, are present in the females and
the apterous males of Phenacoleachia australis ; apart from Pseudococcidae and
Phenacoleachiidae the ostioles are not recorded in the females or the males of any
other coccid family.

The dermal structures of the pregenital segments : In Pseudococcus group, the
dorsal and the ventral setae (ads & avs) are arranged in transverse bands, composed
of both fleshy and hair-like setae ; in the remaining groups where only the hair-like
setae are present, they occur in segmental, transverse, irregular rows. The
abdominal pleural setae (aps) are arranged in lateral groups.

A varying number of abdominal pleural pores (app) is usually present on segments
I to VII ; sometimes one or more segments of the abdomen are also with dorsal
pores (adp) (e.g. 0. africanus, Text-fig. 31), or ventral pores (avp) (e.g. S. sacchari,
Text-fig. 27), or both (e.g. D. alazon, Text-fig. 25). A cup-shaped glandular pouch
(gp) is present on each side of segment VIII near its posterior margin, formed by a
cluster of slightly smaller but numerous and tightly packed disc pores ; within the
pouch several setae (gls) originate, namely :

(a) A pair of long and stout " tail setae " (ts), arising from the centre of the
pouch ; these setae are sometimes comparatively short (Nairobia group), or
moderately long (Ceroputo group), or long (the remaining groups).

(b) There are also other shorter setae associated with the long ones : in Piano-
coccus, Pseudococcus and Saccharicoccus groups, a much shorter seta, but con-
spicuously longer than the body setae, is always present ; in Ceroputo and Nairobia
groups, this seta is subequal in length to the other abdominal setae ; in Octococcus
group, two setae of medium but subequal lengths, and a short one are present.

In Ceroputo group, two pairs of glandular pouches and associated setae are
present, one each on segments VII and VIII.

Genital segment and external genitalia

The genital segment is ventrally modified to form a penial sheath (ps) (or " genital
valve ", Berlese, 1893), which represents the fused lateral parts of the 9th sternum
and distally terminates in a rather short projection, the style (st). A small sclero-
tized area occurs dorsally and probably represents the fused 9th tergite and the
10th segment (Theron, 1958), behind which a small anus (an) is situated just
anterior to the style. The style in lateral view, is slightly curved upwards (e.g.
P. citri), or straight with a pointed apex (N. bifrons), or straight with a rounded
apex (e.g. P. citriculus). Anteriorly, the ventro-lateral margin of the penial sheath
is heavily sclerotized, forming a basal ridge (brps). The latter is usually interrupted

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 25

medially and partly overlapped by the membrane of abdominal segment VIII,
forming there a pair of small internal projections (pr) (Giliomee, 1961) ; in Nairobia
group, the basal ridges of the penial sheath are medially continuous and their
projections are absent. Ventrally, at a short distance behind the basal ridges, the
penial sheath is longitudinally slit open to allow the protrusion of the aedeagus
during copulation ; the edges of this slit are sometimes each produced into well
pronounced and heavily sclerotized processes (pro) (e.g. S. sacchari, Text-figs. 27
& 28), or the processes are vestigal (e.g. M. hirsutus, Text-figs. 15 & 16), or entirely
absent (e.g. P. citriculus, Text-figs. 23 & 24). The term " process " was introduced
by Giliomee (1961) for the so-called " opophysis of the genital valve " used by
Berlese (1893), " paramere-like projections " by Makel (1942) and " lobular exten-
sions " by Theron (1958).

The aedeagus (aed) is connected to the ventral wall of the penial sheath, immedi-
ately behind the basal ridges, and supported there by a heavily sclerotized basal
rod (bra). Dorsally, the aedeagus usually forms a curved tube, almost entirely
concealed in the cavity of segment IX when at rest ; in a few species (M. hirsutus,
Text-figs. 15 & 16 and P. citriculus, Text-figs. 23 & 24) the aedeagus is comparatively
long and strongly curved anteriorly, reaching the cavity of segment VII. The
ductus ejaculatorius enters the aedeagus through the internal genital aperture (iga)
and runs posteriorly towards the gonopore at the apex of the penis (Giliomee, 1961).

The dermal structures

The setae of the genital segment (gts) are small and hair-like ; dorsally, 3 setae
are usually present on each side of the median line, near the base of the style, but
sometimes more than 3 are present, e.g. C. insolitus (Text-figs. 35 & 36). Ventrally,
a varying number of setae occur on each side of the penial sheath. Few minute
setal sensilla (pros) may be found on the process of the penial sheath (or its position
if absent). Similar minute sensilla (sts) may also occur dorsally on the style
(N. bifrons, Text-figs. 37 & 38).

APTEROUS MALES

The only available wingless male form was that of S. sacchari (Text-figs. 29 & 30) ;
these are slightly smaller than their winged counterparts. As a result of the
complete absence of their wings and hamulohalterae, the sclerite and ridge degenera-
tion is more pronounced, particularly in the mesothoracic region. The differences
in body structures as compared with the winged males, are as follows :

Head

The separation of the head from the thorax by a neck region is hardly indicated.
The midcranial ridge with its dorsal, lateral and ventral arms is entirely absent.
The ocular ridges (preocular, interocular and postocular) are weakly developed and
reduced to variable degrees. The dorsal and the ventral simple eyes are completely
missing and the antennae are 8-segmented and comparatively short.

26 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Thorax

Prothorax : The pronotal ridge is more or less reduced and sometimes absent
altogether ; the pronotal sclerites and the prosternum are conspicuously reduced.

Mesothorax : The mesotergum is represented only by a weakly sclerotized median
sclerite ; the dorsal subdivisions of the mesothorax with its associated structures
and apodemes are entirely absent. The mesopleuron is greatly reduced and only
an atrophied pleural ridge supporting anteriorly a small episternum, and posteriorly
a vestigial epimeron, persists. The mesosternum is represented by a weakly sclero-
tized basisternum, whose posterior margin is invaginated into a vestigial furca.

Metathorax : The metanotum is indicated by a transverse median narrow sclero-
tized patch. A short pleural ridge, a small episternum and a small epimeron,
represent the metapieuron. The metasternum is largely membranous and the
metasternal apophyses are absent.

Abdomen

The sclerite degeneration is less pronounced in the abdomen ; transverse median
narrow tergites and sternites are present on all the pregenital abdominal segments.
The genital segment and the external genitalia are identical with those of the
macropterous forms.

ERIOCOCCIDAE

In the following account, only the condition of the structures characteristic of
the males of this family, and which are different from those of Pseudococcidae,
will be discussed.

General Characteristics

The studied species were represented by 3 male forms, the macropterous (E.
araucariae, E. orariensis, E. buxi and 0. agavium), the brachypterous (G. salicicola)
and the apterous (P. fraxini) ; macropterous and brachypterous forms of one
species, G. spuria were also available.

Pupation of the eriococcid males (at least of the species obtained by the writer
in the laboratory) takes place within compact, closely felted and rather flattened
waxy puparia ; the latter are split open along the rear edge, to allow emergence
of the adult males.

Appearance. The macropterous forms are moderately robust, if compared with
most of Pseudoccidae ; the brachypterous males are even stouter, with the body
broadest at the third abdominal segment ; the apterous males are spindle-shaped.
The legs are always well developed.

Size. The males of Eriococcidae share about the same range of variation in size
as the Pseudococcidae ; among the species studied the smallest was P. fraxini

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 27

(518-658, av. 6o2[x long), and the largest the winged males of G. spuria (1442-1512,
av. i484[i long).

Colour. The living specimens show a larger variation in colour than in Pseudo-
coccidae ; sometimes they are light yellowish (P. fraxini), light brown (E.
araucariae) or dark red (0. agavium). Some of the material was alcohol-preserved
(Gossyparia spp.), or already mounted on slides in " Berlese fluid " (E. orariensis),
or dry samples (E. buxi), whose normal colour had changed.

Derm Vestiture. Body setae : The hair-like setae are present on the body and
on the appendages ; the fleshy setae may occur on the antennae only (e.g. Gossy-
paria spp.) or on the antennae and the legs (Eriococcus spp.), but are always absent
on the body itself.

Disc pores : These are entirely absent.

Organs of unknown homology : Peculiar organs (X) appearing as small, irregularly
oval or rounded discs, occur dorsally on each side of the midcranial ridge in some
species (E. araucariae and Gossyparia spp.) ; a histological study is required to
elucidate their obscure nature, which however was beyond the scope of the present
work.

MACROPTEROUS MALES

Head

Head capsule : In front view, the head is usually more or less rounded, but
sometimes subtriangular (E. buxi). The ventral preocular depression is hardly
indicated (Eriococcus spp.) or entirely absent (Gossyparia spp.). The arms of the
midcranial ridge are always joined together at the apex of the head forming a
cruciform structure ; the dorsal arm fades away well before reaching the post-
occipital ridge (por). The latter is comparatively strong and well developed, with
the lateral ends distinctly forked. The pre- and the postocular ridges (procr & pocr)
are widely separated, and the interocular ridge is absent ; in E. buxi, the ventral
part of the preocular ridge is reduced. The corneae of the dorsal and the ventral
simple eyes (dse & vse) are usually subequal in diameter. The lateral pair of ocelli
(o) are usually well developed and close to the postocular ridges ; in 0. agavium,
they are reduced to small spots only, and attached to the postocular ridges by
means of short sclerotized arms. The cranial apophysis (ca) is apically bifurcate,
except in E. buxi where it is truncate. The tentorial bridge (tb) is comparatively
stout ; the anterior tentorial arms (ata) are not fused, each joining the cranial
apophysis separately. A small mouth tubercle (mt) sometimes occurs just behind
the preoral ridge (pror) (e.g. 0. agavium, Text-figs. 45 & 46).

The same groups of head setae which occur in Pseudococcidae are also present
in Eriococcidae.

Antennae

The flagellar segments are comparatively short and barrel-shaped, and some-

28 MORPHOLOGY AND TAXONOMY OF ADULT MALES

times the width of the segments, especially of the terminal, becomes equal to their
length (e.g. E. orariensis, Text-fig. 33), giving the antennae a moniliform rather
than a filiform appearance ; usually 10-segmented (9-segmented only in E. buxi).

The position of the antennal bristles (ab) of the terminal segment is slightly
different from that in Pseudococcidae ; sometimes all three bristles are near the
base of the segment (E. buxi, Text-fig. 43, C), or only the dorsal bristle preapical
and the other two at a greater distance from the apex (the remaining spp.). In
E. araucariae (Text-fig. 39, C) two additional, much smaller bristles are also present
on the terminal segment.

The scape (scp) is always with 4 hair-like setae, and the pedicel (pdc) is com-
paratively short. The terminal segment is distinctly pear-shaped ; in some species
(0. agavium) its apex is conspicuously pointed.

Thorax

Prothorax

The prothorax is short. The proepisternum without any ridge-like sclerotiza-
tions. The prosternum (stnl) is triangular, usually long but sometimes rather
short (G. spuria) ; the prosternal ridge (stnr) is well developed.

The prothoracic setae : The medial pronotal, the post-tergital, the lateral pronotal
and the antespiracular dorsal setae are usually absent.

Mesothorax

The scutum (set) is comparatively short (the prescutum 1-2 times as long) and
evenly sclerotized throughout. The scutellum (scl) is transversely rectangular,
with an additional, strong scutellar ridge (sclr), obliquely traversing the scutellum
on each side and dividing it into three well defined triangular areas, the largest of
which is the median. The width of the scutellum is twice or more its length. The
anterior and the posterior ridges of postalare (apar & ppar) are always joined
anteriorly.

The mesopleural ridge (plr2) is continuous above the coxal articulation, and the
basalare (bas) is comparatively stout. The later opleurite (lpl) is large, sometimes
with a circular or oval membranous area (e.g. E. buxi). The trochantin is absent.

The mesothoracic setae : The postmesostigmatal (pms) and the basisternal setae
(stn2s) were found only in E. buxi ; the other groups of the mesothoracic setae
are usually present. Apart from the setae present in Pseudococcidae, few mes-
episternal setae (eps2s) occur in E. buxi, and a minute lateropleurital seta (lpls) on each
side in E. orariensis.

Metathorax

The suspensorial sclerites are usually present, but in 0. agavium they are missing
as a result of the complete absence of the hamulohalterae. The metapostnotal
ridge is always absent. The precoxal ridge of metathorax (pcr3) is either well
developed (e.g. E. araucariae) or rather weak (e.g. 0. agavium).

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 29

The metathoracic setae : The metapleural (mps) and the postmetastigmatal setae
(eps3s) are present in E. buxi only ; the posterior metasternal setae are always
absent.

Wings and Hamulohalterae

The circular sensoria could not be detected on the wings, and are presumably
absent. The hamulohalterae are sometimes absent (0. agavium, Text-figs. 45 & 46) ;
the hooked seta is usually subequal in length to the hamulohaltera (Eriococcus spp.),
but sometimes considerably shorter (G. spuria).

Legs

The fore and the middle legs are usually subequal in length, but sometimes the
latter are slightly shorter (G. spuriae) ; the hind legs are always longest. The
proximal part of the trochanter (tr) is distinctly longer than the distal. The femur
(fern) of the middle leg is always the shortest, and that of the hind leg the longest.
The claw (cl) is relatively broad at the base and abruptly tapering towards a sharply
pointed tip ; the ungual digitules (udgt) are distinctly apicalfy knobbed.

Abdomen

Pregenital segments

The ostioles are always absent. The abdominal setae are arranged in dorsal and
ventral irregular transverse rows, and in pleural clusters ; one dorsal seta on each
side of segment VIII is noticeably longer than the others. A dorso-median, trans-
verse sclerotization sometimes occurs on segment VIII and anteriorly binds its
tergite (E. buxi) ; similar, longitudinal sclerotizations may be also present ventrally
along the outer margins of the sternites of the same segment (e.g. E. orariensis,
Text-figs. 41 & 42). The glandular pouches (gp) are present only on segment VIII,
and their setae (gls) only include a pair of comparatively short tail setae (ts) (the
body more than 5 times as long).

Genital segment and external genitalia

The structure of the genital segment and the external genitalia is of great
taxonomic significance ; it appears to be the most reliable feature distinguishing
all forms of Eriococcidae males from those of Pseudococcidae. The basal ridges of
the penial sheath are medially continuous and modified into a complete, well
sclerotized ring. The basal part of the penial sheath is rather long, intimately
fused with the 9th tergite + the 10th segment, forming together a compact,
cylindrical genital capsule (gc). The anal opening (an) is comparatively large, and
the style (st) is usually short but sometimes rather long (e.g. E. orariensis, Text-
figs. 41 & 42) ; the style in lateral view, is either sinuate (e.g. E. araucariae) or
straight (e.g. 0. agavium). Ventrally, the slit-like opening of the penial sheath is
removed far posteriorly, and the aedeagus (aed) arises at a comparatively large
distance from the basal ring of the genital capsule. The aedeagus usually consists
of a basal rod (bra) and a sclerotized tube, ventrally accommodated within the

3 o MORPHOLOGY AND TAXONOMY OF ADULT MALES

style ; in E. araucariae the structure of the aedeagus is rather complex, with
secondary sclerotizations, as will be discussed in the individual description.

The dermal structures

The dorsal setae of the genital segment (gts) always consist of a pair of relatively
long and stout setae on each side of the median line, usually near the base of the
style. A variable number of smaller setae also occurs ventrally on each side of the
genital capsule.

It appears that the structure of the genital segment shows closer relationship of
Eriococcidae with Diaspididae than with Pseudococcidae.

BRACHYPTEROUS MALES

In this form, a considerable sclerite degeneration in the meso- and the metathoracic
regions is apparent, as a result of the considerable reduction of the fore wings and
the hamulohalterae.

Head

A comparatively large mouth tubercle (mt) is always present behind the preoral
ridge.

Thorax

All the meso- and the metathoracic structures become greatly reduced. The
scutellum is trapezoidal in dorsal view, with the scuto-scutellar suture weak or
incompletely developed ; the scutellar ridge is entirely absent. The marginal ridge
of the basisternum is slender and partly absent. The suspensorial sclerite is absent,
and the metapleural ridge is dorsally reduced. The hamulohalterae if present, are
atrophied and without setae or ridges.

Abdomen

The glandular pouch and its associated setae are greatly reduced, the total length
of the body being 12 times or more as long as the setae. The genital segment is
similar to that of the macropterous males.

APTEROUS MALES

The form is only represented by the males of Pseudochermes fraxini. The sclerite
degeneration is most pronounced.

The body is spindle-shaped in the general appearance. The head is broadly fused
with the thorax ; the simple eyes are absent and the ocelli represented by small
spots or tubercles. Almost all the thoracic features have disappeared, except for a
short and vestigial pleural ridge on each segment. The legs are comparatively short
and stout ; the trochanter with a long apical seta, but apparently without any
sensilla. The glandular pouches are entirely absent ; the genital capsule is small
and the anal opening minute.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 31

DISCUSSION

This study of the male representatives of Pseudococcidae and Eriococcidae has
revealed many interesting facts regarding their taxonomic status. Moreover, the
material of Pseudoccidae, being fairly representative (20 spp. were considered),
allowed for certain suggestions as to the classification within the family. The
material of Eriococcidae, although inadequate for similar suggestions (7 spp. only),
was sufficient for a general characterization of the males of the family, and for
suggestions regarding the various degrees of morphological relationships between
the studied species.

These conclusions were assessed by taking into consideration as many characters
as possible, as listed in table I. The table includes all the species here studied and
3 others adequately described for the purpose of comparison by Giliomee (1961).
The large number of species studied by Beardsley (i960, 62, 63, 64 and 65) could
not be included in this table since his descriptions contain comparatively few of the
characters here employed. The assessment of the relationships between the studied
species was carried out by using the quantitative evaluation, method adopted by
Ghauri (1962) and followed by Giliomee (1967). In this method all the characters
were regarded as being of equal importance, and those shared by any two species
were counted, and the calculated numbers (shown in table II in the form of a
matrix) were taken as indices of the degree of affinity. This method was found
most practical in the circumstances, although not the most satisfactory from the
statistical point of view. The methods recently discussed by Sokal and Sneath
(1963), in which a correlation coefficient is obtained for each pair of species, is more
accurate, but involves the use of an electronic computer for calculation and the
development of a rather elaborate computer programme.*

The discussion, therefore, will include the following aspects :

(a) The taxonomic status of Pseudococcidae and Eriococcidae.

(b) The taxonomic significance of the systematically important characters, and
the levels at which they could be used.

(c) The classification and interrelationships within each of these groups.

(d) The relationships of the Pseudococcidae and Eriococcidae with other
families of Coccoidea.

(A) Status of Pseudococcidae and Eriococcidae

On examining table I, it was immediately apparent that the pseudococcid and
the eriococcid males exhibit a number of structural differences, comparable with
those separating other families (e.g. Pseudoccidae and Coccidae), thus justifying the
recognition of the two groups as distinct families. This conclusion is in accordance
with Ferris's (1937) concepts of classification, which were also adopted by Hoy
(1962-63).

* This programme has only recently been developed in the Department of Zoology and Applied
Entomology by Dr. G. Murdie, taking the advantage of the computing facilities available in the Imperial
College. It was applied too late to include in the main part of the thesis, but the results of the two
methods are discussed under Analysis.

32

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MORPHOLOGY AND TAXONOMY OF ADULT MALES

The characters distinctly separating the macropterous males of Pseudococcidae
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Puparia

Table A

Pseudococcidae

Fluffy, loosely felted, with no
definite posterior opening.

Postoccipital ridge : Weak, slender, U- or V-shaped.

Terminal antennal
segments :

Proepisternum :

Elongated ; more or less
cylindrical.

With the dorsal margin (and
sometimes also the ventral
margin) heavily sclerotized and
ridge-like.

(5)

Scutellum, in dorsal
view :

Pentagonal.

(6)

Scutellar ridge :

Absent.

(7)

Anterior and
posterior postalar
ridges :

Well separated.

(8)

Mesopleural ridge :

Interrupted above the coxal
articulation.

(9)

Trochanter :

With the basal part longer
than the distal one.

(IO)

Ungual digitules :

With acute tip.

(«)

Penial sheath :

Basal part short, not fused

(12) Aedeagus

with the 9th tergite + the
10th segment ; anterior
margin of the slit-like opening
close to the basal ridge.

Arises ventrally just behind
the basal ridge of the penial
sheath.

Eriococcidae

Compact, closely felted,
flattened, permanently split
along the posterior edge.

Strongly developed, laterally
forked.

Barrel-shaped ; the apical
segment distinctly pear-
shaped.

Without any ridge-like
sclerotization.

Transverse, rectangular.

Present.
Anteriorly joined.

Continuous above the coxal
articulation.

With the basal part shorter
than the distal one.

Apically knobbed.

Basal part long, fused with
the 9th tergite, the 10th
segment forming together
cylindrical basal genital
capsule ; anterior margin of
the slit-like opening removed
far posteriorly.

Removed backwards and
arises at a comparatively
large distance from the basal
ring of the genital capsule.

The two families can be also separated by a number of other less well defined
characters (listed in table B), the taxonomic significance of which may be confirmed
as more information becomes available :

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

53

(i)

(2)

(3)

(4)

(5)

(6)

Tentorial bridge :
Three large bristles
of terminal antennal
segment :
Claws :

Dorsal setae of
abdominal segment
VIII :

Glandular pouch of
abdominal segment
VIII :

Dorsal setae of the
genital segment :

Table B

Pseudoccidae

Comparatively thin.

All preapical, or the dorsal one

removed a greater distance

from the apex.

Rather long, slender, uniformly

tapering to a pointed end.

As long as other abdominal
setae.

With a pair of long setae and

one or more much shorter

ones.

Three or more setae, subequal

in length to the abdominal

setae, occur on each side of

the median line.

Eriococcidae

Rather stout.

All near the base of the

segment, or only the dorsal

one preapical.

Rather short and stout, broad

at base and abruptly tapering

to a pointed end.

Include one conspicuously

long seta on each side of the

segment.

With only one pair of setae.

Two setae, much longer than
the abdominal setae, are
always present on each side.

There is also a number of supporting characters (listed in table C) whose presence
or absence obtains in one of the two families, but whose alternative conditions
occur in the other.

Characters

(I)

Disc pores :

(2)

Dorsal arm of the

midcranial ridge :

(3)

Pre. and post-ocular

ridges :

(4)

Cranial apophysis :

(5)

Anterior tentorial

arms :

(6)

Relative lengths of

prescutum and

scutum :

(7)

Median, longitudinal,

membranous area of

scutum :

(8)

Trochantin :

(g) Metapostnotal ridge :
(io) Ostioles :

Table C

Pseudococcidae

Present, except in Nairobia

group.

Detached anteriorly from other

arms, except in Ceroputo and

Nairobia groups.

Joined by interocular ridge,

except in Ceroputo and

Nairobia groups.

Apically truncate.

Anteriorly fused before

meeting cranial apophysis,

except in Ceroputo and

Nairobia groups.

Prescutum shorter than scutum,

except in Ceroputo and

Nairobia groups.

Present, except in Ceroputo

and Nairobia groups.

Present, except in Nairobia

group.

Present, except in Nairobia

group.

Present or absent.

Eriococcidae
Always absent.

Joined with the lateral and
the ventral arms at the apex
of the head.

Widely separated ; inter-
ocular ridge absent.

Apically bifurcate, except in

E. buxi.

Well separated throughout.

Prescutum 1-2 times as long
as scutum.

Absent.

Absent.
Absent.
Absent.

54

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Apterous Males

This comparison between the apterous males of Pseudoccidae and Eriococcidae
must be regarded as very provisional since it is based only on one representative of
each family (S. sacchari and P. fraxini). They appear to indicate that some of
the characters separating the macropterous males of the two families also differen-
tiate the apterous forms ; these are the characters related to the conditions of the
trochanter, claw, ungual digitules, penial sheath and aedeagus. The other
characters separating the wingless males are given in the following table :

Characters

(i) Appearance in dorso-
ventral view :

(2) Sclerite degeneration :

(3) Disc pores :

(4) Length of body /length
of hind legs :

(5) Length/width of hind
femur :

(6) Length of tibia/length
of tarsus :

(7) Ostioles :

(8) Pleural setae of
abdominal segments
VI, VII and VIII :

(9) Glandular pouches of
segment VIII :

Pseudococcidae
Oblong.

Comparatively less pronounced.
Present.

Body less than 2-5 times as long.

Length more than 35 times
as width.

Tibia more than 1-5 times the
length of tarsus.

Present.

Include one comparatively
long seta (more than twice as
long as other abdominal setae).

Present.

Eriococcidae
Spindle-shaped .

Well pronounced.
Absent.

Body more than 2-8 times as
long.

Length twice the width, or
less.

Tibia less than 1-5 times the
length of tarsus.

Absent.

Pleural and other abdominal
setae subequal in length.

Absent.

The differences between the two families will be compared with the differences
separating other families of Coccoidea later (p. 74).

(B) Taxonomic Significance of the Characters

The taxonomically important characters of both Pseudococcidae and Eriococcidae
(winged forms) are here discussed together, but the limited number of the available
species of Eriococcidae make the significance and levels on which these characters
may operate within this family more tentative than those of Pseudococcidae.
These characters are also listed in tables III, V and VI (pp. 58, 69, 72).

General appearance

Although the appearance of the males does not seem to be strictly characteristic
at any level, most Pseudoccidae are narrow and slender, whereas Eriococcidae are
rather stout and robust ; the hairy appearance of the species will be discussed later.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 55

Size

The size generally varies a great deal, even within the species themselves.
Giliomee (1967), for example, showed that the host plant and the locality had a
considerable influence on the size of Parthenolecanium corni (Bouche). No such
host-determined differences have been found in P. citri, for which material from
several host plants and localities was available. Some genera or species within
certain genera are distinctly smaller than others (e.g. N. nipae is smaller than
N. vastator). The size, therefore, seems to be of some significance at the generic
and specific levels only.

Chaetotaxy

The presence or absence of fleshy setae on the various parts of the body was
found of considerable importance, operating from the level of the groups of genera
downwards to the species. The hairy appearance, which is generally due to the
presence of the fleshy setae on the body itself, separates the Pseudococcus group.
The complete absence of the fleshy setae also separates groups (Nairobia) or genera
(Centrococcus). In N. vastator the fleshy setae are absent on the legs, but in N.
nipae present.

The dermal pores

With the exception of the Nairobia group, the presence of the disc pores is a
distinct character separating the two families studied ; they are always present in
Pseudococcidae and absent in Eriococcidae. Within the former family their
number and distribution also presents a wide range of variation, and can be utilized
at most intrafamily levels.

Head

The general shape of the head appears to be of certain taxonomic significance at
family level, but some minor details also separate some genera. In Pseudococcidae
the head is triangular in dorsal view, whereas in most Eriococcidae it is more or
less rounded ; in the genus Saccharicoccus, the head is dorso-ventrally flattened
and the ventral preocular depression absent. The condition of the midcranial
ridge operates at many levels ; family Eriococcidae and some pseudococcid groups
(Ceroputo and Nairobia) are separated by having all the arms of the midcranial
ridge joined together. The degree of development of the dorsal arm separates
genera. The weak development of the lateral arms separates groups of genera
(Ceroputo and Nairobia), genera (Trionymus) and species (E. buxi). The nature
of the postoccipital ridge distinguishes the two studied families. The relationship
between this ridge and the preocular ridge separates genera and species of Pseudo-
coccoidae. The complete separation of the pre- and postocular ridges separates
the family Eriococcidae and two groups of Pseudococcidae (Ceroputo and Nairobia).
The dorsal reduction of the postocular ridge excludes the Saccharicoccus group.
The relative size of the simple eyes is a useful character separating groups of genera,
genera and species. The vestigial state of the lateral ocelli appear to be character-

56 MORPHOLOGY AND TAXONOMY OF ADULT MALES

istic of one group of Pseudococcidae (Nairobia) and one genus of Eriococcidae
(Ovaticoccus). The apically truncate cranial apophysis separates Pseudococcidae
and one species of Eriococcidae (E. buxi). The nature of the tentorial bridge is
slender in Pseudococcidae and comparatively stout in Eriococcidae. The presence
of a mouth tubercle separates the genus Ovaticoccus.

The head setae provide taxonomically important characters operating on supra-
generic, generic and specific levels ; the presence of fleshy setae for example
separates Pseudococcus group ; the presence of dorsal ocular setae separates genus
Dysmicoccus ; the presence of setae between the ventral eyes excludes E. buxi.

The number and distribution of the head pores also separates groups of genera,
genera and species of Pseudococcidae.

The antennae afford a number of systematically important characters, at all
taxonomic levels from the family down to the species. The shape of the terminal
antennal segments separates the two families ; the relative length of the antennae
and the individual segments, and the presence and distribution of the different
types of setae operates on most intrafamily levels, and the following have been
utilized for this purpose : the ratio of the length of antennae to the length of the
body and to the length of the hind legs ; the ratio of the length of the 3rd to the
terminal segment ; the ratio of width to length of the 3rd and the preterminal
segments ; the presence or absence of the fleshy, the hair-like and the apically
knobbed sensory setae ; the number of setae on the scape ; the number, nature,
relative lengths and position of the antennal bristles of the terminal segments.

Thorax

Prothorax

The absence of ridge-like sclerotization along the margins of the proepisternum
separates the family Eriococcidae ; within Pseudococcidae the presence of this
sclerotization along both dorsal and ventral margins of the propeisternum separates
Ceroputo group of genera. The absence of the prosternal ridge excludes the
Saccharicoccus group of genera.

Mesothorax

The shape of the prescutum separates groups of genera ; in the Ceroputo and
Nairobia groups, the prescutum is triangular in dorsal view. The medio-posterior
continuation of the prescutal ridges isolates the Ceroputo group. The shape and
structure of the scutellum separates the two families. The following ratios were
utilized, for their significance at family and lower levels : length to width of
prescutum ; length of prescutum to length of scutum ; length to width of scutellum ;
length of scutellum to length of scutum.

The conditions of the postalare and the mesopleural ridges are important
characters separating the two families. The weak development of the basalare
separates Eriococcidae and some groups of Pseudococcidae (Octococcus, Ceroputo
and Nairobia). The presence of a large lateropleurite also separates Eriococcidae

OF PSEUDOCOCCIDAE & ERIOCOCCIDA E 57

and some groups of Pseudococcidae (Saccharicoccus, Ceroputo and Nairobia). The
incomplete development of the marginal ridge of the basisternum eliminates
Saccharicoccus group.

Metathorax

The absence of the suspensorial sclerites is accompanied by the absence of the
hamulohalterae, and separates the genus Ovaticoccus. The absence of the meta-
postnotal ridge is an important feature of all Eriococcidae and also the Nairobia
group of genera of Pseudococcidae.

The number and distribution of the thoracic setae and pores provide useful
characters at the generic and the specific levels.

The legs

The legs, like the antennae, afford several characters operating at all levels.
The two families can be separated by different conditions of the following structures :
(A) the proportion between the basal and the distal parts of the trochanter ; (B)
the shape of the claw ; and (C) the nature of the ungual digitules. The relative
length of legs separates groups of genera ; e.g. the Saccharicoccus group has com-
paratively short legs (the hind pair being shorter than half the length of the body).
The absence of the tarsal digitules separates groups of genera (Ceroputo and
Nairobia). The following characters were also found taxonomically important at
the generic and specific levels : the presence of a long apical seta on the trochanter ;
the ratio of width to length of the hind femora ; the ratio of the length of the femur
to the length of the tibia in the fore leg.

The abdomen

The complete absence of the ostioles is characteristic of Eriococcidae, but among
Pseudococcidae the degree of their development, or their absence separates groups
of genera, genera and species. The number and nature of certain abdominal setae
separates the different categories : the presence of a long dorsal seta on each side
of segment VIII excludes Eriococcidae, and the entire absence of the dorsal setae
on this segment characterizes the Nairobia group. The nature of the pleural setae
of segment VIII, the number and distribution of the abdominal pores (within the
Pseudococcidae) separates groups of genera, genera and species. Narrow areas of
heavy sclerotization of abdominal segment VIII separate some species of Erio-
coccidae (E. orariensis and E. buxi).

The presence of two pairs of glandular pouches is a conspicuous character separating
the Ceroputo group of genera. The number and relative length of the tail-forming
setae of the glandular pouch operates at the levels of family and groups of genera.

The structure of the genital segment is an important character separating the
two studied families. The number and size of the dorsal setae of the genital
segment also separate families. The following characters and ratios were also
found of some taxonomic significance in separating groups of genera, genera and
species : the structure of the basal ridge of the penial sheath ; the relative length

58 MORPHOLOGY AND TAXONOMY OF ADULT MALES

to width of the penial sheath or the genital capsule ; the length of the genital
segment to the total length of the body ; the shape of the style ; the presence or
absence of the process of the penial sheath.

Classification of Pseudococcidae

It is apparent that the material was of necessity, rather limited and unevenly
representative. Nevertheless, it was possible to obtain some interesting results
concerning suprageneric classification, although this division is purely tentative
and some of the proposed groups of genera may be subject to further subdivision,
expansion or any other modification that future findings may demand. Careful
examination of table I showed that the studied species of Pseudococcidae could be
divided, on the basis of the different conditions of 58 characters, into 6 distinct
groups of genera. Some of these groups contain several genera (each represented
by one or more species), and the characters shared within each of these groups were
taken as an indication of the suprageneric level of significance of these characters.
In other instances a single genus (even if represented by one species only) was
found by comparison to be distinct enough to constitute the nucleus of a separate
suprageneric taxon. The six groups of genera, therefore, are not evenly represented ;
one group contains six genera, two groups two genera each, and the remaining three
groups consist of only one genus each. In addition, the published descriptions of
Puto yuccae and Rhizoecus falcifer, though incomplete for detailed comparison with
the present data, would certainly suggest two other groups of genera. The actual
status and rank of all the recognized groups will be discussed later.

The 58 selected characters are listed in table III, which also shows their distribu-
tion among the groups. It will be noticed that alternative conditions of a few
characters sometimes occur in the same group (thus only separating genera), but
which separate the other groups. Bold characters indicate their exclusive
conditions for a particular group of genera ; the total number of exclusive
characters for each group is given at the end of the table.

Table III.
Characters separating groups of genera of Pseudococcidae

Groups

Piano-

Pseudo-

Sacchari- Octo-

Cero-

Nairo

Characters

coccus

coccus

coccus coccus

puto

bia

(1)

(2)

(3) (4)

(5)

(6)

(1) Absent.

(2) Present.

2

2

2 2

2

1

Disc pores

Fleshy setae : (1) Absent on the body
itself. (2) Present on the head,
thorax and abdomen, giving the body
a distinctive hairy appearance.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 59

HEAD : (!) (2) (3) (4) (5) (6)

Ventral preocular depression : ( 1 )

Entirely absent. (2) Well marked. 2 2 1 2 2 2

Appearance in front view : (1) Sub-
triangular or rounded. (2) Dorso-
ventrally flattened. 1 1 2 1 1 1

Dorsal arm of midcranial ridge : (1)

Anteriorly detached from lateral and

ventral arms. (2) All arms of the

midcranial ridge joined. 1 1 1 1 2 2

Lateral arms of midcranial ridge : (1)

Represented by weak sclerotization.

(2) Well developed. 1&22 2 2 1 1

Postoccipital ridge : (1) U-shaped.

(2) V-shaped. 1 1 2 1 1 1

Anterior development of postoccipital

ridge : (1) Not reaching preocular

ridges. (2) Confluent with preocular

ridges. 1 & 2 1 & 2 1 1 2 2

Pre. and postocular ridges : (1) Widely

separated. (2) Joined by means of

the interocular ridge. 2 2 2 2 1 1

Postocular ridge : (1) Dorsally

reduced. (2) Well developed. 2 2 1 2 2 2

Lateral ocelli : (1) Vestigial. (2)

Large and well developed. 2 2 2 2 2 1

Anterior tentorial arms : (1) Separated.

(2) Fused before merging into the

cranial apophysis. 22221 1

Head setae on each side :

Dorsal head setae : (1) Usually 7

setae or less. (2) More than 7 setae. 2 2 2 1 2 2

Ventral ocular setae : (1) Absent.

(2) Present. 1 2 1 1 1 1

Ventral head setae in a transverse row
or band anterior to the ventral eyes :

(1) Usually 5 setae or less. (2) More

than 5 setae. 222122

Genal setae : (1)7 setae or less on

each gena. (2) More than 7. 1 2 1 1 1 1

Ventral head pores : (1) Absent.

(2) Present. 1 1 1 1 2 1

Antennae : (1) Comparatively short,

the body 16 or more times as long.

(2) Comparatively long, the body

1-5 or less times as long. 1 1 1 2 2 1

6o MORPHOLOGY AND TAXONOMY OF ADULT MALES

, # . , , , . M < 2 ) (3) (4) (5) (6)

Length oj antenna in relation to length

of hind leg : (i) Subequal or antenna

shorter. (2) Antenna longer. 1 1 1 2 2 1

Relative length of yd and terminal
antennal segments : (1) Segment III
less than 1 -5 times as long ; some-
times both subequal. (2) Segment
III usually more than 1-5 times as

the terminal. 1 1 1 1 2

Width of segment III related to its
length : (1) Length usually 5-3 times
the width or less. (2) Length more

than 5-3 times the width. 1 1 1 1 2 1

Width of preterminal antennal segment
to its length : (1) Length usually less
than 3-3 times the width. (2) Length

33 times the width, or more. 1&21 1 2 2 2

Antennal setae : (1) Comparatively
short ; less than twice as long as
body setae. (2) Long ; twice as

long as body setae or more. 1 & 2 1 1 1 2 1

Capitate, apically knobbed sensory

setae : (1) Absent. (2) Present. 2 2 2 2 1 2

If present : (1) Only on segment X.
(2) Few on segments III to X, but
other setae more numerous. (3) On
segments III to X forming the

majority of the antennal setae. 1 1 1 2 NC 3

Number of hair-like setae on scape :
(1)4 setae almost always present.
(2) More than 4 setae present. 1 & 2 2 2 1 1 1

THORAX :
PROTHORAX :

Proepisternum : (1) Dorsal margin

ridge-like. (2) Dorsal and ventral

margins ridge-like. 1 1 1 1 2 1

Prosternal ridge : (1) Its position

only marked by heavy sclerotization.

(2) Distinct. 2 2 1 2 2 2

Medial pronotal setae : (1) Usually

absent. (2) Present. 2 2 2 1 1 1

Post-tergital setae : (1) Usually

absent. (2) Present. 1&22 2 1 1 1

A ntespiracular dorsal setae : ( 1 )

Absent. (2) Present. 2 2 2 1 1 2

Antespiracular ventral setae : (1)1

seta on each side. (2) 2 setae. 1 1 2 1 1 1

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 61

(I) (2) (3) (4) (5) (6)

Prosternal setae : (1)3 setae or less
on each side. (2) More than 3 seta
present. 1 2 1 1 1 1

Antespiracular dorsal pores : (1)

Absent. (2) Present. 2 2 2 1 1 NC

MESOTHORAX :

Prescntal ridges : (1) Well separated.

(2) Posteriorly joined. 1 1 1 1 2 1

Width to length of presculum : (1)

Width less than 1-3 times the length.

(2) Width 1 3 times the length, or

less. 1&22 2 2 1 1

Scutum : Median longitudinal narrow

membranous area : (1) Absent.

(2) Present. 2 2 2 2 1 1

Relative length of presculum and

scutum : (1) Prescutum shorter.

(2) Prescutum longer. 1 1 1 1 2 2

Relative length of scutellum and

scutum : (1) Scutellum shorter than

scutum. (2) Scutellum longer. 1 1 1 1 2 2

Basalare : (1) Comparatively weak

and slender. (2) Rather stout. 222111

Lateropleurite : (1) Comparatively

narrow. (2) Large. 1 1 2 1 2 2

Trochantin : (1) Absent. (2) Present. 2 2 2 2 2 1

Marginal ridge of basisternum :

(1) Absent. (2) Present. 2 2 1 2 2 2
1'urca : (1) Comparatively small.

(2) Well developed. 2 2 1 2 2 2
Postmesostigmatal setae : (1) Absent.

(2) Present. 2 2 2 2 2 1

METATHORAX :

Metapostnotal ridge : (1) Absent.

(2) Present. 2 2 2 2 2 1

Alar setae : (1) Absent. (2) Present. 2 2 2 2 2 1

LEGS :

Hind legs : (1) Short ; the body

more than 2-2 times as long.

(2) Long ; the body less than 22

times as long. 2 2 1 2 2 2

Femur : (1) Front femur shortest.

(2) Middle femur shortest. 1&21 1 2 2 1

Tarsal digitules : (1) Absent.

(2) Present. 2 2 2 2 1 1

62 MORPHOLOGY AND TAXONOMY OF ADULT MALES

(i) (2) (3) (4) (5) (6)
ABDOMEN :

Ostioles : (i) Absent. (2) Present. 1 & 2 2 2 1 2 1

Dorsal setae of segment VIII :

(1) Absent. (2) Present. 2 2 2 2 2 1

Pleural setae of segment VIII :

(1) All of subequal length. (2)

Include one slightly longer seta. 1&22 2 1 1 1

If disc pores present : (1) Only on

segment I. (2) On segment I and

other abdominal segments. 2 2 2 1 2 NC

Glandular pouches and their setae :

(1) Only on segment VIII. (2) One
set on each side of segments VII and

VIII. 1 1 1 1 2 1

Setae of the glandular pouch of

segment VIII : (1) Include a pair of

comparatively long tail setae and one

much shorter seta. (2) Include a

pair of tail setae, two of medium,

unequal lengths and a short one. 1 1 1 2 1 1

Length of tail setae related to total

length of body : (1) Short ; body

usually more than 5 times as long.

(2) Long ; body usually 5 times as

long, or less. 2 2 2 2 1 1

Basal ridge of the penial sheath :

(1) Medially interrupted and with a

small projection. (2) Medially

continuous ; projection absent. 1 1 1 1 1 2

Total number of exclusive characters 048589

In discussing the groups of genera, an attempt was made to assess their primitive
or specialized status, on the basis of the number of shared specialized and primitive
characters, but this was found to be rather difficult. Theron (1958) discussed the
matter with reference to the conditions of the more primitive Sternorrhynchan
Homoptera and considered 3 main conditions as distinct specializations in
Coccoidea : (1) sclerite degeneration, and consequently (2) the development of
secondary ridges for reinforcing the desclerotized parts, (3) the absence of any of
the primary ridges, or the reduction of other generalized structures.

When the groups of genera of the studied Pseudococcidae were compared with
each other, the different structures showed varying degrees of development, and
complying with the principles already set out by Theron (I.e.) and Giliomee (1967),
the following conditions of the various characters were regarded as specialized :

(a) The presence of : ocular ridges (preocular, postocular and interocular) ;
ridge-like sclerotization of the proepisternum ; short and transverse scutellum ;
ostioles ; basal ridge replacing the basal ridge of the penial sheath.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 63

The presence of the lateral arms of the midcranial ridge is considered as a
secondary specialization since they are absent in primitive families (Margarodidae),
but reappear in most specialized ones such as Coccidae and Diaspididae.

(b) The absence of : various sclerites (as an expression of sclerite degeneration) ;
disc pores ; dorsal and ventral arms of midcranial ridge ; lateral ocelli ; pronotal
ridges ; cervical sclerites ; metapleural ridges ; hamulohalterae ; tarsal digitules ;
glandular pouches and their setae.

On the other hand, the following conditions were considered as being primitive :

(a) The strong development of : postoccipital ridge : distinct neck region ;
basalare.

(b) The presence of : more than 2 pairs of simple eyes ; separated anterior
tentorial arms ; prosternal or metasternal apophyses ; large lateropleurites ;
more than one hamulohalteral seta ; preapical denticle of the claw.

The presence of a small, narrow sclerite identified as the trochantin (first mentioned
by Giliomee, 1961, and noticed but not identified as such by Makel, 1942), and
found in all Pseudococcidae here studied except N. bifrons, should be considered
as a primitive condition. It was not discussed at all by Theron, but Weber (1928)
and Roberti (1946) found it in Aphis.

(c) The absence of : prosternal ridge ; marginal ridge of the basisternum ;
metapostnotal ridge.

a. Groups of genera :

The proposed groups of Pseudococcidae are :

I. The Nairobia group, represented by the genus Nairobia only, is the most
specialized among the Pseudococcidae here studied. The exclusive characters of
this group are :

(1) The disc pores entirely absent.

(2) The lateral ocelli vestigial.

(3) Antennal segments III to X with a dominant number of capitate, apically
knobbed sensory setae.

(4) The trochantin absent.

(5) The metapostnotal ridge absent.

(6) The basal ridges of the penial sheath ventro-medially continuous and their
projections absent.

(7) The absence of the postmesostigmatal and the alar setae, and the abdominal
dorsal setae of segment VIII.

The group exhibits a large number of specializations which include four of the
above-listed exclusive characters (1, 2, 6 and 7) as well as : (5) the reduction of
the prosternal plates, (6) the weak development of the basalare, (7) the absence of the
tarsal digitules, and (8) the reduction of the setae of the glandular pouches.

64 MORPHOLOGY AND TAXONOMY OF ADULT MALES

It also features a few primitive conditions including : (i) the absence of the
metapostnotal ridge (exclusive character No. 5), (2) the weak development of the
lateral arms of the midcranial ridge, (3) the absence of the interocular ridges, (4)
separated anterior tentorial arms, and (5) the absence of ostioles.

This group shares with Eriococcidae the absence of three characters typical of
Pseudococcidae (i.e. the absence of : disc pores, metapostnotal ridge and trochantin),
and probably represents a link between these two families.

As far as the existing literature is concerned, none of the previously described
genera could be assigned to this group.

II. The Ceroputo group, represented by two closely related genera, Ceroputo
and Centrococcus, which share between them a large number of characters (103) ;
this group is less specialized than Nairobia and shows the following exclusive
conditions :

(1) The ventral head pores present.

(2) The 3rd antennal segment ii or more times as long as the terminal

segment.

(3) The 3rd antennal segment more than 5-3 times as long as wide.

(4) The capitate, apically knobbed sensory setae of the antennae entirely

absent.

(5) The dorsal and the ventral margins of the proepisternum heavily sclero-

tized and ridge-like.

(6) The prescutal ridges posteriorly continuous.

(7) The glandular pouches and their setae present on each side of abdominal

segments VII and VIII.

The specializations of the group include two of the above-listed exclusive
characters (5 & 6) and : (3) the presence of a ridge-like projection or sclerotized arm
extending anteriorly from the postocular ridge, (4) the reduction of the scutum,
(5) the weak development of the basalare, (6) the absence of the tarsal digitules,
and (7) the short setae of the glandular pouches.

The primitive features are two of the exclusive characters (1 & 7) and : (3) the
weak development of the lateral arms of the midcranial ridge, and (4) the separation
of the anterior tentorial arms.

From the available generalized descriptions of the males of Phenacoccus aceris
(Sign.) (Sulc, 1943), Heterococcus graminicola Morrison (Morrison, 1945), Phena-
coccus piceae Loew (Jancke, 1955) and Phenacoccus gossypii Townsend & Ckll.
(Beardsley, i960, 1962), it appears that the genera Phenacoccus and Heterococcus
also belong to this group. The descriptions indicate that all these species have
glandular pouches and associated setae on each side of abdominal segments VII
and VIII, which is the most distinctive exclusive character of the Ceroputo group
of genera.

III. The Octococcus group is fairly specialized and represented by one genus
only, Octococcus. Its only representative species (0. africanus) exhibits a number

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 65

of exclusive features justifying its isolation as a separate group. These exclusive
features concern chaetotaxy and the distribution of disc pores, namely : (1) dorsal
head setae 7 or less on each side ; (2) transverse band of ventral head setae include
5 or less setae on each side ; (3) few capitate, apically knobbed sensory setae present
on antennal segments III to X ; (4) abdominal disc pores present on segments I
and VIII only ; and (5) setae of the glandular pouch include a pair of comparatively
long tail-forming setae, two setae of medium, unequal lengths and a short one.

The specializations of the group are : (1) the absence of disc pores on the head
and on abdominal segments II to VII ; (2) the strong development of the lateral
arms of the midcranial ridge ; and (3) the presence of interocular ridges.

The primitive features are : (1) a large scutum, (2) strong basalare, and (3) long
setae of the glandular pouches.

None of the previously studied species could be included in this group.

IV. The Saccharicoccus group, which also comprises only one genus, Sacchari-
coccus. The species studied (S. sacchari) represents a rather primitive group with
the following exclusive characters :

(1) The head and body dorso-ventrally flattened.

(2) The ventral preocular depression entirely absent.

(3) The postoccipital ridge V-shaped.

(4) The postocular ridge dorsally reduced.

(5) The prosternal ridge absent.

(6) The antespiracular ventral setae more than one on each side.

(7) The marginal ridge of the basisternum absent medially.

(8) The furca reduced.

(9) The legs comparatively short ; the body more than 2-2 times as long as

the hind legs.

The specializations of the group are : (1) the strong development of the lateral
arms of the midcranial ridge ; (2) the presence of interocular ridges ; and (3) the
reduction of the furca.

The comparatively large number of primitive features include three exclusive
characters (4, 5, 7) and : (4) large scutum, (5) strong basalare, (6) long setae of
glandular pouches.

No other species could be assigned to the Saccharicoccus group.

V. The Pseudococcus group : Two genera, Pseudococcus and Dysmicoccus
represent this group and share 93 characters. The group is one of the two most
primitive among the material studied, and includes species with most characters in
the generalized pseudococcid condition. It shows only one exclusive, but rather
obvious character, i.e. the presence of fleshy setae on any part of the body itself
(head, thorax and abdomen), usually giving the species a rather hairy appearance.
The specializations of this group are : (1) lateral arms of midcranial ridge strongly
developed, (2) interocular ridge present.

66 MORPHOLOGY AND TAXONOMY OF ADULT MALES

The primitive characters are : (i) scutum large, (2) basalare strong, (3) setae of
the glandular pouches long.

The descriptions of Pseudococcus antricolens Ferris ; P. dorsispinosus Beard. ;
P. floriger Ferris ; P. lycopodii Beard. ; P. montanus Ehrhorn ; P. straussiae
Ehrhorn ; P. comstocki (Kuwana) ; P. peleae Beard. ; Dysmicoccus boninensis
(Kuwana) ; D. brevipes (Ckll.), D. neobrevipes Beard, by Beardsley (i960, 1962,
1965) and those of Pseudococcus fragilis Brain ; P. adonidum (L.) ; and "P.
maritimus (Ehrhorn) type A & B " by Giliomee (1961) show that all these species
have fleshy setae on one or more parts of the body itself, thus confirming the validity
of the exclusive character in the other species of the two genera Pseudococcus and
Dysmicoccus. According to Beardsley (I.e.), the males of Pedronia acanthocauda
Beard., P. cibotii Beard., P. crypta Beard., P. hawaiiensis Ferris, Clavicoccus tribulus
Ferris, Phyloccus oahuensis (Ehrhorn), Laminicoccus giffardi (Ehrhorn), Palmicola
palmarum (Ehrhorn), and Nesococcus pipturi (Ehrhorn) also have fleshy setae on
the main parts of the body. The genera Pedronia, Clavicoccus, Phylococcus, Lamini-
coccus, Palmicola and Nesococcus apparently ought also to be placed in the Pseudo-
coccus group of genera.

VI. The Planococcus group, includes the remaining genera here studied (i.e.
Planococcus, Planococcoides, Nipaecoccus, Maconellicoccus, Ferrisiana and Triony-
mus) ; these genera also exhibit the generalized conditions of almost all characters
and represent one of the most primitive groups among the material studied. Pairs
of genera within the group share a rather large number of characters (88-1 11) and
the members of the group have 71 characters in common. It may be interesting
to note that this group, unlike all the others here recognized, has no exclusive
characters. The two specializations of the group (i.e. strong development of the
lateral arms of the midcranial ridge and the presence of interocular ridge) and the
three primitive conditions (i.e. large scutum, strong basalare and long setae of
glandular pouches) are the same as in the Pseudococcus group, and the two groups,
as will be discussed, are very closely related.

Beardsley's (I.e.) descriptions of Chorizcooccus lounsburyi (Brain) (=Trionymus
lounsburyi Brain), Trionymus multiductus Beard., T. rostellum'Lobde\\,T.danthoniae
Morrison, Nipaecoccus nipae (Mask.), N. vastator (Mask.), N. longispinus Beard,
and Antonina crawii Ckll. indicate that all these species exhibit the generalized
characters of Pseudococcidae and have no fleshy setae on the body itself, thus con-
firming the writer's concept in including Trionymus and Nipaecoccus in the Plano-
coccus group. The descriptions also suggest that Chorizcooccus and Antonina* also
belong to this group of genera.

Beardsley (i960) also described the males of Trionymus refertus Ferris and
showed that they carry few fleshy setae on the head. For this reason, and to
comply with the present definition of the groups, this species should be placed in
Pseudococcus group. This would imply that it may not be congeneric with the

* The writer, in a subsequent study of 5 Antonina species (Ann and Kosztarab, 1967), decided to
remove the genus Antonina from the Planococcus group and assigned it to the Saccharicoccus group of
genera.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 67

other spp. of Trionymus, since the latter definitely belongs to the Planococcus
group. In this respect it may be interesting to point out that this species was
referred to Trionymus with some reservation, and Ferris (see Zimmerman, 1948)
noted that certain characters of the females (presence of 4 conical setae in each
anal lobe cerarius, complete absence of multilocular pores and very few and small
tubular ducts) separate it well from any other species referred to Trionymus.

Other groups of Pseudococcidae

Beardsley (1962) described the males of another two rather interesting pseudo-
coccid species, Puto yuccae (Coquillett) and Rhizoecus falcifer Kunckel d'Herculais,
both of which showed great structural diversities from the usual males here dis-
cussed. It is evident, in view of these descriptions, that neither species could be
accepted in any of the recognized groups, and therefore the introduction of two
additional ones, Puto and Rhizoecus respectively, seems to be inevitable. It must
be remembered that the identity of these groups is based purely on the available
information extracted from Beardsley's paper.

VII. The Puto group probably represents the most primitive extreme within
the Pseudococcidae. The group has the largest number of exclusive primitive
characteristics, namely : (1) sclerite degeneration less pronounced ; (2) a neck
region well developed ; (3) the presence of numerous disc pores ventrally on the
head (about 12 on each side) ; (4) the presence of more than two pairs of simple
eyes ; (5) the ventral arm of the midcranial ridge posteriorly reaching the preoral
ridges, and the lateral arms absent ; (6) the presence of prosternal apophyses ;
(7) the hamulohalterae usually with more than one apically hooked seta ; (8) the
claw with a preapical denticle and a pair of stout spine-like projections on the inner
face, near its base.

The detailed description of the males of Puto anteunatiis Sign, by Reyne (1954)
indicates that this species is definitely congeneric with P. yuccae, with which it
shares almost all the above-listed exclusive characters (the presence of prosternal
apophyses was not mentioned) ; Reyne's account of the males of M acrocerococcus
superbus Leon, and his illustration of the head (dorsal view) also suggest the
inclusion of the genus M acrocerococcus in this group.

The primitive nature of Puto sp. among Pseudococcidae is also supported by
cytological studies (Hughes-Schrader, 1944).

VIII. The Rhizoecus group : Contrary to conditions in Puto, this group repre-
sents the most specialized extreme of the known males of Pseudococcidae. It
exhibits the largest number of exclusive specialized characteristics, which include :
(1) the head broadly joined to the thorax, the neck region being absent ; (2) sclerite
degeneration of the thorax more pronounced ; (3) absence of a distinct midcranial
ridge ; (4) absence of the cervical sclerites ; (5) absence of the lateral ocelli ; (6)
reduction of the pronotal ridges ; (7) absence of the hamulohalterae ; (8) absence
of the metapleural ridge ; (9) absence of the glandular pouches.

The Rhizoecus group also shows the following conditions which are regarded as

68

MORPHOLOGY AND TAXONOMY OF ADULT MALES

primitive : (i) the absence of the preocular ridges ; (2) the presence of a pair of
prosternal apophyses ; and (3) the less pronounced sclerite degeneration of the
abdominal segments.

According to Hambleton (1946), the genera Rhizoecus, Riper siella, Geococcus and
Pseudorthizoecus appear to be elements of a natural group within the mealybugs, to
which he also added 4 new genera : Radicoccus (Radiococcus, misspelling), Brevi-
coccus, Morrisonella and Neorthizoecus. This group was erected on the basis of
certain female characters (geniculate antennae and nature of the anal ring) and
on the fact that all these genera include ground-inhabiting species. Apart from
Rhizoecus, no information about the males of this group is available, and it would
be interesting if Hambleton's conception of this natural group could be confirmed
on the basis of male characters also.

Relationships between the groups of Pseudococcidae

The relationships between the groups of genera is illustrated in table IV which
shows : (1) the number of characters shared by any pair of groups ; (2) the number
of characters exclusive to each of these pairs ; and (3) the number of characters
by which the pairs of groups differ from each other. These affinities are based on
the total number of shared characters, since the specialized or primitive condition
of most of these could not be determined with great certainty.

Table IV
Relationships between the groups of genera

Number of Characters.

Pairs of groups

Planococcus — Pseudococcus
Planococcus — Saccharicoccus
Planococcus — Octococcus
Planococcus — Ceroputo
Planococcus — Nairobia
Pseudococcus — Saccharicoccus
Pseudococcus — Octococcus
Pseudococcus — Ceroputo
Pseudococcus — Nairobia
Saccharicoccus — Octococcus
Saccharicoccus — Ceroputo
Saccharicoccus — Nairobia
Octococcus — Ceroputo
Octococcus — Nairobia
Ceroputo — Nairobia

Examination of this table suggests that :

(a) The Planococcus and Pseudococcus groups are the closest relatives, sharing
the largest number of characters (44) and differentiated by the smallest (4).

(A)

(B)

(C)

Shared

of which
exclusive

Differentiating

44

4

38

10

37

11

25

22

26

22

42

14

36

21

22

35

24

33

3i

27

18

39

20

38

32

4

25

28

30

34

8

24

OF l'SEUDOCOCCIDAE & ERIOCOCCIDAE 69

(b) Planococcus, Pseudococcus, Saccharicoccus and Octococcus groups seem to be
closely related ; they exhibit the generalized typical pseudococcid condition and
share a comparatively large number of characters (31-44).

(c) The Ceroputo group is the closest relative to Nairobia, and vice versa ; the
two groups have the largest number of exclusive conditions (8), and share 34
characters, which is more than each of them shares with any other group.

These relationships, and taking into consideration the available published
information, probably indicate that there are four major sections of Pseudococcidae :

I. The Rhizoecus section which represents the most specialized extreme.

II. The Ceroputo + Nairobia section which is less specialized.

III. The Planococcus, Pseudococcus, Saccharicoccus and Octococcus section, which

includes closely related and rather primitive groups.

IV. The Puto section, which appears to represent the most primitive extreme

of all.

It is difficult to ascribe any definite taxonomic status to these sections, but they
may possibly deserve subfamily ranks, with further subdivision into tribes and
subtribes based on the groups of genera suggested here. However, further investiga-
tions of more widely representative material is required before more accurate
conclusions could be made.

b. Genera

In table V, 73 characters are listed, separating the genera in at least one of the
three groups in which more than one genus was available. When a character can
be used taxonomically within a particular group, it is marked " G " in the table ;
if it holds at the specific level only or at both generic and specific levels, the marks
" S " or " GS " respectively are given ; the mark " — " indicates that the character
does not apply at all within the group.

Table V
List of characters separating genera

Characters

1. Presence and position of fleshy setae on body.

HEAD :

2. Nature of ventral preocular depression.

3. Nature of dorsal arm of midcranial ridge.

4. Degree of posterior development of dorsal arm.

5. Nature of the lateral arms of the midcranial ridge.

6. Nature of the postoccipital ridge. —

7. Condition of the postoccipital ridge anteriorly. GS

Plano-

Pseudo-

Ceroputo

coccus

coccus

group

group

group

GS

—

G

GS

.

G

—

G

G

G

G

G

7 o

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Characters

8. Nature of the ventral rudiment of the preocular

ridge.

9. Distance between the dorsal eyes related to their

corneae.

10. Relative size of dorsal and ventral eyes.

11. Nature of attachment of ocellus to postocular

ridge.

12. Number of dorsal head setae.

13. Number of genal setae.

14. Number of dorsal ocular setae.

15. Presence of setae between ventral eyes.

16. Number of ventral head setae, transversely

arranged in front of the ventral eyes.

17. Presence and position of head pores.

18. Number of disc pores, if present, dorsally only

on the head.

Piano-

Pseudo-

Ceroputo

coccus

coccus

group

group

group

GS

S

G

GS

G
G

GS

G
G

G
G

G

G
G

GS

ANTENNAE :

19. Relative length of antenna to body.

20. Relative length of antenna to hind leg.

21. Relative length of 3rd and terminal segments.

22. Width of segment III related to its length.

23. Presence of fleshy setae on antennae.

24. Relative length of antennal and body setae.

25. Number of hair-like setae on scape.

26. Number of antennal bristles on terminal segment.

27. Relative length of the two lateral bristles of

terminal segment to those on preceding segments.

28. Relative lengths of the three large bristles of

terminal segment.

THORAX :

29. Shape of prosternum.

30. Nature of prosternal ridge.

31. Number of medial pronotal setae.

32. Number of lateral pronotal setae.

33. Number of post-tergital setae.

34. Presence of prosternal setae.

35. Number of medial pronotal pores.

36. Number of lateral pronotal pores.

37. Number of antespiracular dorsal pores.

GS

S

GS

s

GS

s

G

s

GS

G

GS

s

GS

G
G

GS

—

G

GS

—

—

GS

S

—

GS

GS

G

S

G

—

—

G

GS

—

—

s

G

GS

s

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

7i

Characters

38. Number of prosternal pores.

39. Relative length of prescutum to scutum.

40. Appearance of scutellum in dorsal view.

41. Presence of a membranous area on the

lateropleurite.

42. Number of prescutal setae.

43. Number of scutal setae.

44. Number of scutellar setae.

45. Number of tegular setae.

46. Position of postmesostigmatal setae.

47. Number of postmesostigmatal pores.

48. Number of metatergal setae.

49. Number of metapleural setae.

50. Number of anterior metasternal setae.

51. Number of posterior metasternal setae.

52. Presence of metatergal pores.

53. Presence of metaspiracular pores.

54. Presence of anterior metasternal pores.

55. Presence of posterior metasternal setae.

56. Number of alar setae.

57. Number of circular sensoria on fore wings.

58. Relative length of the hamulohalteral seta to

length of the hamulohaltera.

59. Relative length of fore legs.

60. Presence of differentiated apical seta on trochanter.

61. Relative length of fore and middle femora.

62. Relative width of hind femur to its length.

63. Relative lengths of tibia and femur in fore legs.

ABDOMEN :

64. Nature of ostioles.

65. Number of ventral setae on abdominal segment

VIII.

66. Presence of disc pores on abdominal segments.

67. Position of abdominal pores.

68. Relative length to width of penial sheath.

69. Relative length of genital segment to total

length of body.

70. Appearance of style in lateral view.

71. Condition of process of penial sheath.

Piano-

Pseudo-

Ceroputo

coccus

coccus

group

group

group

GS

S

—

—

—

G

—

—

G

—

—

G

GS

—

G

S

S

G

GS

S

—

GS

GS

G

G

S

—

G

S

G

GS

S

G

GS

s

G

GS

s

—

GS

s

—

G

s

—

GS

s

—

GS

s

—

GS

s

—

GS

GS

G

G

s

—

G

—

G

G

—

—

GS

—

—

GS

s

—

G

—

G

G

—

—

GS

GS

S

G

s

GS

GS

GS

s

G

s

G

s

G

s

72

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Piano-

Psendo-

Ceroputo

coccus

coccus

group

group

group

G

S

—

Characters

72. Comparative length of aedeagus.

73. Number of dorsal setae of the genital segment. — — G

c. Species

Table VI includes 57 characters separating the species in at least one of the three
genera in which more than one species was available for study. The characters
operating taxonomically within a particular genus are indicated by the mark

X ", and those not operating by the mark " O ". The table shows that the
species of these genera could be separated by a large number of characters including
differences in the development of various structures, the number and distribution
of the setae and disc pores.

Table VI

2.
3-

4-

5-

6.

9-
10.
II.

12.

13-
14.

15-
16.

17-

18.

List of characters separating species

Characters
1. Presence of fleshy setae on antennae and legs.

HEAD :

Nature of ventral preocular depression.
Anterior development of postoccipital ridge.
Distance between dorsal eyes in relation to

diameter of their corneae.
Relative size of dorsal and ventral eyes.
Number of genal setae.
Number of head pores.

ANTENNAE :
Relative length to body.
Relative length to hind legs.
Relative lengths of 3rd and terminal segments.
Width of segment III related to its length.
Relative length of antennal and body setae.
Number of antennal bristles of terminal segment.
Relative size of the three large bristles on terminal

segment to other setae.
Relative lengths of the three large bristles of the

terminal segment.
Position of the dorsal bristle of terminal segment.

THORAX :

Shape of prosternum.

Nature of prosternal ridge.

Piano-
coccus

O

x
x

X

O
O
x

X

O

X

O
O
O

O

Nipae-
coccus

O
O

x

X
X

O
O

X

O
O
x

X

Pseudo-
coccus

O
O

o

o
o
o

X
X

x
x
x

o

x

o

X

X

X

X

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

73

Characters

19. Number of medial pronotal setae.

20. Number of lateral pronotal setae.

21. Number of post-tergital setae.

22. Number of antespiracular dorsal setae.

23. Number of antespiracular ventral setae.

24. Number of medial pronotal pores.

25. Number of lateral pronotal pores.

26. Presence of post-tergital pores.

27. Number of antespiracular dorsal pores.

28. Number of prosternal pores.

29. Relative length of prescutum to its width.

30. Number of prescutal setae.

31. Number of scutal setae.

32. Number of scutellar setae.

33. Number of tegular setae.

34. Position of postmesostigmatal setae.

35. Number of postmesostigmatal pores.

36. Number of metatergal setae.

37. Number of metapleural setae.

38. Number of anterior metasternal setae.

39. Presence of postmetastigmatal setae.

40. Number of posterior metasternal setae.

41. Presence of metatergal pores.

42. Presence of metaspiracular pores.

43. Presence of anterior metasternal pores.

44. Presence of posterior metasternal setae.

45. Number of alar setae.

46. Number of circular sensoria on fore wings.

47. Presence of apical setae on trochanter.

48. Relative lengths of fore and middle femora.

ABDOMEN :

49. Nature of ostioles.

50. Number of ventral setae on abdominal segment

VIII.

51. Presence of disc pores on abdominal segments.

52. Position of abdominal pores.

53. Relative length to width of the penial sheath.

54. Relative length of genital segment to total length

of body.

55. Lateral appearance of style.

56. Condition of process of penial sheath.

57. Comparative length of aedeagus.

Piano-

Nipae-

Pseudo-

coccus

coccus

coccus

O

X

X

X

X

X

O

O

X

O

O

X

O

O

X

X

X

O

X

X

X

X

X

X

X

O

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

o

X

X

X

X

X

X

X

X

X

X

X

X

X

74 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Eriococcidae

A division of this family into groups of genera, similar to that of Pseudococcidae
could be suggested at the present stage. Only 7 species (representing 4 genera)
belonging to this family were available, and it was not possible to assess correctly
the significance of the morphological differences they exhibit. However, the
distribution of the body setae and the structure of the genital segment appear to
be most significant. Inspection of table II shows that of the 3 spp. of Eriococcus,
E. araucariae shares a comparatively large number of characters with E. orariensis
(118), but each has only 101 and 102 characters respectively, in common with
E. buxi. This would suggest that the latter is less closely related to the other two
species and would support Borchsenius's (1963) view that E. buxi (the type species
of Eriococcus) is not congeneric with the other species hitherto ascribed to the same
genus.

On the other hand, the structure of the genital capsule indicates that the studied
species could be grouped as follows :

a. E. araucariae and Gossyparia spp., where the style is undulated and the base
of the aedeagus distinctly sclerotized latero-dorsally.

b. E. buxi, 0. agavium and P. fraxini, where the genital capsule is comparatively
short and the style rather straight.

c. E. orariensis, where the genital capsule is comparatively elongate (its length
more than 2-5 times its width).

It should be emphasized, however, that the available material was quite
inadequate and the above suggestions are very tentative indeed.

The genera and available species of family Eriococcidae are merely separated by
differences in chaetotaxy and the development of certain structures. These are
listed in table I and will be used in the keys.

Relationships of Pseudococcidae and Eriococcidae with other Coccoidea

According to Balachowsky's (1937, 1942) original designation, margaroids —
the most primitive type of Coccoidea males, comprise Ortheziidae and Margarodidae
(including the aberrant species Steingelia gorodetskia) ; lecanoids comprise all the
other families (including the monotypic family Phenacoleachiidae), except
Diaspididae and Phoenicococcidae, which constituted the diaspidoids. Balachowsky
stressed that the diaspidoids represent a homogenous group fundamentally differing
in the characters of the males and females from the other two groups. Theron
(1958), who gave a detailed list of morphological characters separating the three
types of males, and later also Giliomee (1961, 1967), both concluded, however, that
although the lecanoids are more specialized than the margaroids and less than the
diaspidoids, they are nevertheless more closely related to diaspidoids than to
margaroids.

As far as the relationships of Phenacoleachiidae with other coccid families are
concerned, Morrison and Morrison (1922) maintained that the sole species of this
family, Phenacoleachia zealandica (Maskell) has a number of characteristics of

OF PSEUDOCOCCIDAE & ERIOCOCCID AE 75

Pseudococcidae " e.g. the kinds of pores, the character of the anal ring ", and other
characteristics of Margarodidae " e.g. the number of antennal segments, the
absence of differentiated tarsal digitules ". At the same time it has certain characters
refuting any relationship with either of these families " e.g. absence of dorsal
ostioles disclaiming relationship with Pseudococcidae, and presence of anal ring
with setae and pores and absence of abdominal spiracles, disclaiming affinity with
Margarodidae". As already mentioned, Balachowsky included the Phenacole-
achiidae in the lecanoids (although admitting affinity to the margaroids mainly by
the possession of n-segmented antennae). Theron (1962), however, after detailed
study of the males concluded that Phenacoleachia zealandica " ostensibly belongs
to the margaroid group ", showing a particularly close relationship to the aberrant
margaroid Steingelia gorodetskia. He regarded them as annectant genera linking
the primitive Margarodidae with the more specialized lecanoids. Giliomee (1967),
on the basis of a simple numerical analysis of the characters available for comparison,
was able to demonstrate that Steingelia apparently represents a link between
Margarodidae and Coccidae, while Phenacoleachia a similar link between Margaro-
didae and Pseudococcidae. This suggestion was strongly supported by the recent
discovery and description by Beardsley (1964) of the new second Phenacoleachia
species, P. australis from Campbell Islands, which shows the presence of such
pseudococcid features as " ostioles and cellular anal ring ". These features tempted
Beardsley to suggest that the Phenacoleachiidae are definitely allied to the mealy-
bugs and probably represent a primitive subfamily of Pseudococcidae. The males
(apterous) also described by Beardsley, have two very distinctive pseudococcid
characters (as shown in the present work), i.e. the presence of disc pores and of
dorsal ostioles.

The study of males has therefore revealed that there appear to be at least two
links between the primitive margaroids and the more specialized lecanoids. On the
other hand, a close relationship between Asterolecaniidae (lecanoid) and diaspidoids
was recently suggested on the basis of cytological studies and of some characters of
the female (Brown & McKenzie, 1962). Thus there may exist similar links between
lecanoids and diaspidoids. There is no detailed information about the males of
Asterolecaniidae,* but their study may perhaps also support this suggestion.

Theron (1958), Ghauri (1962) and Giliomee (1967) each discussed the relationships
of various groups of Coccoidea based on their own observations and on the informa-
tion then available. Each contribution naturally allowed for more detailed and
comprehensive comparisons and conclusions. For the present author, Giliomee's
conclusions are of particular interest since he was dealing with the lecanoids, and
included in his discussion not only Coccidae (the main object of his studies) but also
males of 3 species of Pseudococcidae described in detail in his earlier paper (1961).
Incidentally, his data on these species have been included here in the tables,
calculations and discussion. Giliomee pointed out that the Coccidae are more

* The males of an asterolecaniid (Cerococcus deklei Kosztarab and Vest) were later available to the
author. They completely agreed with the lecanoid type of males, as will be shown in a separate
publication.

76 MORPHOLOGY AND TAXONOMY OF ADULT MALES

specialized than Pseudococcidae since they generally share more characters with the
highly specialized Diaspididae. Dealing with lecanoids, Giliomee found it necessary
to introduce minor amendments to Theron's definition of the lecanoid type of
male (these include the presence of the metasternal plates and the abdominal
sclerites). The present study of Pseudococcidae and Eriococcidae has also revealed
that two further minor amendments are necessary, i.e. regarding the presence of a
distinct postoccipital ridge and of the metasternal apophyses.

In an attempt to determine the affinities of Coccoidea, it is now possible to
consider 6 families for which adequate information on the males is now available,
i.e. Margarodidae, Phenacoleachiidae, Pseudococcidae, Eriococcidae, Coccidae and
Diaspididae. Steingelia, although a margarodid, is treated separately because of its
aberrant characters. Thirteen available characters were relevant for this purpose,
and the distribution of their conditions within the families is shown in table VI.
Each of the characters occur either in a specialized condition (marked " X ") or a
primitive one (marked " O ").

Table VI
Male characters whose conditions indicate affinities within coccoidea

•o S o «2 2

"3 .2 -3 ■« "°

o

"v o o y iS !2

iS Jj J3 u) 1- O -JT

g m CU d, W cj Q

Specialized conditions of the characters ( x ) :
i. Disc pores absent. O O XQxO X XO X

2

3
4
5
6

7
8

9
io
ii

12

13

Neck region absent. O O O O O O X

Postoccipital ridge absent. O X O O X O

Interocular ridges present. O O OxOOxOO

Tentorium absent. O O O O O O X

Compound eyes absent. O X X X X X X

Membraneous area of scutum absent. X O O X X X X

Scutellum short and transverse. O X O O X X X

Median ridge of basisternum absent. O O X X X O

Metasternal sclerites absent. O O O X X O O

Metasternal apophysis absent. O O O O O X X

Ostioles present. O OXOxOO O O

Penial sheath elongate. O O O O O X X

Specialized conditions. — 3 3 4 6 5

Primitive conditions. 12 10 8 6 7 6

Both conditions present. 1

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

It should be noted that both conditions may obtain in certain families. At the
bottom of the table are given the total number of occasions on which the specialized,
primitive, and both conditions occur in each group. The proportion of the number
of primitive to specialized conditions of the characters can be taken as indication to
the primitive or specialized nature of the group, e.g. the primitive Margarodidae
have no specializations, 12 primitive characters and one in which both conditions
occur, and the specialized Diaspididae show 8 specializations and 5 primitive
conditions. Table VIA, prepared in the form of two half matrices, in which are
given the total number of characters shared by each pair of groups (upper triangle
of the matrix), and separating each pair (lower triangle). In this table the charac-
ters which occur in both conditions in either member of a pair are omitted, e.g.
between Margarodidae and Steingelia, the character 7 in table VI (membranous area
of scutum) which may be present or absent in Margarodidae, was not taken into
consideration, and therefore only 12 characters are left for comparison.

Table VIA

The number of characters shared by, and separating each pair of families:

V

Cfl

~z

4>

(8

Sharing

•0

B

u

(J

§

01

•5

c

V

u

6

-
to

•5

Separating

u

M

C

S3

c

•a

3
■-

!2
u

5

s

X

P-

P-

W

'-^

P

Margarodidae

9

8

i>

7

5

5

Steingelia

3

7

5

8

9

6

Phenacoleachiidae

2

4

9

9

4

5

Pseudococcidae

3

5

1

9

3

3

Eriococcidae

5

5

2

1

5

7

Coccidae

5

2

6

7

6

7

Diaspididae

7

7

6

7

6

4

It appears from these tables that the Eriococcidae are more specialized than the
Pseudococcidae. They have 6 specializations (Table VI) and share a total number
of 7 characters with the decisively more specialized Diaspididae (Table VIA), while
the Pseudococcidae have only 4 specialized conditions and share only 3 characters
with Diaspididae. It is also interesting to note that the structure of the genital
segment suggests a closer relationship between Eriococcidae and Diaspididae than
between the latter and Pseudococcidae.

78 MORPHOLOGY AND TAXONOMY OF ADULT MALES

From the same tables it appears that Giliomee was justified in assuming that the
Coccidae are more specialized than Pseudococcidae. They have 5 specialized and
6 primitive features against 4 specialized and also 6 primitive in the latter, and
comparison with Diaspididae shows that they share a total number of 7 characters
with Coccidae but only 3 with Pseudococcidae.

Table VIA also indicates that Coccidae are closer to Eriococcidae (sharing 5
characters) than they are to Pseudococcidae (sharing only 3 characters). In the
same way it also appears that the Eriococcidae are more closely related to the
Pseudococcidae (sharing 9 characters) than they are to the Coccidae (sharing only
5 characters).

Table VIB is also derived from table VI and prepared in the form of a matrix.
Here the relationships between the groups are expressed in fractions obtained by
dividing the number of shared conditions by the total number of characters con-
sidered. Those characters occurring in either specialized or primitive conditions
were counted as 1 unit, and if both conditions occurred in any of the compared

Table VIB

Steingelia

Phenaco-
leachiidae

Pseudo-
coccidae

Erio-
coccidae

Coccidae

Diaspi-
didae

Margarodidae

(073)

(073)

(0-62)

(0-58)

(050)

(0-42)

Steingelia

(062)

(050)

(062)

(077)

(0-46)

Phenacoleachiidae

(o-88)

(077)

(0-46)

(0-46)

Pseudococcidae

(o-8i)

(0-42)

(o-35)

Eriococcidae

(0-46)

(0-54)

Coccidae

(0-62)

Diaspididae

groups, they were both given a score of \ unit each, e.g. Margarodidae and Steingelia
share 9 characters and are separated by 3, but in the former there is one character
(No. 7) occurring in both conditions ; the numbers thus corrected will be 9J shared
and 3! separating. The number of the shared characters (9!) was divided by the
total number of characters (13) and the result (073) was entered in the table as a
measure of similarity. The values obtained vary from 0-1 indicating no or complete
similarity respectively, i.e. the higher the values are, the closer the relationship is.

From this table it is obvious that Steingelia and Phenacoleachiidae are similarly
related to Margarodidae, and are closer to it than they are to each other. It is also
apparent that Phenacoleachiidae are more closely related to Pseudococcidae than
they are to Margarodidae, which would support the views of Beardsley. It also
appears that Steingelia is closer to Coccidae than to the Margarodidae.

Giliomee (1964) discussed the taxonomic status of the family Kermococcidae, which
Ferris (1937) placed in the family Eriococcidae, but which Balachowsky (1942)
included as a subfamily in one family (Eriococcidae) together with Pseudococcinae

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

79

and Eriococcinae. The description of the males of Kermococcus quercus (L.) by
Borchsenius (i960) prompted Giliomee to assume that Kermococcidae are more
closely related to Coccidae than to any other Coccoidea, on the basis of 6 characters
shared by the two families. Further investigations on Kermococcidae are required
before Giliomee's assumption can be supported since it was based on one species
only, and since the present study revealed that some of the characters he used are
also shared by Pseudococcidae (separate pre- and postocular ridges) and Eriococcidae
(separate ocular ridges ; short and transverse scutellum).

ANALYSIS

On the suggestion of Dr. K. Boratynski, advantage was taken of the computing
facilities (IBM 7090 computer) available at Imperial College, and the data obtained
in this study (138 characters for 29 species) were subjected to the more elaborate
statistical analysis by some methods of numerical taxonomy. In their book on the
subject, Sokal and Sneath (1963, p. VIII) define the numerical taxonomy as "the
evaluation by numerical methods of the affinity or similarity between taxonomic
units and the employment of these affinities in erecting a hierarchic order of taxa " ;
it " aims to develop methods which are objective and repeatable both in evaluation
of taxonomic affinity and in the erection of taxa ". They pointed out that assess-
ment of the degrees of affinity of the taxa (expressed as a coefficient of similarity)
should be based on a large number of characters (at least 60) to assure objectivity
and repeatability. All these characters should be treated as of equal taxonomic
value and the equivalence of all the characters is one of the basic assertions of
numerical taxonomy. The relationships between taxa thus calculated is " phene-
tic ", i.e. based on overall resemblance and is free of subjective phylogenetic
speculation. But the authors believe that numerical methods may open up a wide
field in the exact measurement of evolutionary rates and may provide a more
critical approach to phylogenetic problems (pp. VIII, 57 and chapter 8).

Method

The method employed in the main part of this thesis contains also an element of
objectivity in the fact that all the characters were treated as being equivalent ;
the assessment of the degree of similarity was based on simple counting of the
characters shared by each pair of species. But this method does not take into
consideration which of the characters are shared by various species ; moreover, a
certain amount of subjective selection of characters was used in the grouping of
taxa. For example, the separation of Planococcus and Pseudococcus groups of
genera was based on one exclusive character, i.e. the absence or presence of fleshy
setae on the main parts of the body.

The comparison of these results with those obtained by more accurate calculation
of coefficient of similarity between each pair of species, was thought to be of interest.

It should be pointed out that the application of numerical taxonomy is still in
an experimental stage, and the exact value of the various suggested methods for

80 MORPHOLOGY AND TAXONOMY OF ADULT MALES

satisfactory classification is to be investigated further. A number of papers have
been published, in which numerical taxonomy was utilized. In Entomology it was
applied by Stroud (1953) for Termites ; Sokal (1958) and Sokal and Michener (1958)
for the classification of Hoplitis complex of Bees ; Rohlf (1963) for Mosquitoes ;
and Sheals (1964) for Acarina.

The computer programmes were available for two methods of statistical analysis,
which were used for my data :

The Principal Component (vector analysis) method, using the BIGMAT programme
developed by M. Elson and R. E. Funderlic of Central Data Processing Facility,
Oak Ridge, Tennessee, U.S.A., and available at the Imperial College computer
unit (Share pamphlet). This programme generates first a Covariance matrix
calculated from

I(X ij -X j )(X lh -X ll )

i = l

which basically is a coefficient of similarity ; and then the subroutine BIGMAT
Fortran IV matrix eigensystem solver calculates eigenvalues (latent roots) and
eigenvectors (latent vectors) of real symmetric matrices and can operate up to a
matric limit of 235 X 235. The " normal " analysis (Q-technique) which ordinates
the species (OTU's) was used.

The Taxon analysis method as described by Sokal and Sneath (1963, p. 195 and
296), with clustering by the weighted pair-group method using Spearman's sums of
variables method (WPGM) (p. 309). This method uses the correlation coefficients
between each pair of OTU's based on standardized characters and calculated by
the computational formula :

|XuX ik -i(|x ij )(|x ik )

i = l v i = l ' J L i = 1 X i = l '

This programme was developed in the Department of Zoology, Imperial College,
by Dr. G. Murdie specially for the purpose of these studies. The author, who is
not very familiar with the statistical methods involved, is grateful to Dr. Boratynski
for his guidance and to Dr. Murdie for developing the Taxon programme.

The method based on taxonomic distance coefficient, which in some cases was
shown to give better results, could not be applied since the appropriate computer
programme was not ready in time.

Procedure

The 29 species included in this study are listed on the right side of the dendogram
of Fig. (A), and the order or numbering corresponds to the degree of relationships
and the classification suggested by the analysis in the main part of this thesis.
Family Pseudococcidae include the first 21 species and family Eriococcidae species

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 81

22-29. Species 1-9 = Planococcus group of genera ; species 10-15 — Pseudococcus
group ; species 16 and 17 = Saccharicoccus group ; species 18 = Odococcus group ;
species 19 and 20 = Ceroputo group, and species 21 = Nairobia group.

The 138 characters taken into consideration are the same as those listed in
table I (see Discussion, p. 31). Their different state conditions were expressed in
numerical code ranging from 1 to 5, but in most cases only 2-3 conditions were
used. The better the structure is developed the higher the value given, e.g. the
absence of a certain ridge was given 1, its weak development 2 and its strong
development 3 ; if the absence of setae or pores on a particular part of the body was
given the value of 1, the presence of 1-3 setae was given 2, and 4-6 for example
will be 3, and 6-10 will be 4 ; the small size (e.g. body) was given 1, intermediate 2,
and large 3. These conditions were punched on to the cards for processing on the
IBM 7090 Fortran IV electronic computer available at Imperial College.

In the Principal Component method, the computer generates covariance matrix
but prints only the diagonal values, the sum of which (Trace) represents the amount
of total variance (Text-fig. I). Example of Pseudococcidae above.

The BIGMAT calculates any desired number of latent roots (eigenvalues) and the
corresponding latent vectors (eigenvectors) ; as usual 10 were calculated, but only
the first three were used for plotting (Text-fig. II).

Each latent root represents an axis which is perpendicular to all the other axes,
and the latent vectors represent the co-ordinates on these axes, thus locating the
species in the N-multiple superspace. The value of the latent root, expressed as a
percentage of the sum of diagonal values, gives the amount of variance (Trace)
accounted by each root. This amount is the highest for the first root and gradually
decreasing in the other roots. The first three in this case (29-41%, 15-96% and
11-33%, respectively) account for 56-7% of total variance, and these were used
for plotting (the remaining 7 roots account for 6-71%, 6-12%, 5-27%, 3-89%,
3'7°%» 3'4°% an< i 2-87%, respectively). The values of the co-ordinates are
calculated by multiplying the eigenvalues by the square root of the corresponding
latent root of each species. For the purpose of plotting an integral number was
added to the latent vectors of each latent root to eliminate negative values, and in
this case 6, 5 and 6, respectively were added to the three roots. Table 1 shows the
data for the pseudococcid species prepared for plotting.

The 3-dimensional block diagrams of the first three vectors were made separately
for (1) all forms of Pseudococcidae and Eriococcidae studied, (2) Pseudococcidae
and Eriococcidae excluding the apterous males, and (3) Pseudococcidae only
(Figs. 1, 2 and 3, respectively). In these diagrams the space was divided by
horizontal planes. The first vector (axis) was represented by one side of the square
base (horizontal, I in Figures), the third by the other side (III, oblique in the
Figures), and the second vector by the vertical axis (II). For the purpose of easier
reference to the actual location of the species, each side of the base (and other planes)
was divided into 3 sections marked A, B, C for axis I, and a, b, c for axis III. The
resulting 9 squares are defined by the reference to the appropriate sections on the

EIGENVECTOR CORRESPONDING TO
2.4G15854E-G2 7 .82261 04E-03
1.0492194C-02 -1.5792798E-01

22.592
24.392
38.1*92
24.992
34.292
54.492

28.

45.

37.

25.

33.

46.

52.

54.

44.

54.

73.

95.
101.
123

892
092
992
792
592
692
092
292
592
392
592
592
792
49?

210 1017.14 Total variance (Trace)

Text-fig. I. Trace of 20 spp. of Pseudococcidae.

2.991 3983E ;2 (= Latent root)

- 1.295 1852E-03 -3. 5644727F-02 5.6835338E-02

-2.238852BE-G1 -2.65624f.4E-0 1 -3. 1 658460E-C 1

1.2530954E-01 -5.9564865E-02 7. 64500 1 6E-03
•3. 1533943E-01 -2.4942465 E-C 1 -8. 29552 10E-02

2.9370538F-01 4 . 4PU3477E-C 1 3. 1 U38629H-0 1 U.236C780E-G1

EIGENVECTOR CORRESPONDING TO 1.6239042E 02

-2.2098C02E-0 1 -2 .3 159923E-01 -3. 09C3342E-0 1 -2. 75861 03E-Q1

-1.521U205E-C1 1.3926755E-02 1 .6055679E-01 1 . 9671679F-0 1

3.1300302E-02 1.8158900E-01 2. 5952574E-C 1 3.7i*38265E-01

EIGENVECTOR CORRESPONDING TO
-7.8359791E-C2 5.4741075E-02
1.C589229E-01 7 . 54C2325E-02
-2.5461 197E-01 2. 7366 1 96E-0 1

1.1523669E 02

8.2U61756E— j2 -9. 1954241E-02

1 . 606450.1 1-02 4.6854463E-02

6.4068813E-C1 -4 . 8C 7 5832E-0 1

-2.8919946E-01 -3.6976436E-01 - 1 . 2965085E-0 1
2.3084065E-01 2.213579 1E-0 1 1 .9066565E-01

1.5312027E-02 -4. 1155443E-03 4. 81 77828E-02
1.3160295E-02 1 .0 1 58375E-0 1 - 1 . 1463530E-02

TfiXT-FIG. II. Latent vectors corresponding to first three latent roots for 20 spp. of Pseudococcidae. The actual values of
the co-ordinates for plotting are calculated by multiplying each eigenvector by the square root of the corresponding latent root.

1.3466497E-01
■2.0218696E-02

-3.4256143E-01
-2. 1068075E-01

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

83

8 4

MORPHOLOGY AND TAXONOMY OF ADULT MALES

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

85

86

MORPHOLOGY AND TAXONOMY OF ADULT MALES

1

H 1

1
o

o

1

o

i
o

o

CD

0)

*>

CO

1

1

1

i

1

(1) P.citri

(2) P.kenyae

(4) P.ireneus

(3) P.dioscoreae

(7) M.hlrsutus

(9) T.newsteadi

(5) N.vastator

(6 ) N .nipae

(8) F.virgata

(10) P.obscurus

(11) p:Vnaritlrau3 A"

(12) P.citriculus

(13) P .adonldum

(14) P.fragilis

(15) D.alazon

(16) S.sacchari

(17) « '• (apterous)
(29) P. fraxini (apterous)

(18) O.africanus

(19) C.pilosellae

(20) C.inaolitus

(21) N.blfrona

(22) E.araucariae

(23) E.orariensis

(25) O.agavium

(26) G. spuria

(27) '» '» (brachypterous)

(28) G.salicicola ( n )

(24) E.buxi

Dendrogram A.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

87

Correlation coefficient scale

0000

K) i*» <H Q9

1

M

1

1

O

1

1

•

O

00

o>

»

to

r

1

1

(1)

(2)
(4)

H (3)
(7)
(9)
(5)
<6)
(8)
(10)
(11)
(12)
(13)
(14)
(15)
(16)
(18)
(19)
(20)
(21)
(22)
(23)
(26)
(27)
(28)
(25)
(24)

Dendrogram B.

88

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Correlation coefficient scale

-0.8
-1.0

-0.2
-0.4
-0.6

D

O

.

to

o

o

o>

O H
CD O

i I i i i

1 1

1

1

(1)

(2)

(4)

(3)

(7)

(9)

(5)

(6)

(8)

(10)

(11)

(12)

|-

(13)
(14)
(15)
(16)
(18)
(19)

(20)
(22)
(23)

(26)
(27)

1

1

(28)

(25)
(24)

i i

i i i

1

i i i

Dendrogram C.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

89

Correlation coefficient scale
1 1 • •

0000 0000
. . . . o •

QSOi^tO tO 4>> 0> OD

Dendrogram 1).

go

MORPHOLOGY AND TAXONOMY OF ADULT MALES

sides ; the vertical position is referred to as " level " above which the species is
located. This way of plotting was suggested by Dr. K. Boratynski.

In the Taxon programme the usual correlation coefficient matrix is generated
(each diagonal value = i), and a series of clustering cycles are made. After each
cycle, a new correlation coefficient matrix is recalculated. In the employed weighted
pair-group method (WPGM) only the two most highly correlated stems are allowed
to join at each clustering cycle. Members of a cluster include either the individual
species or the smaller clusters to form another cluster of a higher rank. The
growing clusters join the more remote relatives at a lower point of similarity (or
correlation coefficient). The dendograms of Figs. A, B, C, D and E are prepared to
illustrate the values of correlation coefficient at which the OTU's or their groups
are joined together.

Correlation coefficient scale

Dendrogram E.

OF PSEUDOCOCCIDAE & ERIOCOCCID A E

9i

Table I
Data of the species of Pseudococcidae prepared for plotting

Latent root

1

II

III

% of variance (Trace)

29-4i°o

i5-96°o

n-33%

Integral number added to make the latent
values positive.

4 6

~5

1-6

Species

Latent values

1

P. citri

2

P. kenyae

3

P. dioscoreae

4

P. ireneus

5

N. vastator

6

N. nipae

7

M. hirsutus

8

F. virgata

9

T. newsteadi

10

P. obscurus

11

P. "maritimus A

12

P. citriculus

13

P. adonidum

M

P. fragilis

15

D. alazon

16

S. sacchari

18

O. africanus

19

C. pilosellae

JO

C. insolitus

21

N. bifrons

642

2-18

516

613

2 05

6-59

5 98

1 06

689

538

1-48

502

698

1-3*

617

8-17

029

5 96

497

335

652

613

672

232

618

306

7-M

327

5-i8

681

213

705

617

141

751

650

052

7 94

614

o-55

7-82

709

1 69

743

588

456

474

374

1 1 08

5 4°

327

1 3 62

7-31

8-94

11-51

831

12-87

1333

981

084

Results

The results of the Principal Component analyses are shown in Figs. 1, 2 and 3.
In Fig. 1, in which all species were considered (including the apterous forms), the
macropterous forms are situated in the upper part of the diagram, and except
S. sacchari (16) above the third level, while the two apterous forms (S. sacchari,
17 and P. fraxini, 29) near the base, below the second level. This is attributed to
the fact that these males, unlike the apterous males of other families (e.g.
Diaspididae, Ghauri 1962) are greatly degenerate and have lost most of their
morphological structures. They show considerable differences as compared with
the macropterous forms even of the same species. It has been decided therefore
to exclude these two species, and another set of calculations for the macropterous
males only was made ; the results are illustrated in Fig. 2. The two diagrams are
virtually identical as far as the relative position of the species are concerned, except
for S. sacchari (16) whose position is closer to Planococcus group (spp. 1-9). The
relative position of the species, as shown in Fig. 2, will be discussed in greater detail
since it appears to illustrate the relationships more clearly. Pseudococcidae (1-21)
are well separated from Eriococcidae (22-29) by a vertical place passing diagonally

92 MORPHOLOGY AND TAXONOMY OF ADULT MALES

across the base through the near left and the far right edges of the block diagram.
The Pseudococcidae are situated beyond this plane and the Eriococcidae in front
of it. Nairobia (21) lies in the plane near Eriococcidae. Pseudococcus group
(10-15) occupy space (Ab) on the " ground floor level " ; Planococcus group (1-9)
space (Aa) on the " first floor level " except F. virgata (8) which occupies space (Ab)
on the same floor. The position of Saccharicoccus (16) indicates that it is probably
more closely related to Planococcus group than is F. virgata (8). All these species,
however, are concentrated in the left-hand corner of the block diagram forming
a rather close group of genera. Octococcus (18) occupies a solitary position in the near
left upper part of space (Bb) of the second floor, and Ceroputo and Centrococcus
(19 and 20) are closely approximated in space (Be) of the third floor. The Erio-
coccidae form a fairly compact group in the spaces (Ca) and (Cb) on the ground
floor, with 0. agavium (25) isolated on the second level in space (Cb). Nairobia (21)
is situated in the same space (Bb), second level) as Octococcus (18), but very near
the right margin and close to Eriococcidae. The diagram of Pseudococcidae alone
(Fig. 3) shows similar relationships, only with differences regarding the position of
the groups ; in this diagram F. virgata (8) is also removed from Planococcus group.
The results of the Taxon analyses are basically in accord with those of the
Principal Component. The dendogram of Fig. (A) also shows that the apterous
males of S. sacchari (17) and P. fraxini (29) are highly correlated. In Fig. (B),
where both species were excluded, it is apparent that the two brachypterous forms
of Gossyparia (27 and 28) are the most closely related (with a similarity coefficient
0-97), i.e. more than the macropterous (26) and the brachypterous (27) forms of
G. spuria. This again indicates the considerable differences apparently resulting
from the conditions of the wings. This dendogram shows that F. virgata (8) is far
removed from Planococcus group and is closer to Pseudococcus group. When a
phenon line was drawn at the correlation coefficient point of 0-45, the six groups of
genera suggested for Pseudococcidae were distinctly separated (except for the
position of F. virgata). It was also possible to draw another phenon line at 0-2
correlation coefficient, which roughly separates the two suggested major taxa or
sections, i.e. Planococcus, Pseudococcus and Saccharicoccus on the one hand, and
Ceroputo and Nairobia on the other (leaving Octococcus isolated). These dendo-
grams illustrate the most noteworthy conclusion. They indicate that Octococcus,
Ceroputo and Nairobia groups of genera have greater phenetic affinity with Erio-
coccidae than with Pseudococcidae. It was already suggested that N. bifrons (21)
ostensibly represents a link between the two families. But when a third programme
was run excluding this species, similar results were obtained (Fig. C). When the
state conditions of most characters were reduced so that almost all characters had
only 2-3 states, the results were still not much different (Fig. D). The dendogram
of Fig. (E) is only concerned with Pseudococcidae, where the relationships between
the species are amplified and where no substantial differences occurred.

DESCRIPTION OF SPECIES

The following descriptions, conforming with those of Ghauri (1962) and Giliomee

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 93

(1967) for Diaspididac and Coccidae respectively, contain all morphological details
which could be observed ; some of these might seem unnecessary, but it was done
deliberately since the evaluation of the importance of the various characters has
yet to be decisively determined.

PSEUDOCOCCIDAE

PLANOCOCCUS Group

PLANOCOCCUS Ferris, 1950

Planococcus citri (Risso)

(Text-figs. 4, 5)

The males of this species were described with varying degrees of accuracy by
Newstead (1903), Leonardi (1920), Green (1922), Makel (1942), Jancke (1955),
Theron (1958) and Beardsley (i960).

The winged forms only are known ; living individuals light brown in colour, with much
darker thorax and a blackish tinge on the wings. A narrow and slender species, moderately
long, with moderately long antennae and legs. When mounted, 966-1232 (ii2o)\t long,
266-308 (280) (j. wide at mesothorax and 2142-2380 (2282)^ wing expanse.

Body setae and derm pores : The antennae and the legs with numerous fleshy and few hair-
like setae of subequal lengths, about 31^ long on the antennae and 24a. long on the legs ; the
body itself with hair-like setae only, also about 24U. long. Quadrilocular and occasionally
trilocular and quinquelocular disc pores, about 6;x in diameter present on the head, the thorax
and the abdomen.

Head : Irregular tetrahedron ; widest across the genae, tapering anteriorly and ventrally ;
ventral preocular depression moderately pronounced. Length from apex to postoccipital ridge
113-128 (i22)u. ; from apex to neck 153-174 (i65)u. ; width across the genae 183-214 (198)0..
Dorsal arm of midcranial ridge anteriorly sepaiated from the other arms and reaching the
posterior margins of the dorsal eyes. Ventral arm anteriorly giving off the two lateral arms of
midcranial ridge, forming together a distinct Y-shaped ridge (Text-fig. 4,B). Postoccipital
ridge slender, U-shaped and with the anterior ends extending towards, but not reaching the
preocular ridges ; this ridge was described by Theron (1958) as two short meniscate ridges
but in the specimens examined, the ridge was found medially continuous. Dorsomedial part
of epicranium slightly raised. Preocular -j- interocular ridges strong, joining the postocular
ridge just below the ocellus ; a sclerotized rudiment of the preocular ridge below the articular
process, well marked. Genae membranous. Eyes : Corneae of the dorsal simple eyes 18-24
(2i)u in diameter and both separated by 92-104 (95)0. i.e. 3-8-5-6 (41) times their diameter
apart. Ventral simple eyes slightly larger, 24-31 (28) u in diameter and 24-43 (31)0. apart.
Lateral ocelli well developed. Cranial apophysis apically truncate ; tentorial bridge slender
and the anterior tentorial arms fused just before meeting the apophysis. Dorsal head setae
7-12 (9-3) on each side of the median line ; each gena always with 3 genal setae. Ventral head
setae present in 3 well defined groups on each side : a group of 2-3 (26) setae forming on both
sides an irregular longitudinal row between the ventral eyes ; a group of 6—10 (7-9) setae
forming with their partners of the other side a transverse band in front of the ventral eyes ;
a group of 2-4 (3-2) setae present anteriorly in a longitudinal row on each side of the ventral
arm of the midcranial ridge. Head pores : Two dorsal head pores always occur on each side
near the base of the antennal scape.

94 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Antennae : Filiform ; normally io-segmented but sometimes two or more adjacent segments
of the flagellum fused or incompletely separated ; 589-631 (616) (x long, i.e. somewhat longer
than half the body length (the ratio 1 : 1-6-1-9, av. i-8), and somewhat shorter than the hind
legs, the ratio 1 : 1-1-1-2 (1-16). Scape 37-43 (40)^ long and just as wide at the base ; with
4-7 (5-6) h.s. Pedicel 58-64 (61) [x long and 34-37 (35) fx wide ; with 19-26 (22-9) f.s., 3-10
(6-9) h.s. and a sensillum placodeum. Flagellum : Segment III club-shaped, about as long
as segment X, and both being longest of all (the ratio lengths of segments III to X 1 : 0-9-1-1,
av. i-o) ; the ratio width to length of segment III 1 : 3-3-4-1 (3-9). Segments IV to X
cylindrical and 21-24 (22) (A wide ; the ratio width to length of segment IX being 1 : 2-4-3-0
(2-8). In the following table are shown the lengths of segments III to X and the number of
setae on each:

Ill

IV

V

VI

VII

VIII

IX

X

lengths in [x

76-89

49-55

52-61

55-64

55-04

61-64

58-64

78-89

(av.)

(82)

(52)

(55)

(61)

(61)

(63)

(61)

(82)

f.s.

10-17

11-18

13-18

13-18

13-20

12-16

12-17

13-20

(av.)

(12-8)

(12-9)

(14-8)

(15-5)

(147)

(14-2)

(M-i)

(i6-i)

h.s.

2-5

i-3

i-3

i-3

1-2

1-2

1-2

1-2

(av.)

(3)

(i-8)

(2-2)

(i'9)

(i-6)

(17)

(1-4)

(2)

Antennal bristles easily distinguishable from the fleshy setae ; segments VIII and IX each
with a ventral bristle, about 43 [x long. Segment X with 3 preapical bristles, two lateral, also
about 43 jx long and one dorsal, slightly shorter ; this segment also with two capitate subapical
setae, about 40 jx long, and one apical hair-like seta.

Thorax : 426-540 (486) jx long. Prothorax : Pronolal ridges medially interrupted at a weak
point. Lateral pronotal sclerites and post-tergites small. Proepisternum + cervical sclerite
well developed, with the usual constriction which indicates the point of fusion between the
two component parts ; Theron (1958) called it propleuron -f- cervical sclerite, regarding it
as a ridge-like structure, but according to the writer's observations, the small episternum is
distinct and its dorsal margin only is ridge-like. It articulates posteriorly with the coxa by
means of a short propleural ridge, above which a small apophysis is invaginated. Prosternum
triangular, 31-46 (40) jx long; posteriorly bounded by a transverse, 70-92 (79) [x long pr asternal
ridge. Prothoracic setae on each side : Medial pronotal setae 1-2 (1-4) ; lateral pronotal setae
usually absent but one seta may occasionally occur on either side (av. 0-2). Antespiracular
dorsal setae 2—4 (2-9), and one antespiracular ventral seta always present. Prosternal setae usually
one and occasionally two (av. i-i). Prothoracic disc pores on each side : Medial pronotal pores
1-2 (1-2) ; lateral pronotal pores 3-6 (4-3) ; antespiracular dorsal pores 2-3 (2-2). Ventral
prosternal pores 0-2 (0-5).

Mesothorax : Prescutum subrectangular in dorsal view ; 70-82 (79) jx long and 107-119
(n6)(x wide (ratio 1 : 1-4-1-6, av. 1-5) ; laterally bounded by the prescutal ridge and posteriorly
by the prescutal suture. Scutum comparatively large, with the antero-lateral extremities
heavily sclerotized and with a narrow median longitudinal membranous area ; the scutum
107-116 (no)[x long, i.e. the ratio lengths of prescutum to scutum 1 : 1-3-1-5 (1-4). Prealare,
prealar ridge, triangular plate, tegula, anterior and posterior notal wing processes well developed.
Scutellum pentagonal, 49-61 (55)0. long and 89-104 (95)^ wide, the ratio being 1 : 1-6-1-9 (17)
and the ratio its length to the length of the scutum 1 : 1-8-2-3 (2-0). Postalare with well
separated anterior and posterior postalar ridges ; postnotal apophysis strong. Mesopleuron :
Mesopleural ridge interrupted at a short distance above the coxal articulation ; Theron (1958)
neither commented on this interruption nor showed it in his illustrations. Mesopleural
apophysis, mesopleural wing process, basalare and subepisternal ridge well developed ; subalare
small. The two parts of the episternum weakly sclerotized ; later opleurite thin ; mesepimeron
small. Mesosternum : Basisternum 134-143 (140) [x long and 156-183 (171) \l wide ; bounded
antero-laterally and posteriorly by the marginal and the precoxal ridges respectively ; furca
strong. Mesotkoracic spiracles 15-21 (i8)jx wide at opening, with 31-37 (34)^ long supporting

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

95

bar. Mesothoracic setae on each side : Prescutal setae 2-5 (33) ; scutal setae 9-12 (io-i) ;
scutellar setae 2-4 (3-1) ; tegular setae 1-3 (1-9). Postmesostigmatal setae absent medially and
occur laterally in two groups behind the spiracle, the dorso-lateral group comprising 1-3 (1-7)
setae, and the ventro-lateral with 4-6 (51) setae. The basisternal setae 9-16 (126). Meso-
thoracic disc pores : Mesospiracular pores 1-2 (1-3) behind each spiracle.

spl ahp dhs Irncr

Fig. 4. Planococcus citri (Risso), dorsal and ventral view.

56

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Metathorax : Metapostnotal sclerites medially connected by the transverse metapostnotal
ridge ; the latter, which appears to be of taxonomic importance, was overlooked by Theron
(1958). Metapleural ridge attenuates near the middle at a small pleural apophysis ; the
metapleural ridge dorsally merges into a small metapleural wing process. Metepisternum
dorsally supports the precoxal ridge of metathorax ; metepimeron irregularly triangular. Meta-
sternal apophysis, whose absence was emphasized by Theron (1958) and regarded as a specialized
feature of the lecanoid type of male (which includes P. cilri as a representative of Pseudo-
coccidae), is definitely present. Metathoracic spiracle identical with the mesothoracic. Meta-
thoracic setae on each side : A submedian group of 3 metatergal setae always present ; meta-
pleural setae usually absent, but one seta was found in one specimen (av. 01) ; anterior
metasternal setae 1-2 (1-9) forming, with their partners of the other side, a short median trans-
verse row ; posterior metasternal setae 1-2 (17) in a similar arrangement. Metathoracic pores :
Metaspiracular pores 1-2 (11) behind each spiracle, and one anterior metasternal pore may
occur on either side (av. 0-5) ; posterior metasternal pores absent.

Wings : 938-1050 (1008) jjl long and 378-476 (420)0. wide ; usually with 3 alar setae (4 setae
observed in one occasion, av. 31); a compact row of 2-3 (2-2) minute circular sensoria also
apparent. Hamulohalterae 70-76 (74) (x long and about 15U, wide; with a weak ridge and an
apical, 61-67 (64) [i l° n g hooked seta, i.e. the seta slightly shorter than the hamulohaltera, the
ratio their lengths being 1 : 11-1-2 (1*15).

Legs : Comparatively long and moderately stout ; the ratio length of the hind leg to the
total body length 1 : 1-3-1-7 (i-6). Coxa and trochanter about 46 and 24(1 wide ; the proximal
part of the latter longer than the distal and with 3 circular sensilla on each side. Femur about
37 [/. wide ; that of the fore leg shortest and that of the hind leg longest ; the ratio width to
length of the hind femur 1 : 5-2-5-8 (5-4). Tibia 2iu. wide ; with 2 strong apical spurs and
2-4 smaller spines ; in front leg, the femur shorter than the tibia, the ratio their lengths
1 : I-I-I-2 (113). The distal segment of the tarsus about 2I[a wide ; tarsal digitules apically
knobbed, 34-40 (37) (x long. Claw gradually tapering ; ungual digitules extremely fine, about
I5[x long. The following table shows the lengths of the leg segments in microns and the
number of fleshy and hairlike setae on each :

200 -~

100.

Fig. 5. Planococcus citri (Risso), lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

97

Coxa

length

f.s.

h.s.

Fore Leg

37-46 (40)
10-18 (13-2)
5-9 (6-8)

Middle Leg
40-49 (43)
12-19 (15-1)
5-10 (6-2)

Hind Leg

46-52 (49)
13-22 (173)
6-1 1 (8-8)

Trochanter

length

f.s.

h.s.

55-61 (59)
3-8 (4-3)
2-4 (2-9)

55-61 (60)
3-9 (5-9)
2-4 (3-o)

58-64 (62)
4-9 (6-4)
2-5 (3-2)

Femur

length

f.s.

h.s.

177-183 (180)

22-31 (269)

4-7 (5-o)

183-198 (189)

25-36 (28-6)

4-8 (5-9)

192-21 1 (198)

32-45 (364)

5-9 (6-9)

Tibia

length

f.s.

h.s.

201-21 1 (204)

36-47 (4 1 -6)

5-9 (6-8)

217-232 (223)

43-54 (47 6 )

6-9 (73)

256-278 (265)

52-64 (58-3)

5-9 (7-i)

Tarsus
Claw

length
f.s.
h.s.
length

89-95 (92)
21-27 (237)

2-5 (31)
34-4° (37)

89-98 (95)
22-34 ( 20 '5)

3-6 (4-2)
34-40 (37) '

101-107 (104)
24-37 (288)

3-7 (4-6)
34-40 (37)

Total length

of leg

598-619 (607)

628-662 (641)

695-735 (7M)

Abdomen : 304-494 (395) u. long and 220-289 (258) u. wide. The tergites of segments I and II
small ; that of segment VIII large and that of segments IX -f- X distinct. Sternites of segment
VIII weakly sclerotized. Ostioles well developed, 27-34 (30) u. long at orifice. Abdominal
setae on each side : Dorsal setae 2-3 on segments I and VII, 3-4 on segments II to VI, and
always 2 on segment VIII. Pleural setae 3-5 on segments I to VII, and 2-3 on segment VIII,
in addition one conspicuously longer seta. Ventral setae 1-2 on segment II, 2-3 on segments
III to VII, and absent on segment VIII. Abdominal disc pores only in lateral groups on each
side ; segments I to VII with n-18 (141), 2-4 (31), 2-4 (27), 2-4 (27), 2-4 (23), 3-6 (40)
and 5-8 (6-2) pores, respectively.

Glandular pouches well developed ; setae of glandular pouch include a pair of long tail setae
299-342 (323) \l long and one seta, 70-104 (82) u. long, i.e. the ratio lengths of the tail setae to
the length of the body 1 : 3-0-37 (35).

Genital segment comparatively small ; conical in dorsal view, with a short and narrow style,
which is curved upwards in lateral view. Penial sheath 107-113 (109)11 long and 73-82 (76)11
wide (the ratio 1-3-1-5, av. 14 : 1, and the ratio its length to the total body length 1 : 8-8-11-4,
av. 10-2). Basal ridge of penial sheath with a small projection. Process of penial sheath well
pronounced and sclerotized, with 3-4 minute setal sensilla. Aedeagus relatively short, dorso-
posteriorly curving from its basal rod towards its pointed apex ; internal genital aperture
distinct. Setae of genital segment : Dorsally, 3 setae always present on each side near the base
of the style ; ventrally a group of 3-5 (4-1) setae occur on each side of the penial sheath.

Material : 10 specimens studied, collected by M. Kosztarab, on Coleus sp. (in
greenhouses), in Ohio State University, Columbus, Ohio, U.S.A., 6.V.1961.

Other material : The writer also obtained adult males of this species from the
following sources :

On Ivy (Hedera helix), in Week's Hall, one of Imperial College residential buildings,
South Kensington, during the second week of June, 1963.

98 MORPHOLOGY AND TAXONOMY OF ADULT MALES

On Solatium sp. and on Rhapes sp., in the gardens of Manial Palace, Cairo, Egypt,
U.A.R., during the second and third weeks of August, 1964.

On Croton sp., in the Botanic gardens of Zohreya (greenhouses), Cairo, Egypt,
U.A.R., also during the second week of August, 1964.

On potato sprouts received from Dr. D. J. Williams, 30.1.1965. The culture
was sent from Angola, West Africa where the insect was reported causing tremendous
damage to potatoes. Dr. Williams, although he identified the females as P. citri,
was somewhat doubtful ; the pleural prothoracic group of ducts, normally found
anterior to, and around the spiracles in this species, were absent. When the males
were mounted and examined, they completely agreed with the typical specimens.

Planococcus kenyae (Le Pelley)
(Text-fig. 6)

The macropterous forms only are known ; narrow and slender, of medium size, with
comparatively short antennae and long legs. Mounted specimens 1092-1204 (1162)0. long,
252-294 (280)0. wide at mesothorax and 2352-2576 (2478)0 wing expanse.

Body setae and pores : Antennae and legs with numerous fleshy and a few hair-like setae,
about 310 long on the former and 240 on the latter ; the body itself with hair-like setae only,
about 240 long. Quadrilocular and sometimes trilocular or quinquelocular disc pores, about
60 in diameter, present on the head, thorax and abdomen.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression well pronounced in lateral view. Length from apex to postoccipital ridge 113-134
(122)0 ; from apex to neck 153-174 (168)0 ; width across the genae 180-207 ( I 95)( jl - Dorsal
arm of the midcranial ridge anteriorly detached from other arms, and almost reaching the
posterior level of the dorsal eyes. Ventral and lateral arms forming together a Y-shaped ridge.
Postoccipital ridge U-shaped, not reaching the preocular ridges anteriorly. Dorsomedial part
of epicranium slightly raised. Preocular + interocular ridges well developed, posteriorly fused
with postocular ridge below the ocellus. A distinct ventral rudiment of the preocular ridge
present below the articular process. Preoral ridge slender. Eyes : Dorsal simple eyes not
projecting beyond the outer margins of the head ; their corneae 21-27 ( 2 4) a m diameter, and
both separated by 85-98 (92)0, i.e. 3-5-4-0 (3-8) their diameter apart. Ventral simple eyes
larger and much closer, 27-34 (31)0 in diameter and 18-24 (21)0 apart. Lateral ocelli well
developed. Ocular sclerite traversed by the interocular ridge. Cranial apophysis apically
truncate. Tentorial bridge slender. Dorsal head setae 9-12 (10-2) on each side anterior to the
postoccipital ridge ; each gena usually with 3 and occasionally 2 (av. 2-8) genal setae. Ventral
head setae on each side : 2-4 (3-4) between the ventral eyes ; 7-10 (8-6) in the area of the
ventral preocular depression, forming with their partners of the other side a transverse band ;
anteriorly, 3-4 (3-3) in a longitudinal row on each side of the ventral arm of the midcranial
ridge. Head pores : Usually 2 pores, and occasionally 3 or 4 (av. 2-3), present dorsally near
the base of each antenna.

Antennae : Filiform ; 10-segmented ; 509-641 (577)0 long, i.e. about half the body length,
the ratio being 1 : 1-8-2-2 (2-0), and slightly shorter than the hind leg, the ratio 1 : i-i— 1-3
(1-2). Scape 37-43 (40)0 long and 34-40 (37)0 wide at base ; with 5-7 (5-9) h.s. Pedicel
55-64 (60)0 long and 31-37 (34)^ wide at widest ; with 12-23 (17-3) f.s., 6-n (89) h.s. and a
sensillum placodeum. Flagellum : Segment III club-shaped, longest of all (the ratio its length
to the length of segment X 1-1-1-3, av. 1-2 : 1) and about 210 wide (the ratio its width to its
length i : 3-8-5-2, av. 4-3). Segments IV to X cylindrical and 18-21 (20)0 wide (the ratio

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 99

width to length of segment IX 1 : 2-4-2-9, av. 2-5). The lengths of the flagellar segments and
the number of setae on each as follows:

III IV V VI VII VIII IX X

length in ^ 73-98 40-58 46-58 46-58 46-58 49-64 46-61 67-82

(av.) (85) (52) (52) (53) (53) (55) (52) (73)

f.s. 5-12 6-1 1 8-15 9-17 8-16 9-18 9-14 10-18

(av.) (87) (8-9) (ll-a) (13-1) (12-4) (12-9) (n-8) (137)

h.s. 3-6 1-3 1-3 1-4 1-4 1-4 1-3 1-4

(av.) (4-1) (2-1) (2-3) (1-9) (2-1) (2-2) (1-9) (2-3)

Antennal bristles much stouter than the fleshy setae ; segments VIII and IX each with a
ventral bristle ; segment X with two preapical lateral bristles and a slightly shorter dorsal one,
at a greater distance from the apex. Terminal segment also with two subapical sensory setae
and one apical hair-like seta.

Thorax : 470-555 (524)0. long. Prothorax : Pronotal ridges medially interrupted by a weak
sclerotization ; lateral pronotal sclerites moderately large and well sclerotized. Post-tergites
narrow. Proepisternum with a ridge-like dorsal margin. Prosternum triangular, 31-43 (37) u.
long ; prosternal ridge 79-95 (85) u. long. Prothoracic setae on each side of the median line as
follows : One medial pronotal seta usually present, but sometimes absent on either or both
sides (av. o-8) ; lateral pronotal and post-tergital setae absent. Antespiracular dorsal setae
1-3 (2-1) ; antespiracular ventral setae and prosternal seta always one. Prothoracic disc pores
on each side : Medial pronotal pores 2-4 (3-2) ; lateral pronotal pores 2-6 (41) ; antespiracular
dorsal pores 2-5 (3-4). Prosternal pores usually 1-2, but sometimes 3 pores may occur on either
side or the pores may be absent altogether (av. i-6).

Mesothorax : Prescutum subrectangular in dorsal view ; 76-89 (82)0. long and 104-122
(119) [i wide, the ratio being 1 : 1-3-1-6 (1-4) ; prescutal ridges well developed and prescutal
suture distinct. Scutum heavily sclerotized antero-laterally and with a median longitudinal
narrow membranous area ; scutum 95-116 (104)0. long, i.e. the ratio length of prescutum to
scutum 1 : 1-1-1-4 ( I- 3)- Prealare, prealar ridge, triangular plate, tegula, anterior and posterior
notal wing processes well developed. Sciitelhun pentagonal, 46-58 (52) u. long and 82-101
(92)0 wide, i.e. the ratio 1 : 1-7-1-9 (i-8) and the ratio its length to the length of scutum
1 : 1-9-2-2 (2-0). Anterior and posterior ridges of postalare well separated ; postnotal apophysis
strong. Mesopleuron : mesopleural ridge interrupted above the coxal articulation ; meso-
pleural apophysis, mesopleural wing process, basalare and subepisternal ridge well developed.
Mesepisternum, mesepimeron well-defined, and later opleurite narrow. Mesosternum : Basi-
sternum 128-146 (140)0, long and 156-183 (i7i)(x wide ; marginal and precoxal ridges well
developed ; furca strong. Mesothoracic spiracle 15-21 (18) u. wide at opening, with a 34-40
(37) (i. long supporting bar. Mesothoracic setae on each side : Prescutal setae 4-6 (4-6) ; scutal
setae 6-9 (7-5) ; scutellar setae 2-3 (2-5). Tegular setae 2-3 (2-1) ; postmesostigmatal setae
4-7 (5-1), occurring laterally behind each spiracle. Basisternal setae 9-14 (11-4). Mesothoracic
pores : 1-2 mesospiracular pores usually present behind each spiracle, but sometimes entirely
absent (av. 09).

Metathorax : Metapostnotal sclerites and metapostnotal ridge distinct. Metapleural ridge
attenuating near the middle at the position of the metapleural apophysis ; metapleural wing
process small. Precoxal ridge of metathorax well developed ; metepisternum , metepimeron and
metasternal apophysis well defined. Metathoracic spiracle similar to the mesothoracic. Meta-
thoracic setae on each side : Metatergal setae 2-3 (2-7) ; metapleural setae absent ; anterior
metasternal setae 1-3 (2-3) ; posterior metasternal setae 0-2 (0-4). Metathoracic disc pores on
each side : Metathoracic pores 0-1 (0-3) ; anterior metasternal pores 0-2 (1-2).

Wings : 1064-1162 (ii2o)[a long and 406-462 (434)0- wide ; usually with 4 alar setae and
occasionally 3 or 5 (av. 3-9) ; a compact row of 2-3 (2-3) minute circular sensoria also present.

ioo MORPHOLOGY AND TAXONOMY OF ADULT MALES

Axillary and additional sclerites distinct. Hamnlohalterae 67-89 (73)^1 long and 15-18 (16) (x
wide ; with a slender ridge and a 61-67 (64) pi long, apically hooked seta, i.e. the ratio length
of seta to the length of the hamulohaltera 1 : 1—1-2 (i-i).

Fig. 6. Planococcus kenyae (Le Pelley), dorsal, ventral and lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 101

Legs : Comparatively long and moderately stout ; the ratio length of the hind legs to the
total body length i : 1-5-1 -8 (1-7). Coxa and trochanter about 46 and 24^ wide respectively.
Femur about 37 u. wide ; that of the middle leg shortest and that of the hind leg longest ;
the ratio width to length of the hind femur 1 : 5-5-6-3 (5-7). Tibia about 21 [i wide ; with
2 apical strong spurs and 2-4 smaller spines ; in the fore legs the femur is slightly shorter than
the tibia, the ratio their lengths 1 : 1-04-1-2 (i-i). Tarsus about 21 u. wide ; tarsal digitules
apically knobbed, 37-43 (40)^ long. Claw gradually tapering towards a pointed end ; ungual
digitules with acute tips. In the following table, the lengths of the leg segments (in microns)
and the number of setae on each are given :

Fore leg

Middle leg

Hind leg

Coxa

length

f.s.

h.s.

40-46 (44)
13-19 (156)
4-8 (6-2)

43-49 (46)
14-19 (16-2)
5-8 (6-4)

46-52 (48)
15-21 (17-8)
5-10 (7-8)

Trochanter

length

f.s.

h.s.

58-64 (62)
5-8 (6-6)
2-4 (3-2)

58-64 (62)
5-9 (62)
2-4 (3-o)

61-70 (65)
6-9 (7-6)
2-4 (3'2)

Femur

length

f.s.

h.s.

174-189 (183)

24-32 (27-6)

4-8 (5-8)

171-186 (177)

24-30 (26-4)

4-8 (5-2)

183-211 (192)

31-43 (37-2)

4-9 (6-8)

Tibia

length

f.s.

h.s.

180-214 (195)

33-46 (406)

5-8 (6-6)

192-223 (207)

39-51 (454)

5-9 (7'2)

238-265 (253)

47-58 (53-6)

6-9 (7-4)

Tarsus

length

f.s.

h.s.

85-92 (88)
20-26 (23-2)
3-5 (3-6)

85-92 (89)
21-28 (24-8)
3-6 (4-4)

92-101 (95)

24-31 (27-4)

3-6 (4-8)

Claw

length

34-40 (37)

34-4° (37)

34-40 (37)

Total length

of leg

577-634 (604)

586-637 (613)

665-738 (692)

Abdomen : 388-441 (418)0. long and 236-274 (251)11 wide. Tergites of segments I and II
small and narrow ; those of segments VIII and IX + X distinct. Sternites of segment VIII
weak. Ostioles well developed, 31-37 (34) \i long at orifice. Abdominal setae on each side : Dorsal
setae 2-3 on segments I to VII and always 2 on segment VIII. Pleural setae 3-5 on segments
I to VII, and always 3 on segment VIII, including one slightly longer seta. Ventral setae
always 1 on segment II, usually 2 and sometimes 3 on segments III to VII, and absent on
segment VIII. Abdominal disc pores : Segments I to VII with 9-16 (12-2), 3-5 (37), 2-4
(2-9), 2-5 (31), 2-4 (2-9), 3-4 (3-4), and 3-6 (46) pleural pores respectively ; segment VI also
with 0-1 (04) and segment VII with 0-2 (i-i) ventral pores.

Glandular pouches well developed ; setae of glandular pouch consist of a pair of 336-366
(35i)u, long tail setae, and one seta of medium length (70-92, av. 79(1 long) ; i.e. the ratio
length of the tail setae to the total body length 1 : 3-1-3-4 (3-3).

Genital segment small ; triangular in dorsal view, with a short narrow style ; the latter
curving upwards in lateral view. Penial sheath 98-110 {io^)\j. long and 70-82 (76) u. wide, i.e.
the ratio length to width 1-2-1-5 ( I- 4) : *• an d tne ratio its length to the total body length
1 : 10-1-12-5 ( II-2 )- Basal ridge of penial sheath with a small projection. Process of penial
sheath well pronounced and usually with 3-4 minute setal sensilla. Aedeagus relatively long
and broad, with a wide internal genital aperture. Setae of the genital segment : Dorsally, 3 setae
always present on each side near the base of the style ; ventrally 3-4 (3-5) slightly smaller
setae occur on each side of the penial sheath.

102 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Material : 10 specimens examined, collected by T. J. Crowe, on Coffea arabica,
in Ruiru, Kenya, 25. ix. 1962.

Remarks : This species, although closely related to P. citri, could be separated
by having : the third antennal segment longer than the terminal ; the femur of
the middle leg shortest ; a ventral pore, at least on one side of abdominal segment
VII ; a comparatively stouter aedeagus.

Planococcus dioscoreae Williams
(Text-figs. 7, 8)

Only the winged forms known ; living specimens not available. The mounted males narrow
and slender, of medium or large size, with short antennae and moderately long legs. The
total length of the body 1246-1400 (1330) fx, the width at mesothorax 280-308 (290) [x and the
wing span 1960-2128 (2044) u..

Body setae and pores : Numerous fleshy and few hair-like setae occur on the antennae and
the legs ; the body itself with hair-like setea only. All the setae about 21 [i long. Quadri-
locular and few quinquelocular disc pores present on the head, thorax and abdomen, about
6[x in diameter.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression well pronounced in lateral view. Length from apex to postoccipital ridge 116-122
(120) [x ; from apex to neck 168-183 ( I 74)t i '• width across the genae 198-229 (207) ;x. Dorsal
arm of midcranial ridge weak, anteriorly detached from the other arms and posteriorly reaching
the hindmost level of the dorsal eyes. Ventral and lateral arms of midcranial ridge forming
together a Y-shaped ridge. Postoccipital ridge (por) slender, anteriorly continuous with the
preocular ridges ; the postoccipital ridge binds the dorsomedial part of epicranium (dmep)
posteriorly. Preocular and interocular ridges intimately joined to the postocular ridge ; ventral
rudiment of the preocular ridge well marked by a short sclerotization just below the articular
process. Preoral ridge slender. Genae large and membranous. Eyes : Dorsal simple eyes
not projecting beyond the outer margins of the head in dorsal view ; their corneae 21-27 ( 2 4)( JL
in diameter and separated by 98-107 (104) jx, i.e. 39— 46 (4-3) times as much as the corneae
apart. Ventral simple eyes slightly larger and much closer, 27-31 (29) y. in diameter and
21-27 ( 2 4)[ JL apart. Lateral ocelli comparatively large and dorsally supported by a slender
ocellar ridge. Ocular sclerites traversed by the interocular ridges. Cranial apophysis apically
truncate. Tentorial bridge slender. Dorsal head setae 8-12 (9-7) on each side of the median
line ; each gena with 2-3 (2-7) genal setae. Ventral head setae on each side : 3-4 (3-4) between
the ventral eyes ; a group of 9-1 1 (10-4) setae forming with their partners of the other side a
transverse band in the area of the ventral preocular depression ; 2-3 (2-4) setae on each side
of the ventral arm of the midcranial ridge. Head disc pores : Dorsally, 5-8 (6-4) pores present
on each side near the base of the antennal scape ; ventral pores absent. Antennae : Filiform ;
normally 10-segmented ; 616-665 (641) n long, i.e. about as long as half the body length and
slightly shorter than the hind legs, the ratios being 1 : 2-0-2-1 (2-05) and 1 : i-i respectively.
Scape 43-46 (45) (x long and 43-49 (46) y. wide at base ; with 4-7 (5-0) h.s. Pedicel 61-67 (63) y
long and 34-37 (35) u. wide at widest ; with 15-21 (18-4) f.s., 5-12 (9-3) h.s., and a distal
sensillum placodeum. Flagellum : Segment III club-shaped, being the longest of all, and
about 25 [x wide ; the ratio lengths of segments III to X i-i : 1 and the ratio width to length
of segment III 1 : 3-6-3-7 (3-65). Segments IV to X cylindrical and about 23(x wide ;
segments IV-IX subequal in length, and the terminal segment slightly longer ; the ratio
width to length of segment IX 1 : 2-7-2-9 (2-8).

The following table shows the lengths of the flagellar segments and the number of setae
on each :

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 103

III IV V VI VII VIII IX X

length in (x

79-89

61-64

55-6i

58-64

61-64

61-67

61-64

73-79

(av.)

(85)

(62)

(58)

(61)

(62)

(64)

(62)

(76)

f.s.

11-16

12-14

11-16

12-14

15-16

13-15

14-16

13-16

(av.)

(13-5)

(13-0)

(13-5)

(13-0)

(15-5)

(140)

(i5-o)

(M-5)

h.s.

3-4

2-4

2-3

2-3

2-3

3-4

3-4

4-5

(av.)

(3-5)

(3-o)

(2-5)

(2-5)

(2-5)

(3-5)

(3-5)

(4-5)

Antennal bristles : Segments VIII and IX each with a ventral antennal bristle, about 34(x
long ; segment X with 3 similar, preapical bristles, the dorsal of which is slightly shorter and
at a greater distance from the apex. Terminal segment also with two capitate sabapical
sensory setae, about 37 ;x long, and one apical hair-like seta.

Thorax : 509-547 (524)^ long. Prothorax : Pronotal ridges interrupted in the middle ;
lateral pronotal sclerites and post-tergites small. Proepisternum with ridge-like dorsal margin ;
propleural ridge short and propleural apophysis small. Prosternal ridge double barred, 76-92
(85) (x long. Prothoracic setae on each side of the median line : One medial pronotal seta usually
present (av. 09) ; one lateral pronotal seta also may be present or absent (av. 08) ; post-
tergital setae absent. Antespiracular dorsal setae 1-2 (i-8) h.s. One antespiracular ventral seta
and one prosternal seta always present. Prothoracic disc pores on each side : Medial pronotal
pores 5-10 (8-4) ; lateral pronotal pores 3-4 (3-8). Antespiracular dorsal pores 5-9 (7-4) ;
post-tergital pores 2-6 (4-4). Ventrally, 1-2 (i-8) prosternal pores occur medially.

Mesothorax : Prescutum subrectangular in dorsal view : 82-85 (84) H long and 122-143
(134) u. wide, i.e. the ratio length to width 1 : 1-4-1-7 (1 : i-6). Prescutal ridge strong and
prescutal suture distinct. Scutum with heavily sclerotized anterolateral extremities and with
a longitudinal median narrow membranous area. Scutum 98-116 (104)^ long, i.e. the ratio
lengths of prescutum to scutum 1 : 1-2-1-4 ( l '■ 1*22). Prealare, prealar ridge, triangular
plate, tegula, anterior and posterior notal wing processes well developed. Sattellum pentagonal,
58-67 (6i)(jl long and 98-116 (104)0. wide, the ratio length to width 1 : 1-6-1-7 (168), and the
ratio its length to the length of scutum 1 : 1-6-1-7 ( I- 67). Postalare with well separated
anterior and posterior postalar ridges ; postnotal apophysis strong. Mesopleuron : Mesopleural
ridge interrupted above the coxal articulation ; mesopleural apophysis, mesopleural wing
process and basalare well developed. The mesepisteruttin anteriorly bounded by the sub-
episternal ridge ; lateropleurite narrow ; mesepimeron small and heavily sclerotized. Meso-
slernum : Basisternum 137-156 (143)0. long and 180-204 ( J 89)|x wide ; the marginal and
precoxal ridges, as well as the furca strongly developed. Mesothoracic spiracles 18-21 (20) u,
wide at opening, with 31-40 (35) [x. long supporting bar. Mesothoracic setae on each side of the
median line : Prescutal setae 3-4 (3-8) ; scutal setae 8-10 (9-4) ; scutellar setae 2-4 (2-8).
Tegular setae 2-3 (2-6) ; postmesostigmatal setae 4-5 (4-4) in a lateral group behind each spiracle.
Basisternal setae 10-13 ( II- 8). Mesothoracic disc pores : 2-4 (3-0) mesospiracular pores present
behind each spiracle ; postmesostigmatal pores absent.

Metathorax : Metapostnotal sclerites well developed and metapostnotal ridge distinct. Meta-
pleural ridge provides a small metapleural apophysis near the middle ; metapleural wing process,
small. Precoxal ridge, metepisternum, metepimeron and metasternal apophysis well developed.
Metathoracic spiracle identical with that of the mesothorax. Metathoracic setae on each side :
Metatergal setae 3-5 (4-0), in a sublateral group ; metapleural setae absent. Anterior meta-
sternal setae 4-6 (5-0) ; posterior metasternal setae 0-2 (i-o). Metathoracic pores on each side :
Metaspiracular pores 2-3 (2-4) ; anterior metasternal pores 3-4 (34) ; and posterior metasternal
pores also 3-4 (3-5).

Wings : 854-924 (896) jx long and 308-364 (336) u, wide ; axillary and additional sclerites
well developed. 3 alar setae and 2 circular sensoria always present. Hamulohalterae 67-73
(70) [x long and 15-18 (17) [x wide ; with a slender ridge and an apically hooked seta, 40-46
(43) {x long (the ratio its length to the length of the hamulohaltera 1 : 1-6-1-7 ( I- 64)-

104

MORPHOLOGY AND TAXONOMY OF ADULT MALES

dhp dhs

dmcr ; / vhs

pocr

-vse

h '

L_ procr

E?_ - - ca

/

A pocr

/ — - P rnr

r _ pepcv

p \

^— E

^!S3-- CX '

L<; 1 ^asvs

1 ^

stn,«

/-' „-stn,r

^

-- ^ s PjP

Aj? \^ ' ^ P m s

'*yi " " 5Cr

' /t~ - €ps i

V /A -- plr i

In,

\?t- ~~ ~~ pa

>»

^cK^" CXl

pcf

(/j3D ^sp 3

)

^'K-' 5P3P

• i _ — amss

J-

"/jl _ — pla 3

«t

~"C plr 3

sla

M~ ~ ~ epm 3

Fig. 7. Planococcus dioscoreae Williams, dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

105

Legs : Comparatively long and moderately stout ; the ratio length of the hind leg to the
total length of the body about 1 : 19. Coxa about 49 and trochanter about 3051 wide. Femur
about 43 [x wide ; that of the fore leg shortest and that of the hind leg longest ; the ratio width
to length of hind femur 1 : 4-5-5-2 (4-9). Tibia about 24;x wide ; with 2 apical stout spurs and
2-4 smaller spines ; the ratio length of femur to tibia in fore leg being 1 : i-i. Tarsus about
21 (x wide ; tarsal digilules apically knobbed, about 34^ long. Claw gradually tapering to a
sharp point ; ungual digilules extremely fine. The lengths of the leg segments and the number
of setae on each are given in the following table, in microns :

Fore leg

Middle leg

Hind leg

Coxa

length

f.s.

h.s.

40-46 (43)
8-12 (100)
4-6 (5-o)

40-46 (43)
9-11 (100)
5-7 (6-o)

43-49 (46)
10-13 (n-5)

5-8 (6-5)

Trochanter

length

f.s.

h.s.

64-73 (70)
3-4 (3-5)
0-1 (0-5)

64-73 (70)
2-4 (3-o)
1-2 (i-5)

67-76 (73)
3-4 (3-5)
1-2 (1-5)

Femur

length

f.s.

h.s.

168-189 (180)

16-23 (195)

7-10 (8-5)

171-189 (180)

19-25 (22) .

9-1 1 (100)

180-207 (195)
22-32 (27)
7-10 (8-5)

Tibia

length

f.s.

h.s.

183-204 (195)

32-38 (35)

6-9 (7-5)

198-217 (207)

33-40 (365)

6-10 (8-o)

250-268 (259)

46-51 (48-5)

7-1 1 (9-0)

Tarsus

length

f.s.

h.s.

76-85 (82)
9-12 (105)
8-10 (90)

79-85 (82)
9-1 1 (100)
7-10 (85)

92-98 (95)
11-14 (125)
9-12 (105)

Claw

length

27-31 (29)

27-31 (29)

27-31 (29)

Total length of leg

558-628 (592)

580-641 (610)

659-729 (695)

Abdomen : 471-532 (502) jx long and 274-319 (296) [x wide. Tergites of segments I and II
small ; that of segment VIII large and that of segments 9+10 distinct. Sternites of segment
VIII small and weak. Ostioles prominent and well developed, 37-46 (40) jx long at orifice.
Abdominal setae on each side : Dorsal setae on segments I to VIII usually 2-3 and occasionally
4. Pleural setae 2-4, including one slightly longer seta on segment VIII. Ventral setae 1

I ,' I 1 1 1 \ \ \ \ 1 \ 1 I \ \ \ \ \

Q I " f I S f * - i * u S" e ' e e =

w «n i v n a. Ol

200^

Fig. 8. Planococcus dioscoreae Williams, lateral view.

106 MORPHOLOGY AND TAXONOMY OF ADULT MALES

on segment II and 2 on segments III to VIII. Abdominal disc pores : Segment I with 10-17
(13-5), and segments II to VII with 3-5 pleural pores ; dorsal and ventral abdominal pores
absent.

Glandular pouch well developed ; setae of glandular pouch consist of the two long tail setae,
305-320 (314) [i long, and one much shorter seta, 55-92 (73) \l long ; the ratio length of the tail
setae to the total length of the body 1 : 4-1-4-4 (1 : 4-2).

Genital Segment : Comparatively small : triangular in dorsal view ; style short, curving
upwards in lateral view. Penial sheath 122-128 (125)11 long and about 76(1 wide, i.e. the ratios
its length to its width about i-6 : 1, and its length to the total body length 1 : 10-2-10-9
(1 : io-6). Basal ridge of penial sheath well developed, with a small projection. Process of
penial sheath well pronounced, with 3-4 (3-6) minute setal sensilla. Aedeagus gradually tapering
to a sharply pointed tip. Setae of genital segment on each side : Dorsally, 3 setae always
present near the base of the style ; ventrally 4-5 (4-5) setae occur on the penial sheath.

Material examined : 3 specimens only of this species were available for study,
collected by H. Standfast, on Yam (Dioscorea sp.), in Sepik district, Tenteguna,
New Guinea, on 23. vi. 1959 (received from D. J. Williams).

PLANOCOCCOIDES Ezzart & McConnell, 1956

Planococcoides ireneus De Lotto

(Text-figs. 9, 10)

Winged forms only known ; living specimens not available. The males narrow and slender,
of medium size, with comparatively short antennae and moderately long legs. When mounted,
total body length 1008-1414 (1162) [i, width at mesothorax 280-350 (322)11 and wing expanse
1932-2422 (2114)11.

Body setae and pores : The appendages clothed with many fleshy and few hair-like setae,
about 39p. long ; the body itself with hair-like setae only, about 24 [x long. Numerous
quadrilocular and occasionally quinquelocular disc pores, about 90. in diameter, occur on the
head, the thorax and the abdomen.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression hardly indicated in lateral view. Length from apex to postoccipital ridge 98-122
(113)11 ; from apex to neck 153-180 (165) y. ; width across genae 174-214 (192)11. Dorsal
arm of midcranial ridge slender, anteriorly detached from other arms and posteriorly reaching
the posterior level of the dorsal eyes. Ventral and lateral arms of midcranial ridge well
developed, forming together a Y-shaped ridge. Postoccipital ridge anteriorly separated from
the preocular ridges. Dorsomedial part of epicranium (dmep) slightly raised. Preocular +
interocular ridges meet the postocular ridge below the ocellus ; ventral rudiment of the former
entirely absent. Preoral ridge slender. Genae large. Eyes : Dorsal simple eyes not projecting
beyond the outer margins of the head in dorsal view ; the diameter of their corneae 21-27
(24)11, and separated by 79-101 (89)11, i.e. 3-1-4-7 (3-6) times as much as their corneae apart.
Ventral simple eyes 24-31 (29)11 in diameter and 18-40 (25)11 apart. Lateral ocelli well
developed. Ocular sclerite traversed by the interocular ridge. Cranial apophysis truncate ;
tentorial bridge slender. Dorsal head setae 8-1 1 (9-9) on each side of the median line ; each
gena always with 3 genal setae. Ventral head setae on each side : 3-5 (3-9) between the ventral
eyes ; 7-1 1 (7-7) in the area of the ventral preocular depression, forming a transverse band
continuous with those of the other side ; 3-5 (4-3) anteriorly on each side of the ventral arm
of the midcranial ridge. Head disc pores : 2-3 (2-4) dorsal pores present near the base of each
antenna.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

107

Antennae : Filiform ; normally 10-segmented but fusion between adjacent segments very
common ; 494-580 (545) (x long, i.e. equal or slightly shorter than half the body length, the
ratio 1 : 2-0-2-5 f 1 : 2-1 ) ; somewhat shorter than the hind leg, the ratio 1 : 1-1-1-3 (1 : 1-2).
Scape 31-40 (37) (i long and 40-46 (42) u. wide at base ; always with 5 hair-like setae. Pedicel
5 2 -55 (54)^ l° n g an d 31-37 (34)^ wide ; with 12-77 ( I 5' 1 ) f- s -. 5~ 10 (6-8) h.s. and a sensillum
placodeum. Flagellum : Segment III club-shaped, usually the longest (sometimes equal in

drip dmcr
1 I

n H — a5 *

-h

\'J — \ ~ ~ DrDS

/N l

1/ ~~ -

/ X N ""-Sis

'

f N N pro

\

St

aed

Fig. 9. Planococcus ireneus De Lotto, dorsal and ventral view.

io8

MORPHOLOGY AND TAXONOMY OF ADULT MALES

length to terminal segment), the ratio length of segments III to X being 10— ii (1-05) : 1 ;
segment III about 25^ wide, the ratio its width to its length being 1 : 2 6-3 3 (3-1). Segments
IV-X cylindrical and about 23^1 wide ; the ratio width to length of segment IX 1 : 2-9-3-6
(3-3). In the following table the length of the flagellar segments and the number of setae on
each are given :

Ill

IV

V

VI

VII

VIII

IX

X

length in ;x

70-82

43-52

43-52

46-58

46-58

49-58

49-58

70-79

(av.)

(76)

(49)

(49)

(52)

(52)

(55)

(55)

(73)

f.s.

12-15

8-14

9-14

11-17

13-17

12-18

13-16

14-20

(av.)

(H-i)

("•3)

(I2"5)

(147)

(15-4)

(14-8)

(147)

(i8-3)

h.s.

2-5

i-3

0-2

0-2

O-I

O-I

O-I

0-2

(av.)

(3-9)

(2-0)

(I-I)

(07)

(o-4)

(07)

(07)

(o-8)

Antennal bristles : Each of antennal segments VIII and IX with one ventral bristle.
Terminal segment with 3 preapical bristles the dorsal of which is slightly shorter ; terminal
segment also with two capitate subapical sensory setae and one apical hair-like seta.

Thorax : 395-578 (464) pi long. Prothorax : Pronotal ridges medially interrupted ; lateral
pronotal sclerites small. Post-tergites of medium size. Proepisternum with ridge-like dorsal
margin ; propleural ridge short and propleural apophysis small. Prosternum subtriangular,
narrow medially, and 37-43 (39) \i long ; with a transverse, 85-101 (89) pi long prosternal ridge
Prothoracic setae on each side : One medial pronotal seta usually present, but two setae may
occasionally occur (av. i-i) ; lateral pronotal seta 0-1 (o-6) ; post-tergital setae absent. Ante-
spiracular dorsal setae 3-5 (3-7) and antespiracular ventral setae 1-2 (1-2). Prosternal setae
2-3 (2-5). Prothoracic disc pores on each side : Medial pronotal pores 4-10 (6-3) and lateral
pronotal pores 3-1 1 (5-6) ; these two groups sometimes become rather difficult to separate.
Antespiracular dorsal pores 4-8 (5-5) ; post-tergital pores absent. Prosternal pores 1-3 (i-6)
forming with those of the other side a median group.

Mesothorax : Prescutum subrectangular in dorsal view ; 64-76 (70)0. long and 11 6-1 3 7
(i25)[x wide, the ratio being 1 : 1-6-2-0 (1 : i-8) ; prescutal ridges well developed and prescutal
suture well marked. Scutum with well sclerotized anterolateral extremities, and with a longi-
tudinal median narrow membranous area. Scutum 95-119 (ioi)pi long, i.e. the ratio length of
prescutum to scutum 1 : 1-3-1-8 (1-4). Prealare, prealar ridge, triangular plate and tegula well
developed. Scutellum pentagonal in dorsal view ; 55-70 (58) pi long, and 98-125 (107) u. wide,
the ratio being 1 : 1-7-1-9 (1 : i-8), and the ratio its length to the length of scutum 1 : i-6-
i-8 (1 : 1-7). Postalare with well separated anterior and posterior postalar ridges. Postnotal
apophysis strong. Mesopleuron : Mesopleural ridge interrupted above the coxal articulation ;
basalare well developed. Mesepisternum semimembranous ; lateropleurite narrow ; mesepimeron
well sclerotized. Mesosternum : Basisternum 122-143 (128) pi long and 183-229 (195) pi wide ;
marginal, precoxal ridges and furca well developed. Mesothoracic spiracles 18-24 (21) [*■ wide
at opening, and with a 37-46 (43)0. long supporting bar. Mesothoracic setae on each side :
Prescutal setae 1-2 (1-7) ; scutal setae 7-10 (8-i) ; scutellar setae usually 2 and occasionally 3
(av. 2-1) ; tegular setae 2-3 (2-2). Postmesostigmatal setae 3-5 (4-3), in a latero-ventral group
behind the spiracle ; basisternal setae 7-1 1 (9-2). Mesothoracic disc pores : Mesospiracular
pores comparatively numerous, 4-7 (51) posteriorly associated with each spiracle.

Metathorax : Metapostnotal sclerites and metapostnotal ridge well developed. Metapleural
ridge with the usual attenuation near the middle at the point of origin of the metapleural
apophysis ; metapleural wing process small. Precoxal ridge of metathorax, metepisternum and
metepimeron distinct ; metasternal apophysis small. Metathoracic spiracles identical with the
mesothoracic. Metathoracic setae on each side : Metatergal setae always include a group of
3 sublateral setae, and one isolated submedially ; one metapleural seta usually present.
Anterior metasternal setae 2-4 (3-1) ; posterior metasternal setae 1-3 (1-9). Metathoracic disc

pores on each side : Metaspiracular pores 2-4 (2-6) ; anterior metasternal pores 1-2 (1-5) ;

posterior metasternal pores absent.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

109

Wings : 854-1050 (924) u. long, and 322-434 (364)^1 wide ; usually with 3 alar setae
(occasionally 2, av. 2-q(, and 3-4 (3-3) circular sensoria. Hamulohalterae 64-73 (70) (x long,
and 12-18 (15) (i wide ; with a weak slender ridge and one, apically hooked, 52-58 (55) n long
seta ; i.e. the ratio length of seta to the length of the hamulohaltera 1 : 1-2-1-4 (1 : 1-3).

Legs : Moderately long ; the fore legs shortest, and the hind legs longest ; the ratio length
of the hind leg to the total body length 1 : 1-8-2-1 (1 : 1-9). Trochanter about 24^ wide.
Femur about 40^ wide, the ratio width to length of hind femur 1 : 4-2-5-0 (4-5). Tibia about
24 (x wide, with 2 strong apical spurs and 2-4 smaller spines ; the femur in front leg longer
than the tibia, the ratio their lengths being 1-01-1-4 ( I-1 ) : *■ Tarsus about 21^1 wide ;
tarsal digitules apically knobbed, about 34 \l long. Claw uniformly tapering to a sharply
pointed apex ; ungual digitules finely pointed. The following table shows the lengths of the
leg segments (in microns) and the number of setae on each:

Coxa

Trochanter
Femur
Tibia

Tarsus

Claw

Total length of leg

length

f.s.

h.s.

length

f.s.

h.s.

length

f.s.

h.s.

length

f.s.

h.s.

length

f.s.

h.s.

length

Fore leg

37-43 (40)
6-1 1 (90)

3-7 (4-3)

61-67 (62)

3-5 (4-4)

i-3 (2-0)

153-174 (162)

17-24 (20-1)

3-6 (3-9)

137-162 (153)

21-30 (256)

3-5 (36)

73-79 (76)
18-23 (203)
0-2 (i-o)

37-46 (40)
503-570 (534)

Middle leg

37-48 (40)
8-14 (9-6)
3-6 (4-i)

61-67 (64)
3-7 (47)
1-2 (i-3)

156-177 (168)

16-26 (209)

3-6 (43)

153-183 (168)

24-32 (261)

3-5 (40)

73-79 (76)
16-23 (19-4)
0-2 (0-9)

37-46 (40)
519-598 (555)

Hind leg

43-52 (46)
9-17 (12-0)
4-7 (5-o)

67-73 (70)
4-10 (63)

i-4 (2-4)

165-198 (180)

20-29 (246)

3-6 (41)

189-217 (204)

33-40 (35-6)

3-6 (43)

76-82 (79)
19-26 (216)
o-3 (10)

37-46 (40)
577-665 (619)

1
1

1
1

\
\

a.

c

t

"

200

y«

Fig. 10. Planococcus ireneus De Lotto, lateral view.

no MORPHOLOGY AND TAXONOMY OF ADULT MALES

Abdomen : 380-562 (448) u. long and 281-342 (319) fx wide. Tergites of segments I, II and
III small ; those of segments VIII and IX + X large. The sternites of segment VIII distinct.
Ostioles well developed, with a 52-61 (55) [x long orifice. Abdominal setae on each side :
Dorsal setae 2-4 on segments I to VII, and always 2 on segment VIII. Pleural setae 3-5 on
segments I to VIII, including one conspicuously longer seta on the latter. Ventral setae 1-2
on segment II, 2-3 on segment III, always 2 on segments IV to VII, and absent on segment
VIII. Abdominal disc pores : Segments I to VII with 9-23 (14-1), 4-11 (7-4), 4-11 (7-4).
5-10 (6-7), 5-10 (6-9) 4-9 (7-3), and 5-12 (7-8) pleural pores, respectively.

Setae of glandular pouch include two, 275-305 (290) jx long tail setae, and one, 79-107 (92) y.
long seta ; the ratio length of the tail setae to the total length of the body 1 : 3-7-4-6 (4-0).

Genital segment : Conical in dorsal view, extending posteriorly into a tubular, apically
rounded style ; the latter slightly curving upwards in lateral view. Penial sheath 11 3-1 34
(i22)[x long and 85-98 (89)fx wide, i.e. the ratio 1-3-1-5 : 1 (1-4 : 1), and the ratio its length to
the total body length 1 : 8-9-10-6 (1 : 9-5). Basal ridge of penial sheath with a small internal
projection ; process of penial sheath well developed, with 3-4 (3-6) setal sensilla. Aedeagus
tapering to a pointed end ; internal genital aperture large. Setae of genital segment slightly
smaller than the other body setae ; the dorsal always includes 3 h.s. on each side, near the
base of the style ; the ventral group consists of 3-4 (3-4) h.s. on each side of the penial sheath

Material : 10 specimens examined, collected by D. N. McNutt, on roots of Coffea
arabica, in Buwagogo, Uganda, 23.viii.1961.

NIPAECOCCUS Sulc, 1945

Nipaecoccus vastator (Maskell)

(Text-figs, n, 12)

Only the winged males known. A slender species, of medium size, with moderately long
antennae and legs. Mounted specimens 1 148-1358 (1246) [x long, 280-322 (294)0. wide and
2310-2660 (2520) |x wing expanse.

Body setae and disc pores : The antennae with fleshy and hair-like setae, approximately
37 (x long ; the body itself and the legs with hair-like setae only, about 28 jx long. Quadrilocular
and occasionally quinquelocular disc pores, about 6jx in diameter, occur on the head, thorax
and abdomen ; Beardsley (i960) observed few pores with 6 peripheral oculi.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression hardly indicated in lateral view. Length from apex to postoccipital ridge 1 07-1 31
(n6)jx ; from apex to neck 177-198 (186) (x ; width across the genae 207-244 (229) jx. Dorsal
arm of midcranial ridge anteriorly detached from other arms by a short distance, and
posteriorly meeting the postoccipital ridge ; ventral and lateral arms forming a T-shaped ridge.
Postoccipital ridge U-shaped and anteriorly continuous with the preocular ridges. Dorsomedial
part of epicranium weakly sclerotized. Preocular -f- interocular ridges posteriorly joined to
postocular ridge below the ocellus ; preocular ridge without any apparent ventral rudiment.
Preoral ridge slender. Eyes : Dorsal simple eyes project beyond the outer margins of the
head in dorsal view ; their corneae 34-46 (40) [x in diameter and separated by 107-125 (ii3)(x,
i.e. 2-5-3-5 (2-8) times as much as their diameter apart. Ventral simple eyes only 31-43 (37) jx
in diameter, and separated by 27-40 (34) u.. Lateral ocelli large. Cranial apophysis apically
truncate. Tentorial bridge slender. Dorsal head setae 10-14 ( IJ '8), on each side of the median
line ; each gena almost always with 3 genal setae (occasionally 2). Ventral head setae on each
side as follows : 5-6 (5-4) between the ventral eyes ; 8-12 (io-i) forming on both sides a

length in y

85-104

73-95

76-95

(av.)

(93)

(85)

(85)

f.s.

14-24

16-30

20-31

(av.)

(19-6)

(224)

(24-3)

h.s.

3-8

3-6

2-4

(av.)

(5-5)

(4-3)

(2-8)

79-104

89-101

73-92

85-101

(92)

(95)

(82)

(95)

19-28

22-29

16-27

23-29

(24-6)

(24-6)

(2i-3)

(25-5)

3-6

4-6

3-5

i-4

(4-2)

(47) "

(4-o)

(2"5)

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE m

transverse band across the area of the ventral preocular depression ; 4-6 (5-1) in a longitudinal
row on each side anteriorly. Head disc pores : One dorsal pore always present near the base
of each antenna. Antennae : Filiform ; 10-segmented ; 744-903 (82i)y. long, i.e. longer
than half the total length of the body (ratio 1 : 1-4-1-6, av. 15), and longer than the hind legs
(ratio 1-2-1-4, av. 1-3 : 1). Scape 43-49 (46)0. long and 46-52 (49) u. wide at base ; usually
with 4 h.s. (occasionally with 5 or 6, av. 4-2). Pedicel 58-73 (67) [* long and 40-46 (43)0. wide ;
with 20-36 (27-2) f.s., 5-1 1 (7-6) h.s. and a sensillum placodeum. Flagellum : Segment III
club-shaped ; subequal in length or somewhat shorter than the terminal segment (ratio 1 : i-i-i,
av. 1 03) ; about 3i[x wide, i.e. the ratio its width to its length 1 : 2-8-3-4 (3-0). Segments
IV to X cylindrical, and about 24 u. wide, the ratio width to length of segment IX being
1 : 3-3-8 (3-4). The following table shows the lengths of the flagellar segments and the number
of setae on each :

III IV V VI VII VIII IX X

79-101

(89)

17-32

(23-2)

4-6

(47)

Antennal bristles slightly stouter than the fleshy setae. One ventral bristle present on each
of antennal segments VIII and IX. Terminal segment with 3 subapical bristles and 2 much
smaller, lateroventral ones, at a greater distance from the apex. The segment also with two
capitate subapical sensory setae and one apical, hair-like seta.

Thorax : 456-570 (524)^ long. Prothorax : Pronotal ridges with the usual interruption
medially ; lateral pronotal sclerites and post-tergites distinct. Proepisternum with a ridge-like
dorsal margin. Prosternum subtriangular, 37-52 (43) [z long ; prosternal ridge 92-113 (104) y
long. Prothoracic setae on each side of the median line : Medial pronotal and lateral pronotal
setae 0-1 (av. 0-2 and 0-3 respectively) ; post-tergital setae absent ; antespiracular dorsal setae
2-4 (2-7). One antespiracular ventral seta always present. Prosternal setae 1-3 (1-9).
Prothoracic disc pores on each side : Medial pronotal pores 1-3 (1-9) ; lateral pronotal pores
1-3 (2-1) ; antespiracular dorsal pore always one. Prosternal pores usually one, but sometimes
either side may be with 2 pores or none at all (av. 0-9).

Mesothorax : Prescutum 70-95 (79) y. long and 1 10-134 ( I22 )u- wide (ratio 1 : 1-4-1-7, av.
1-5). Prescutal ridges well developed and prescutal suture distinct. Scutum with heavy antero-
lateral sclerotizations, and with a medial longitudinal narrow membranous area ; 89-113
(i04)u. long, i.e. the ratio lengths of prescutum to scutum 1 : 1-1-1-4 ( I- 3)- Prealare, prealar
ridge and triangular plate well developed. Scutellum 49-64 (58) y. long and 85-104 {98) y. wide,
i.e. the ratios its length to its width 1 : 1-6-1-9, av. 17, and its length to the length of the
scutum 1 : 1-6-2, av. i-8. Postalare with well separated anterior and posterior postalar ridges.
Mesopleuron : Mesopleural ridge interrupted above the coxal articulation ; basalare strong.
Other pleural structures typical of the family. Mesosternum : Basisternum 122-153 ( I 37)( jL
long and 171-214 (189) y wide ; marginal and precoxal ridges strong. Mesothoracic spiracle
18-21 (20) (x wide at opening, with 40-46 (43) y long supporting bai . Mesothoracic setae on each
side : Prescutal setae 3-5 (3-6) ; scutal setae 7-1 1 (8-i) ; scutellar setae 2-4 (2-8). Tegular
setae 4-6 (4-9). Postmesostigmatal setae 5-8 (5-9), occurring in a lateral group behind each
spiracle. Basisternal setae 6-9 (7-3). Mesothoracic disc pores : 1-2 (1-2) mesospiracular pores,
and 0-1 (0-9) postmesostigmatal pores present on each side of the median line.

Metathorax : Metapostnotal sclerites and metapostnotal ridge well developed. Pleural ridge
attenuated near the middle, where a pleural apophysis originates ; pleural wing process small.
Episternum, epimeron, precoxal ridge and metasternal apophysis distinct. Metathoracic spiracle
similar to the mesothoracic one. Metathoracic setae on each side : Metatergal setae 2-5 (2-6) ;

H2 MORPHOLOGY AND TAXONOMY OF ADULT MALES

metapleural setae usually absent, but one seta may occasionally occur on one side (av. 02).
Anterior metasternal setae 1-3 (1-9) ; posterior metasternal setae 1-2 (1-3). Metathoracic disc
pores on each side : Metaspiracular pores 2-3 (2-2) ; anterior metasternal pores 0-2 (09), and
posterior metasternal pores 0-2 (i-o).

Fig. 11. Nipaecoccus vastator (Maskell), dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

113

Wings : 980-1120 (1050)0. long and 378-448 (406)^ wide. Alar lobe, axillary and additional
sclerites well developed. Alar setae usually 3, but 2 or 4 may occur (av. 2-9) ; with 3 circular
sensoria in a compact row. Hamulokalterae 76-92 (80) u. long and 15-18 (17) u. wide ; with a
58-67 (61 )u. long apically hooked seta, i.e. the ratio lengths of seta to hamulohaltera 1 : 1-2— 1-4
(1-3)-

Legs : Moderately long and slender ; the ratio length of the hind leg to the total length
of the body 1 : 1-9-2-1 (20). Coxa and trochanter about 55 and 27^ wide respectively ; the
latter with a long apical seta. Femur about 40^ wide ; that of the middle leg shortest ; the
ratio width to length of hind femur 1 : 4-3-51 (47)- Tibia about 23^ wide, with 2 apical
strong spurs and 3-5 smaller spines ; the ratio lengths of femur to tibia in front leg 1 : i-i-i
(104). Tarsus about 2i[i wide and tarsal digitules about 400. long. Claw gradually tapering
towards a pointed end ; with a pair of finely pointed ungual digitules. The lengths of the leg
segments (in microns), and the number of setae on each are given below :

Coxa

lengths
h.s.

Fore leg
40-46 (43)
9-14 («-2)

Middle leg

40-49 (46)
10-15 (12-8)

Hind leg

43-52 (49)
10-16 (130)

Trochanter

lengths
h.s.

58-67 (64)
4-7 (5'4)

58-67 (64) .
4-6 (52)

61-70 (67)
4-7 (5-6)

Femur

lengths
h.s.

156-183 (171)
28-41 (352)

146-177 (162)
22 36 (276)

156-186 (171)
25-40 (314)

Tibia

lengths
h.s.

150-192 (177)
32-43 (364)

168-207 (i 86 )
33 47 (38-8)

192-250 (220)
35-50 (423)

Tarsus

lengths
h.s.

82-92 (85)
20-27 (236)

82-92 (85)
22-28 (24-6)

85-101 (92)
24-31 (266)

Claw

length

31-37 (34)

31-37 (34)

31-37 (34)

Total length

of leg

522-613 (570)

522-628 (577)

567-689 (631)

Abdomen : 448-517 (479)^ long and 258-304 (281) (ji wide. Tcrgites of segments I and II
small, and those of segments VI 1 1 and IX X distinct. A weak sternite on each side of
segment VIII present. Ostioles ill-defined. Abdominal setae on each side : Dorsal setae 1-2

OH <w^; ■ •

:^^A^J±1J^^

f \ \ o o

w o. a

200.-

Fig. 12. Nipaecoccus vastator (Maskell), lateral view.

ii4 MORPHOLOGY AND TAXONOMY OF ADULT MALES

on segment I, 2-4 on segments II to VII, and always 1 on segment VIII. Pleural setae 2-3
on segment I, 3-5 on segments II to VII, and always 3 on segment VIII, including one slightly
longer seta. Ventral setae 1-2 on segment II, 2-3 on segments III to VII, and absent on
segment VIII. Abdominal disc pores on each side : Pleural pores 2-4 on segment I and 1-3
on segments II to VII. One ventral pore usually also present on segments III to VII, but
sometimes two occur on segments IV and VI, and sometimes the pores are absent on segment
VII.

Glandular pouch well developed ; glandular pouch setae include a pair of 244-300 (268) \j.
long tail setae, and one, about 70^ long seta, i.e. the ratio length of the tail setae to the total
length of the body 1 : 4-2-5-2 (4-6).

Genital segment small ; subtriangular in dorsal view ; style curving upwards in lateral view.
Penial sheath 101-119 {ioj)y. long and 64-76 (70)^1 wide (ratio 1-5-1-6, av. 1-52 : 1, and the
ratio its length to the total body length 1 : 11-1-12-3, av - n - 6). Basal ridge of penial sheath
its projection and the process of penial sheath well developed. Aedeagus tapering posteriorly
towards a pointed tip. Setae of genital segment on each side : Dorsally 3 setae always present
near the base of the style ; ventrally, 4-7 (5-4) setae occur on the penial sheath, and 3-4 (3-6)
setal sensilla on its process.

Material : 10 specimens examined, collected by G. M. Das, in Cinnamara, Assam,
India, during October, 1961 ; host plant not stated.

Nipaecoccus nipae (Maskell)
(Text-figs. 13, 14)

Macropterous forms only known. A narrow and slender species ; comparatively very
small, with moderately long antennae and legs. When mounted 840-980 (896) pi long, 210-224
(214) (x wide and 1974-2240 (2086) \l wing expanse.

Body setae and pores : Fleshy and hair-like setae rather difficult to separate ; those occurring
on the antennae about 31U. long and those on the legs slightly shorter ; the body itself only
with hair-like setae, about i8(x long. Disc pores present on the head, thorax and abdomen ;
these about 6fj. in diameter, usually with 5 or 6, and occasionally 4 peripheral loculi.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression hardly indicated in lateral view. Length from apex to postoccipital ridge 85-98
(92)^ ; from apex to neck 128-143 (137) (i ; width across the genae 162-177 (171) [i. Dorsal
arm of midcranial ridge distinct, anteriorly separated from the other arms and posteriorly
meeting the postoccipital ridge. Lateral and ventral arms forming a T-shaped ridge. Post-
occipital ridge weakly developed and anteriorly confluent with the preocular ridges. Dorso-
medial part of epicranium slightly raised. Preocular + interocular ridges joining postocular
ridge below the ocellus ; ventral rudiment of preocular ridge absent. Eyes : Dorsal eyes not
projecting beyond the outer margins of the head in dorsal view ; 18-21 (19) u. in diameter and
separated by 79-89 (85) jx, i.e. 4-1-4-8 (4-6) times as much as their diameter apart. Ventral
eyes 24-27 (26) \l in diameter and 18-21 (20) \l apart. Lateral ocelli well developed. Ocular
sclerites weakly sclerotized. Cranial apophysis apically truncate. Tentorial bridge slender.
Dorsal head setae 8-10 (8-8) on each side of the median line ; each gena always with 2 genal
setae. Ventral head setae on each side : 4-5 (4-1) between the ventral eyes ; 9-13 (io-i)
forming on both sides a transverse band in the area of the ventral preocular depression ; 2-3
(2-3) in a row on each side of the ventral arm of the midcranial ridge. The head also with
1-2 (1-3) dorsal pores on each side near the antennal base.

Antennae : Filiform ; 10-segmented ; 479-540 (509) (x long, i.e. somewhat longer than half

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 115

the length of the body (ratio 1 : 1 -6-1-9, av. J, 8) and as long as, or slightly longer than the
hind leg (ratio i-o-i-i, av. 1-05 : 1). Scape 34-37 (36)[z long and 31-34 (33) jj- wide at base ;
always with 4 h.s. Pedicel 52-55 (54) u. long and 31 [jt. wide ; with 14-21 (17-2) f.s., 8-1 1 (9-4)
h.s. and a sensillum placodenm. Flagellum : Segment III club-shaped and about as long as
segment X (ratio 1 : i-i-i (104) ), both being the longest of all ; segment III about 20jx
wide, i.e. the ratio its width to its length 1 : 2-7-3-0 (2-9). Segments IV to X cylindrical and
about i8(i wide ; the ratio width to length of segment IX being 1 : 2-3-2-8 (2-5). The following
table shows the lengths of the flagellar segments and the number of setae on each :

III IV V VI VII VIII IX X

length in u.

58-64

43-52

46-55

49-58

49-55

46-55

43-52

58-67

(av.)

(61)

(46)

(51)

(52)

(52)

(51)

(46)

(62)

f.s.

8-14

10-15

12-17

15-18

15-19

15-20

n-16

18-25

(av.)

("•4)

(120)

(15-2)

(16-6)

(17-2)

(16-8)

(M-5)

(2I-I)

h.s.

3-5

2-4

2-3

2-3

2-3

1-2

1-2

1-2

(av.)

(4-8)

(2-5)

(2-4)

(2'4)

(2-3)

(i-6)

(1-4)

(x-4)

Antennal bristles slightly stouter than the fleshy setae, and of subequal length ; segments
VIII and IX each with one ventral bristle. Terminal segment with 3 preapical bristles, two
capitate subapical sensory setae and one apical hair-like seta.

Thorax : 372-418 (395) |x long. Prothorax : Pronotal ridges medially interrupted at a weak
point. Lateral pronotal sclerites and post-tergites small. Dorsal margin of proepi sternum
ridge-like. Prosternum subtriangular. anteriorly forming a weakly sclerotized ring ; 31-37
(34) (i long and posteriorly bounded by a transverse, 70-79 (76) u. long prosternal ridge.
Prothoracic setae on each side : Medial pronotal setae 0-2 (i-i) ; lateral pronotal and post-
tergital setae absent ; antespiracular dorsal setae 1-3 (1-2). One antespiracular ventral seta,
and one prosternal seta always present. Prothoracic disc pores on each side : Medial pronotal
pores 8-1 1 (9-6) ; lateral pronotal pores 2-3 (2-2) ; antespiracular dorsal pores 1-2 (1-2). One
median prosternal pore usually present, but sometimes one pore occurs on each side of the
median line (av. o-8).

Mesothorax : Prescutum 67-76 (73) jx long and 82-92 (89) ;x wide, the ratio being 1 : 1-2-1-3
(1-22). Prescutal ridge strong and prescutal suture distinct. Scutum heavily sclerotized
antero-laterally, with a medial longitudinal narrow membranous area ; 70-76 (74) [a long, i.e.
the ratio lengths of prescutum to scutum 1 : i-o-i-i (102). Prealare, prealar ridge and
triangular plate well developed. Scutellum 34-40 (37) u. long and 64-73 (70) u. wide, i.e. the
ratios its length to its width 1 : 1-8-2-1 (1-9), and its length to the length of scutum 1 : 1-9-2-3
(2-0). Postalare with well separated anterior and posterior ridges. Mesopleuron : Mesopleural
ridge interrupted above the coxal articulation ; basalare stout and subepisternal ridge well
developed. Mesepisternum and mesepimeron distinct ; lateropleurite narrow. Mesosternum :
Basisternum 92-107 (io2)|j. long and 131-137 (135) ^x wide ; marginal and precoxal ridges strong.
Mesothoracic spiracle about 150. wide at opening, with 31-34 (32)^ long supporting bar. Meso-
thoracic setae on each side ; Prescutal setae 2-3 (2-4) ; scutal setae 2-3 (2-6) ; scutellar seta o— 1
(o-6). Tegular setae 2-3 (2-2). Postmesostigmatal setae 2-3 (2-4), in a latero-ventral group.
Basisternal setae 6-8 (6-6). Mesothoracic disc pores : Mesospiracular pores 1-2 (1-2) behind
each spiracle ; postmesostigmatal pores always one.

Metathorax : Metapostnotal sclerites and metapostnotal ridge distinct. Pleural ridge attenuated
near the middle where a small pleural apophysis originates, and dorsally supports the wing
process. Metepisternum and metepimeron distinct. Precoxal ridge well developed and meta-
sternal apophysis small. Metathoracic spiracles identical with those of mesothorax. Meta-
thoracic setae on each side : Metatergal setae 1-2 (1-4) ; metapleural setae usually absent,
although one seta was found in one specimen. Anterior metasternal setae absent, and posterior
metasternal seta 0-1 (0-3). Metathoracic disc pores : Metatergal and metaspiracular pores
missing ; anterior and posterior metasternal pores 1-2 (averages i-i and 13, respectively).

n6

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Wings : 840-952 (896) u. long and 322-378 (336) [x wide ; always with 3 alar setae and 2
minute circular sensoria. Hamulohalterae 55-61 (58) \l long and 12-15 ( I 4)t JL wide ; with a
slender ridge and one apically hooked, 37-46 (41) \l long seta, i.e. the ratio lengths of the seta
to the hamulohaltera 1 : 1-2-1-6 (1-4).

dmep dm , cr dhs

dse
dhp-^

Fig. 13. Nipaecoccus nipae (Maskell), dorsal and ventral view.

OF PSr-IlDOCOCCIDAE & ERIOCOCCIDAE

117

Legs : Comparatively long and slender ; the ratio length of the hind leg to the total length
of the body 1 : 18 1-9(1-84). Coxa and trochanter about 37 and i8(x wide respectively. Femur
about 27(jl wide ; that of the middle leg shortest and that of the hind leg longest ; the ratio
width to length of the hind femur 1 : 4-7-5-2 (5-0). Tibia about i8(x wide ; with 2 apical
spurs and 2 4 smaller spines ; the ratio length of femur to length of tibia in the front leg
1 : 1 -o-i-i (103). Tarsus about i6(z wide ; tarsal digitules apically knobbed, about 31JX long.
Claw gradually tapering to a sharply pointed apex ; ungual digitules extremely fine. The
following table shows the lengths of the leg segments (in microns) and the number of setae
on each :

Coxa

length

f.s.

h.s.

Fore leg
24-27 (26)
7-io (8-7)
4-6(50

Middle leg

24-27 (26)
7-1 1 (96)
4-6 (49)

Hind leg

27-31 (29)
9-12 (103)
4-7 (5-6)

Trochanter

length

f.s.

h.s.

46-49 (48)
2-4 (2-9)
3-4 (37)

46-49 (48)
2-4 (2-5)
2-4 (3-4)

49-52 (51)
2-5 (31)
3-4 (37)

Femur

length

f.s.

h.s.

125-137 (131)

18-23 (204)

6-9 (72)

119-131 (125).
15 19 (169)
5-7 (6-i)

I3I-M3 (137)
16-21 (195)

5-8 (7-o)

Tibia

length

f.s.

h.s.

125-140 (134)

19-28 (244)

6-9 (7-6)

1 <; 146 (143)

21 29 (26-7)

7-10 (7-8)

162-183 (171)

24-32 (28-2)

7-11 (8-i)

Tarsus

length

f.s.

h.s.

55-58 (57)
12-16 (136)

4-6 (5-2)

55-58 (57)
11-16 (129)
4-6 (49)

61-67 (63)
13-18 (15-1)
4-7 (5'8)

Claw

length

31-36 (34)

^1-36 (34)

31-36 (34)

Total length

of leg

418-442 (427)

424-445 (433)

470-512 (485)

app

aps

200^*

100.

Fig. 14. Nipaecoccus nipae (Maskell), lateral view.

n8 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Abdomen : 274-357 ii 12 )^ l° n g an d 198-213 (205)[x wide. A small tergite on each side of
segments I and II present ; tergites of segments VIII and IX -f X large. A weak sternite
on each side of segment VIII present. Ostioles entirely absent. Abdominal setae on each
side : Dorsal setae 1-3 on segments I to VIII. Pleural setae usually 3 on segments I to VIII
(sometimes 2 setae occur on segment I, and 4 on segment III) ; pleural setae of segment VIII
include a slightly longer seta. Ventral setae always 1 on segment II, 2 on segments III to VII,
and absent on segment VIII. Abdominal disc pores : Segments I to VII with 10-15 (12-2),
4-5 (4 - 5). 4- 6 (4- 8 ). 4-7 (53). 4-7 (5'2). 4-7 (4-9), and 4-6 (5-1) pleural pores respectively.

Setae of glandular pouch consist of a pair of tail setae, 238-259 (250)51 long, and one, 37-46
(40)11 long seta, the ratio length of the tail seta to the total length of the body being 1 : 3-2-3-9
(3-6).

Genital segment small ; subtriangular in dorsal view. Style curving upwards in lateral
view. Penial sheath 76-82 (79)^ long and 55-61 (58)51 wide, the ratio being 1-3-1-5 (1-4) : 1,
and the ratio its length to the total body length 1 : 10-4-11-5 (11-2). Basal ridge of penial
sheath with a small projection ; process of penial sheath well pronounced. Aedeagus gradually
tapering towards a pointed tip. Setae of genital segment : Dorsally 3 setae always present on
each side near the base of the style ; ventrally 3-4 (3-3) setae occur on each side of the penial
sheath, and 3-4 setal sensilla on its process.

Material : 10 specimens examined, collected by J. Munting, on Palm, in Durban,
South Africa, i.x.1963.

MACONELLICOCCUS Ezzat, 1958

Maconellicoccus hirsutus (Green)

(Text-figs. 15, 16)

Macropterous forms only known ; living specimens light brown or yellowish. The males
narrow and slender, of medium size, with comparatively short antennae and moderately long
legs. Mounted specimens 1022-1428 (1232)51 long, 252-322 (294)11 wide and 1708-2310 (2016) 51
wing expanse.

Body setae and disc pores : Antennae and legs with many fleshy and few hair-like setae, about
2151 long ; body itself with only somewhat shorter hair-like setae. Quadrilocular and
occasionally trilocular or quinquelocular pores present on the head, thorax and abdomen.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression hardly pronounced in lateral view. Length from apex to postoccipital ridge 1 13-134
(125)51 ; from apex to neck 153-192 (174)11 ; width across the genae 186-226 (214)51. Dorsal
arm of midcranial ridge anteriorly detached from the other arms, and posteriorly meeting the
postoccipital ridge. Lateral arms forming with the ventral arm a Y-shaped ridge. Postoccipital
ridge with a sclerotized area at its medio-posterior base ; anteriorly confluent with preocular
ridges. Dorsomedial part of epicranium slightly raised. Preocular and interocular ridges join
the postocular ridge below the ocellus ; preocular ridge without any ventral rudiment. Eyes :
Dorsal simple eyes slightly projecting beyond the outer margins of the head in dorsal view ;
their corneae 24-37 (31)11 in diameter, and separated by 104-128 (119)51, i.e. 3-3-4-4 (3-7) times
their corneae apart. Ventral simple eyes 27-40 (34)51 in diameter and 21-31 (28) [i apart.
Lateral ocelli well developed. Ocular sclerites weakly sclerotized. Cranial apophysis with
truncate apex. Tentorial bridge slender. Dorsal head setae 12-16 (13-4) on each side of the
median line ; each gena usually with 3-4, and occasionally 5 genal setae (av. 3.3). Ventral
head setae on each side : 1-2 (1-9) between the ventral eyes ; 13-16 (14-1) forming on both
sides a transverse band across the area of the preocular depression ; 2-3 (2-2) anteriorly on

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 119

each side of the ventral arm of the midcranial ridge. Dorsal head pores 1-2 (1-2) near the base
of each antenna.

Antennae : Filiform ; normally 10-segmented ; 470-653 (573) (i long, i.e. as long as, or
shorter than half the total length of the body (ratio 1 : 2-0-2-3, av. 2-2), and shorter than the
hind leg (ratio 1 : 11-12, av. 1-17). Scape 40-49 (43)^ long and 37-46 (43)10. wide at base ;
always with 4 h.s. Pedicel 55-67 (64)0. long and 31-40 (37);x wide ; with 26-36 (31-7) f.s.,
10-18 (12-3) h.s. and a sensillum placodeum. Flagellum : Segment III club-shaped and longest
of all (the ratio lengths of segments III to X 1-2-1-4, av - I- 3 : x ) '• the segment about 231*
wide, i.e. the ratio its width to its length 1 : 2-9-3-8 (3-4). Segments IV to X cylindrical and
about 21 (x wide, the ratio width to length of segment IX being 1 : 2-1-3-3 ( 2- 7)- The lengths
of the flagellar segments and the number of setae on each are given in the following table :

III IV V VI VII VIII IX X

length in [i

61-85

43-67

43-6i

43-6i

43-63

46-70

46-58

52-7°

(av.)

(76)

(55)

(52)

(52)

(53)

(56)

(52)

(59)

f.s.

13-18

13-20

13-20

12-18

13-19

12-19

12-16

10-16

(av.)

(15-9)

(i6-3)

(16-1)

(14-9)

(164)

(150)

(14-2)

(136)

h.s.

3-7

2-5

2-3

2-5

2-6

3-6

3-6

3-6

(av.)

(47)

(3-4)

(2-5)

(3'5)

(3 9)

(4-4)

(4-4)

(4-5)

Antennal bristles easily distinguishable ; segments VIII and IX each with one ventral
bristle. Terminal segment with 3 preapical bristles, the dorsal of which is about half as long
as the others ; the segment also with two capitate subapical sensory setae and one apical hair-
like seta.

Thorax : 418-600 (540) \i long. Prothorax : Pronotal ridges medially constricted at a weak
point. Lateral pronotal sclerites moderately large and post-lergites narrow. Proepisternum
with a ridge-like dorsal margin. Prosternum triangular, 31-46 (38)11 long ; prosternal ridge
85-104 (96) (x long. Prothoracic setae on each side : Medial pronotal setae 0-2 (1-3) ; lateral
pronotal setae 0-3 (i-i) ; post-tergital setae absent, and antespiracular dorsal setae 1-3 (20).
Antespiracular ventral seta always one, and prosternal setae 1-3 (1-7). Prothoracic disc pores
on each side : Medial pronotal pores 2-5 (3-1) ; lateral pronotal pores 1-3 (2-1) ; antespiracular
dorsal pores 1-2 (1-7). Prosternal pores 1-2 (i-i).

Mesothorax : Prescutum 61-89 (76)1* long and 104-143 (131)11 wide (ratio 1 : 1-6-1-8, av.
1-7) ; presculal ridge strong and prescutal suture well marked. Scutum heavily sclerotized
antero-laterally, and with a median longitudinal narrow membranous area ; 85-125 (1 13) pt
long, i.e. the ratio length of prescutum to scutum 1 : 1-3-1-5 (146). Pleural structures typical
of the family. Scutellum pentagonal, 46-64 (58) (i long and 85-116 (104)0. wide, the ratio being
1 : 1-7-1-9 (i-8) ; subequal to half the length of the scutum, ratio 1 : 1-8-2-1 (1-9). Postalare
with well separated anterior and posterior postalar ridges. Postnatal apophysis large. Meso-
pleuron : Mesopleural ridge interrupted above the coxal articulation ; basalare stout. Sub-
episternal ridge well developed. Mesepisternum and mesepimeron distinct ; lateropleunte
narrow. Mesosternum : Basisternum 122-174 (152)11 long and 146-201 ( 1 80) |i wide ; marginal
and precoxal ridges well developed. Mesothoracic spiracle 18-21 (19)11 wide at opening, with a
37-43 (40) (i long supporting bar. Mesothoracic setae on each side of the median line : Prescutal
setae 5-9 (6-3) ; scutal setae usually 8-1 1, although 13 setae were found on one side of one
specimen (av. io-o) ; scutellar setae 3-6 (3-4). Tegular setae 3-5 (4-1). Postmesostigmatal
setae 4-9 (6-i), in a latero-ventral group. Basisternal setae 7-15 (9-9). Mesothoracic pores :
Mesospiracular pores 0-1 (o-8) ; one postmesostigmatal pore almost always present submedially
on each side, but none or two pores may occur on one side (av. 10).

Metathorax : Metapostnotal sclerites and metapostnotal ridge well developed. Metapleural
ridge attenuated at the point of origin of the metapleural apophysis. Precoxal ridge well
developed and metasternal apophysis well defined. Metathoracic spiracle similar to mesothoracic.
Metathoracic setae on each side : Metatergal setae 3-5 (4-1) ; metapleural setae 0-2 (o-8).

i2o MORPHOLOGY AND TAXONOMY OF ADULT MALES

Anterior and posterior metasternal setae 2-4 (3-1) and 0-1 (o-6), respectively. Metathoracic
pores : Metatergal pores absent, and metaspiracular pores 0-1 (07). Anterior metasternal
pores 1-2 (i-i), and posterior metasternal pores 0-1 (o-g).

dse_

Fig. 15. Maconellicoccus hirsutus (Green), dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 121

Wings : 742-1008 (910) [i long and 294-392 (350) \i wide ; with 4-5 (4-2) alar setae and a
compact row of 3-4 (3-2) minute circular sensoria. Hamulohalterae 61-79 (73) (x long and
12-15 ( I 4)i i wide ; with a slender ridge and one, 43-55 (49)^ long apically hooked seta, i.e.
the ratio length of seta to the length of hamulohaltera 1 : 1-4-1-6 (1-5).

Legs : Moderately long and slender ; the ratio length of the hind leg to the total body
length 1 : 1-8-1-9 (i"85). Coxa about 49 and trochanter about 27^ wide. Femur about 37^
wide ; that of the front leg shortest, and that of the hind leg longest ; the ratio width to length
of the hind femur 1 : 5-0-5-9 (5-6). Tibia about iq jjl wide ; with two apical strong spurs and
3-5 smaller spines ; in front leg, the femur always shorter than the tibia, the ratio their lengths
being 1 : 10-11 (112). Tarsus about ij\i wide ; tarsal digitules apically knobbed, 31-37
(34) (x long. Claw gradually tapering to a pointed tip ; ungual digitules fine. The following
table shows the lengths of the leg segments (in microns), and the number of setae on each :

Fore leg

.Middle leg

Hind leg

Coxa

length

f.s.

h.s.

37-43 (40)
8-17 (130)
5-9 (60)

37-43 (40)
9-16 (126)

5-8 (6-4)

40-49 (46)
10-20 (148)
6-8 (6-8)

Trochanter

length

f.s.

h.s.

40-64 (58)
4-6 (4-6)
3-6 (3-8)

49-64 (58)
4-9 (6-2)
3-6 )4o)

55-70 (64)
5-8 (6-4)
3-6 (4-2)

Femur

length

f.s.

h.s.

137-198 (177)

32-44 (38-4)
6-9 (76)

143-204 (183)

33-44 (39o)

6-9 (77)

153-214 (!92)

38-54 (464)

7-9 (7-6)

Tibia

length

f.s.

h.s.

159-226 (198)

33-48 (394)

4-8 (5'4)

165-241 (211)

38-58 (47-8)

5-8 (6-2)

214-299 (256)

45-69 (54 8)

6-9 (6-8)

Tarsus

length

f.s.

h.s.

70-89 (82)
10-17 (13-6)
5-8 (6-6)

70-89 (82)
1 1-21 (150)
6-8 (6-8)

82-98 (92)
13-25 (170)
6-8 (6-8)

Claw

length

21-27 (24)

21-27 (24)

21-27 (24)

Total length of leg

467-644 (580)

479-665 (598)

564-750 (671)

* /

- - L i. i. ' '

200

Fig. 16. Maconellicoccus hirsutus (Green), lateral view.

122 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Abdomen : 334-494 (403) pi long and 236-312 (281) pi wide. Tergites of segments I and II
small ; those of segments VIII and IX + X large. Sternites of segment VIII weak. Ostioles
well developed, 37-46 (40)10. long at orifice. Abdominal setae on each side : Dorsal setae 2-3
on segments I and VIII, and 3-4 on segments II to VII. Pleural setae 2-4 on segment I,
4-6 on segments II to VII, and 3-4 on segment VIII, including one slightly longer seta.
Ventral setae 1-2 on segment II, 2-3 on segments III to VII, and 0-1 on segment VIII.
Abdominal disc pores : Pleural pores 2-4 (2-8), 0-1 (0-2), 0-1 (o-i), and 0-1 (o-i) on segments
I to VI respectively, and absent on segments VII and VIII. One ventral pore occurs at least
on one side of segments III to VII, forming together a longitudinal sublateral row on each side.

Glandular pouch well developed ; setae of glandular pouch include a pair of 229-296 (265) pi
long tail setae and one, 52-82 (61) pi long seta, i.e the ratio length of the tail setae to the total
length of the body 1 : 4-0-4-9 (4-5).

Genital segment moderately large ; subtriangular in dorsal view ; style more or less straight
and apically rounded in lateral view. Penial sheath 140-180 (162) pi long and 67-89 (82) pi
wide, the ratio being 1-8-2-4 ( 2 '°) : x > an d tne ratio its length to the total body length 1 : 7-1-
8-4 (7-6). Basal ridge of penial sheath with a small projection. Process of penial sheath vestigial
and hardly indicated. Aedeagus comparatively long, anteriorly bent to reach the cavity of
abdominal segment VII. Setae of genital segment : Dorsally, 3 setae always occur on each
side near the base of the style ; ventrally 4-6 (5-1) setae present on each side of the penial
sheath, and 3-4 (3-8) setal sensilla on its vestigial process.

Material : 10 specimens examined, collected by myself, on Psidium guava, in
Fayoum, Egypt, U.A.R., during the second week of August, 1964 ; females were
identified by A. I. Ezz and confirmed by Y. M. Ezzat.

FERRISIANA Takahashi, 1929

Ferrisiana virgata (Cockerell)

(Text-figs. 17, 18)

Macropterous forms only known ; comparatively long and slender, with moderately long
antennae and long legs. When mounted, total body length 1274-1596 (1386) pi, width at
mesothorax 294-364 (322) pi, and wing expanse 2296-2576 (2422) pi.

Body setae and pores : Antennae and legs with numerous fleshy and very few hair-like setae,
about 55 pi long ; the body itself with only hair-like setae, about 28 pi long. Quadrilocular and
occasionally trilocular or quinquelocular disc pores, about 6pi in diameter, occur on the thorax
and abdomen.

Head : Irregularly tetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression well pronounced in lateral view. Length from apex to postoccipital ridge 1 13-137
(i25)pi ; from apex to neck 174-204 (i86)pi ; width across genae 204-253 (22o)pi. Dorsal
arm of midcranial ridge only indicated by a median longitudinal heavy sclerotization. Lateral
and ventral arms forming together a Y-shaped ridge. Postoccipital ridge anteriorly confluent
with the preocular ridges. Preocular -\- interocular ridge join the postocular ridge below the
ocellus ; ventral rudiment of the preocular ridge weakly indicated. Eyes : Dorsal simple
eyes not projecting beyond the outer margins of the head in dorsal view ; their cornea 24-28
(26)pi in diameter and separated by 98-110 (i04)pi, i.e. 3-6-4-5 (4-2) times their diameter apart.
Ventral simple eyes 28-31 (29) pi in diameter and 25-31 (28) pi apart. Lateral ocelli well
developed. Cranial apophysis apically truncate. Tentorial bridge slender. Dorsal head
setae 11-14 (12-5) on each side of the median line ; each gena with 3-4 (3-3) genal setae. Ventral
head setae on each side : 2-4 (3-2) between the ventral eyes ; 5-8 (6-7) forming with their

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 123

partners of the other side a band across the area of the preocular depression ; 2-4 (3-3)
anteriorly on each side of the ventral arm of the midcranial ridge.

Antennae : Filiform ; normally 10-segmented ; 674-848 (760)^ long, i.e. slightly longer
than half the total body length (ratio 1 : 1-7-1-9, av. i-8), and shorter than the hind legs (ratio
1 : I-I-I-2, av. 1-14). Scape 37-46 (43)0. long and 46-52 (49)^ wide at base ; always with
4 h.s. Pedical 64-79 (70)^ long and 34-40 (37) (x wide ; with 12-19 ( J 6'9) f.s., 5-8 (6-o) h.s.
and a sensillum placodenm. Flagellum : Segment III being longest (ratio its length to the
length of segment X 1-2-1-6, av. 1-4 : 1) ; the segment about 23^ wide, the ratio its width
to its length being 1 : 4-3-5-6 (5-1). Segments IV to X cylindrical and about 2ijx wide, the
ratio width to length of segment IX being 1 : 2-7-3-1 (2-9). The following table shows the
lengths of the flagellar segments and the number of setae on each :

III IV V VI VII VIII IX X

length in (i

92-131

64-89

73-95

67-92

67-82

64-76

58-73

73-89

(av.)

(116)

(76)

(79)

(82)

(73)

(70)

(67)

(82)

f.s.

13-19

9-15

13-17

14-19

12-17

12-17

11-15

9-15

(av.)

(15-9)

(12-2)

(14-5)

(17-0)

(M-i)

(14-2)

(12-9)

(12-6)

h.s.

0-2

0-2

0-2

1-2

1 -2

0-2

O-I

O-I

(av.)

(0-7)

(o-8)

(o-3)

(i-8)

(1-4)

(i-o)

(O-I)

(01)

Antennal bristles conspicuously stout ; segments VIII and IX each with a ventral bristle.
Terminal segment with 3 preapical bristles, the dorsal of which slightly shorter and at a greater
distance from the apex, two capitate subapical sensory setae, and one apical hair-like seta.

Thorax : 555-737 (608).) jjl long. Prothorax : Pronolal ridges medially interrupted at a
weakly sclerotized point. Lateral pronotal sclerites and post-lergites comparatively large.
Dorsal margin of proepisternum ridge-like. Prosternum triangular, 31-46 (37) [j. long ;
posteriorly supported by a transverse, 107-128 (1 13) (J. long prosternal ridge. Prothoracic setae
on each side of the median line : Medial pronotal setae 2-3 (2-6) ; lateral pronotal setae 2-4
(2-9) ; post-tergilal setae 0-3 (1-4) ; antespiracular dorsal setae 2-4 (3-1). One antespiracular
ventral seta and one prosternal seta always present. Prothoracic pores on one side : Medial
pronotal pores 2-6 (3-2) ; lateral pronotal pores 1-3 (i-8) ; antespiracular dorsal pores 0-1 (0-5).
Prosternal pores absent.

Mesothorax : Prescntum 92-113 (98)^ long and 128-165 ( I 47)( A wide (ratio 1 : 1-4-1-6, av.
I- 5) '• prescutal ridge well developed and prescutal suture distinct. Scutu»i heavily sclerotized
antero-laterally, with a median longitudinal narrow membranous area ; 116-156 (i3i)(x long,
i.e. the ratio lengths of prescutum to scutum 1 : 1-2-1-4 ( I- 3)- Prealare, prealar ridge, triangular
plate, anterior and posterior notal wing processes well developed. Scutellum pentagonal, 49-64
(58) [i. long and 101-131 (ii6)(a wide (the ratio being 1 : 1-9-2-2, av. 2-0, and the ratio its length
to the length of the scutum 1 : 2-1-2-6, av. 2-3). Poslalare with well separated anterior and
posterior postalar ridges. Postnotal apophysis strong. Mesopleuron : Mesopleural ridge
interrupted in the usual manner ; basalare stout. Subepisternal ridge, mesepislernum, latero-
pleurite and mesepimeron typical of the family. Mesosternum : Basisternum 143-198 (162) |z
long and 180-229 ( 2ll )y- wide ; bounded by the marginal and precoxal ridges. Mesothoracic
spiracle 18-21 (20) \l wide at opening, with a 37-43 (40) (i. long supporting bar. Mesothoracic
setae on each side : Prescutal setae 2-4 (3-1) ; scutal setae 6-7 (6-3) ; scutellar setae 3-4 (3-2).
Tegular setae 1-3 (1-7). Postmesostigmatal setae 2-3 (2-3), occurring only latero-ventrally
behind each spiracle. Basisternal setae 6—9 (6-8). Mesothoracic disc pores : Mesospiracular
pores 0-2 (o-8) ; postmesostigmatal pores absent.

Melathorax : Metapostnotal sclerites and metapostnotal ridge well developed. Metapleural
ridge attenuated at the point of origin of the metapleural apophysis ; metapleural wing process
small. Precoxal ridge and metasternal apophysis well developed. Metathoracic spiracle
identical with that of the mesothorax. Metathoracic setae on each side : Metatergal setae
4-5 (4-2). Dorsostigmatal setae 0-1 (03), and metapleural seta always one behind each spiracle.

I2 4

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Anterior and posterior metasternal setae 1-2 (1-5) and 2-3 (27), respectively. Metathoracic
pores : 1-2 (1-2) sub-marginal metatergal pores, and 2-3 (2-2) metaspiracular pores present ;
anterior and posterior metasternal pores absent.

Wings : 1008-1120 (1064) pi long and 420-532 (462) ji. wide ; with 3, and occasionally 4

Fig. 17. Ferrisiana virgata (Cockerell), dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

125

(av. 31) alar setae, and a compact row of 3 minute circular sensoria. Hamulohalterae 76-89
(82)(jl long and 15-21 (i8)[x wide ; with a 61-70 (64)11 long apically hooked seta, i.e. the ratio
length of seta to hamulohaltera being 1 : 1-2-1-5 ( I- 3)-

Legs : Comparatively long and slender ; the ratio length of the hind leg to the total body
length 1 : 1-5-1-7 (i-6). Coxa and trochanter about 51 and 27(1 wide respectively. Femur
about 37 u. wide ; that of the fore leg shortest and that of the hind leg longest ; the ratio width
to length of hind femur 1 : 6-8-7-6 (70). Tibia about 2 1 (j. wide ; with two apical spurs and
5-8 smaller spines ; in front leg, femur shorter than tibia, the ratio their lengths being 1 : 11-
1-3 (12). Tarsus about 2ifjt wide ; tarsal digitules apically knobbed, 34-43 (37)(x long. Claw
gradually tapering to a pointed tip ; ungual digitules fine. The lengths of the leg segments
(in microns) and the number of setae on each are given below :

Fore leg

Middle leg

Hind leg

Coxa

length

f.s.

h.s.

43-49 (46)
I3-I8(I53)

2-5 (37)

46-52 (49)
12-18 (140)
3-5 (4-0

52-58 (55)
12-23 (175)
2-5 (3-6)

Trochanter

length

f.s.

h.s.

58-67 (64)
3-5 (3-6)
i-4 (27)

61-67 (64)
3-5 (40) •
i-3 (i-6)

65-70 (67)
3-5 (4-3)
i-3 (i-8)

Femur

length

f.s.

h.s.

198-253 (226)

25-40 (32 -6)

2-6 (36)

207-259 (232)

27-34 (3°- 1)

2-3 (2-5)

232-278 (256)

32-41 (35-6)

2-5 (30)

Tibia

length
f.s.

h.s.

220-290 (262)

30-43 (37 8)

i-3 (2-0)

238-308 (281)
39-48 (435)

o-3 (i'4)

317-397 (357)

47-59 (530)

2-5 (2-9)

Tarsus

length
f.s.

79-92 (89)
18-22 (196)

85-95 (9i)
20-23 ( 2I- 3)

98-110 (104)
19-24 (221)

h.s.

o-3 (17)

0-2 (i-o)

0-2 (09)

Claw

length

24-31 (28)

24-31 (28)

24-31 (28)

Total length

of leg

622-778 (708)

670-805 (741)

796-946 (872)

200.

100^-

Fig. 18. Ferrisiana virgata (Cockerell), lateral view.

126 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Abdomen : 448-593 (509)0 long and 274-327 (304)0 wide. Tergites of segments I, II, III
and sometimes IV present ; those of segment VII and segments IX + X large. Sternites of
segment VIII weak. Ostioles well developed, 37-46 (40) o long at orifice. Abdominal setae
on each side : Dorsal setae 3-4 on segments I to VIII. Pleural setae 2-3 on segments I to VIII,
including one slightly longer seta on the latter segment. Ventral setae usually 1 and sometimes
2 on segment II, 2-3 on segments III to VII, and absent on segment VIII. Abdominal disc
pores : Segments I to VII with 1-3 (1-3), 1-2 (12), 1-3 (i-8). 1-3 (2-1), 1-3 (1-9), 1, and 1
pleural pores respectively.

Setae of glandular pouch include a pair of 409-427 (421)0 long tail setae and one, 70-101 (82)0
long seta, i.e. the ratio length of the tail setae to the total length of the body 1 : 3-0-3-6 (3-3).

Genital segment comparatively small ; subtriangular in dorsal view, with a broadly rounded
apex ; style more or less straight in lateral view. Penial sheath 122-137 ( I2 8)o long and
76-92 (79)0. wide ; i.e. the ratio length to width 1-5-1-7 (i-6) : 1, and the ratio its length to the
total body length 1 : io-o-n-6 (io-8). Basal ridge of penial sheath and its projection well
developed ; process of penial sheath absent. Aedeagus rather stout ; internal genital aperture
large. Setae of genital segment : Dorsally, 3 setae always present on each side near the base
of the style ; ventrally, 3-5 (3-8) setae, and 3-4 (3-3) minute setal sensilla occur on each side
of the penial sheath.

Material : 10 specimens examined, collected by A. I. Ezz, on Croton sp., in
Cairo, Egypt, U.A.R., 22. xi. 1962.

TRIONYMUS Berg, 1899

Trionymus newsteadi (Green)

(Text-figs. 19, 20)

Winged forms only known ; newly emerged males light brown, becoming darker soon
afterwards, with dark brown or blackish eyes. They are narrow and slender, comparatively
large with short antennae and moderately long legs. When mounted 1 190-1498 (1372)0 long,
308-378 (350)0 wide at mesothorax and 2926-3542 (3318) o wing expanse.

Body setae and disc pores : Antennae with numerous fleshy setae, about 310 long, and a
few slightly longer hair-like ones ; legs and main parts of the body with hair-like setae only.
Quadrilocular and occasionally quinquelocular disc pores, about 6y. in diameter, occur on the
thorax, abdomen and sometimes on the head.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; with a hardly indicated
ventral preocular depression. Length from apex to postoccipital ridge 116-125 (122)0. ; from
apex to neck 183-201 (192)0. ; width across the genae 214-250 (235)0. Dorsal arm of mid-
cranial ridge anteriorly separated from other arms and posteriorly meeting the postoccipital
ridge. Lateral and ventral arms forming a Y-shaped ridge. Postoccipital ridge anteriorly
confluent with preocular ridges. Preocular -\- interocular ridges joining the postocular ridge
below the ocellus ; preocular ridge with a well defined ventral rudiment. Preoral ridge slender.
Eyes : Dorsal simple eyes not projecting beyond the outer margins of the head in dorsal view ;
their corneae 21-27 (24)0 in diameter and separated by 1 13-134 (125)0, i.e. 4-6-6-1 (5-1) times
their corneae apart. Ventral simple eyes 24-34 (3°)^ m diameter and 24-31 (27)0 apart.
Lateral ocelli well developed. Cranial apophysis with truncate apex. Tentorial bridge slender.
Dorsal head setae 13-16 (14-7) on each side anterior to the postoccipital ridge ; each gena
usually with 4 and occasionally 3 or 5 genal setae (av. 3-8). Ventral head setae on each side :
1-2 (i-8) between the ventral eyes ; 11-16 (12-4) forming on both sides a transverse band in the

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 127

area of the ventral preocular depression ; 3-4 (3-8) on each side of the ventral arm of the
medcranial ridge. Head disc pores : Dorsal pores usually absent, although one pore was found
on one side of a few specimens near the base of the antennae (av. 03).

Antennae : Filiform ; 10-segmented ; 610-769 (705)^1 long, i.e. subequal to half the total
length of the body (ratio 1 : 1-8-21, av. 19), and subequal to the hind legs (ratio 1 : o-8-i-i,
av. 095). Scape 46-55 (52) u. long and 46-49 (47) y. wide at base ; always with 4 h.s. Pedicel
61-70 (66)(i long and 37-43 (40)11 wide ; with 19-29 (231) f.s., 10-14 ( I2- i) h.s. an d a sensillum
placodeum. Flagellum : Segment II I club-shaped, about as long as segment X, both being the
longest (ratio approximately 1:1); segment III about 29(1 wide, the ratio its width to its
length being 1 : 2-5-3-2 (29). Segments IV to X cylindrical and 24-27(1 wide, the ratio width
to length of segment IX being 1 : 2-4-2-9 (2-7). The lengths of the flagellar segments and the
number of setae on each are given in the following table :

III IV V VI VII VIII IX X

length in |i

70-92

61-76

58-79

58-89

61-89

58-76

58-73

76-92

(av.)

(82)

(67)

(70)

(73)

(73)

(70)

(67)

(82)

f.s.

13-21

16-24

19-30

21-33

21-31

20-26

17-24

16-27

(av.)

(170)

(18-8)

(23-9)

(25-9)

(257)

(22-6)

(21 -O)

(20-1)

h.s.

5-"

2-5

2-5

3-5

- 5

3-6

2-5

1-2

(av.)

(7-4)

(2-6)

(3-9)

(4-3)

(37)

(4-2)

(3-D

(i-7)

Antenna! bristles well-defined ; se£

'mentsVIII and IX

each \\ ith

one ventral bristle.

Terminal

segment with 3 preapical bristles and two much smaller ones near the base of the segment ;
terminal segment also with a pair of capitate subapical sensory setae ami one apical hair-like seta.

Thorax : 532-707 (722)11 long. Prothorax : Pronotal ridgis with the usual median interrup-
tion. Lateral pronotal sclerites and post-tergites small. Proepisternum with a ridge-like dorsal
margin. Prosternum 43-58 (52) (i long and comparatively narrow ; posteriorly bounded by
the transverse, 92-110 (104) [i long prosternal ridge. Prothoracic setae on each side : Medial
pronotal setae 1-2 (17) ; lateral pronotal setae 1-3 (21) ; post-tergital setae almost always
absent, but one seta was found on each side of one specimen (av. 01 h.s.) ; antespiracular
dorsal setae 2-4 (2-9). One antespiracular ventral seta always present. Prosternal setae 2-4
(2-5). Prothoracic pores on each side : Medial and lateral pronotal pores 1-3 (averages 2-2 and
2-1 respectively) ; antespiracular dorsal pores 1-2 (i-i). Prosternal pore usually one, but
sometimes two or none at all on either side (av. 11).

Mesothorax : Prescutum 92-119 (io4)ji long and 131-168 (153)11 wide (ratio 1 : 1-4-1-7,
av. 1-5) ; laterally bounded by the prescutal ridges and posteriorly by the prescutal suture.
Scutum heavily sclerotized antero-laterally, with a medial longitudinal narrow membranous
area ; 110-137 (i28)(i long, i.e. the ratio lengths of prescutum to scutum 1 : 1-1-1-3 ( I-2 ).
Prealare, prealar ridge, triangular plate, anterior and posterior nolal processes well developed.
Scutellum 61-73 (70)11 long and 98-125 (n6)(i wide, the ratio being 1 : 1-6-1-7 (i'°5) and the
ratio lengths of scutellum to scutum 1 : 1-7-2-0 (i-8). Postalare with well separated anterior
and posterior ridges.

Mesopleuron : Mesopleural ridge with a short interruption above the coxal articulation ;
basalare stout and subepisternal ridge well developed. Later opleurita narrow. Mesosternum :
Basisternum 162-204 ( x 86)|i long and 168-235 (207)11 wide ; marginal and precoxal ridges well
developed. Mesothoracic spiracle about 2 1 (i wide at opening, with a 40-46 (43) (i long supporting
bar. Mesothoracic setae on each side : Prescutal setae 4-7 (5-7) ; scutal setae 8-12 (io-i) ;
scutellar setae 1-2 (i-i). Tegular setae 3-5 (4-2). Postmesostigmatal setae in two separable
groups, a latero-ventral cluster of 8-12 (9-6) setae behind each spiracle, and 1-3 (1-7) setae
submedially, forming with their partners of the other side a transverse irregular row. Basi-
sternal setae 10-16 (12-7). Mesothoracic disc pores on each side : Mesospiracular pores 1-2
(1-5) ; one postmesostigmatal pore usually present submedially, but sometimes absent on one
side (av. o-8).

128

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Metathorax : Metapostnotal sclerites and metapostnotal ridge well developed. Metapleural
ridge attenuated at the position of the pleural apophysis ; with a small metapleural wing process.
M etepisternum and metepimeron well defined. Precoxal ridge strong and metasternal apophysis
distinct. Metathoracic spiracles similar to the mesothoracic ones. Metathoracic setae on each

dmep

Fig. 19. Trionymus newsteadi (Green), dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

129

side : Metatergal setae 1-3 (17) ; metaplenral setae 1-2 (i-i) ; anterior metasternal setae 3 and
posterior metasternal setae 2-3 (2-2). Metathoracic pores : Metaspiracular pores 2-3 (2-3) behind
each spiracle ; one anterior metasternal pore, and one posterior metasternal pore almost always
present on each side (averages 08 and 09 respectively).

Wings : 1330-1610 (1484)^1 long and 462-532 (504) u. wide ; usually with 3 alar setae
(sometimes 4 or 5, av. 3-4), and a row of 4 minute circular sensoria. Hamnlohalterae 104-122
(no)(i long and 21-31 (25)^ wide ; with one apically hooked, 46-55 (52)(x long seta, i.e. the
ratio lengths of the seta to the hamulohaltera 1 : 1-9-2-5 (21).

Legs : Moderately long and slender ; the ratio length of the hind leg to the total length of
the body 1 : 1-7-2-1 (2-0). Coxa and trochanter about 49 and 24^ wide respectively ; the
latter with one differentiated long apical seta. Femur about 43^ wide ; that of the front leg
shortest and that of the hind longest ; the ratio width to length of the hind femur 1 : 41-50
(4-6). Tibia about 24JJ. wide ; with 2 apical strong spurs and 3-4 smaller spines ; the ratio
lengths of femur to tibia in front leg 1 : 10-11 (105). Tarsus about 2i|z wide ; tarsal
digitules apically knobbed, about 37 y. long. Claw uniformly tapering to a pointed apex ;
ungual digitules extremely fine. In the following table the lengths of the leg segments (in
microns) and the number of setae on each are given :

Fore leg

Middle leg

Hind leg

Coxa

length
h.s.

43-49 (4 6 )
n-14 (122)

43-49 (4 6 )
9-13 (114)

46-52 (49)
10-15 (126)

Trochanter

length
h.s.

58-70 (64)
4-6 (5-1)

58-70 (64)
4-6 (50)

64-73 (70)
4-6 (5-2)

Femur

length
h.s.

153-189 (174)

17-25 (20-2)

159-201 (183)
10-26 (22-2)

168-220 (198)

19-25 (22-8)

Tibia

length
h.s.

168-198 (183)

24-35 (28-6)

174-229 (204)
23-33 (28-2)

223-278 (253)
25-37 (30-6)

Tarsus

length
h.s.

76-92 (85)
15-19 (17-2)

76-92 (85)
13-20 (166)

82-IO4 (95)
I5-20 (17-4)

Claw

length

27-31 (29)

27-31 (29)

27-31 (29)

Total length

of leg

525-619 (580)

537-668 (613)

610-753 (692)

Abdomen : 365-532 (456) u. long and 266-327 (312) \l wide. Tergites of segments I and II
small ; those of segment VIII and segments IX + X large and distinct. Sternites of segment
VIII ill-defined. Ostioles entirely absent. Abdominal setae on each side : Dorsal setae 2-3
on segment I, 3-4 on segments II to VI, 2-3 on segment VII, and always 1 on segment VIII.
Pleural setae 3-4 on segment I, 4-5 (and occasionally 6) on segments II to VII, and always 3
on segment VIII. Ventral setae 1-2 on segment II, 3-4 on segments III to V, 2-3 on segment
VI, 2 on segment VII, and absent on segment VIII. Abdominal disc pores on each side :
Pleural setae 2-5 (3-7), 0-2 (i-i), 0-1 (0-5), 0-1 (o-6), 0-1 (0-7), 0-1 (0-5), and 0-2 (0-7) on
segments I to VII respectively. One ventral pore also usually occurs submarginally on
segments III to VII (averages o-8, 07, 07, 0-3, and 02 respectively).

Glandular pouch well developed ; its setae include a pair of tail setae, 229-305 (272) (x long,
and one seta, 40—92 (64)^ long (the ratio lengths of the tail setae to the total body length
1 : 4-6-5-7, av. 50).

Genital segment comparatively small ; subtriangular in dorsal view ; style curving upwards
in lateral view. Penial sheath 113-131 (125) \l long and 79-95 (89) y. wide, i.e. the ratio length

13°

MORPHOLOGY AND TAXONOMY OF ADULT MALES

to width 1-3-1-4 (1-38) : 1, and the ratio its length to the total body length 1 : 10-2-12-0 (n-o).
Basal ridge of penial sheath and its projection well developed ; process of penial sheath well
pronounced. Aedeagus gradually pointed. Setae of genital segment : Dorsally, 3 setae always
present on each side near the base of the style ; ventrally 3-4 (3-7) setae occur on each side
of the penial sheath, and 4-5 (4-3) setal sensilla on its process.

Material : 10 specimens examined, collected by myself, on beech trees (Fagus
sylvatica), in the Imperial College Field Station, Silwood Park, Sunninghill, Berk-
shire, England, iii-iv.1964.

Remarks : Although the present study is mainly concerned with morphology and
taxonomy, some biological observations were also made. According to Blaire
(1958), the second stage larvae of this species, usually males, migrate from the
small twigs towards the main branches and trunk of the trees between February
12th and April 10th ; however, the writer was able to produce adult males from
larvae collected during the last two weeks of April. It was found that the second
stage larvae take about 48 hours to reach the third stage, and the latter about
another 48 hours to become prepupae ; the prepupae take about 7 days to develop
to pupae, and the pupae about 8-10 days to produce males with well developed
antennae, legs and wings, but still in a stationary condition or with very limited
movement. The males became active after about 48 hours, after which the long
waxy tassels covering the abdominal segment VIII were produced ; they were
then ready to emerge from their puparia and search for the females. The males
mated 5-10 times with females introduced to them in the same glass tube ; each
copulation usually lasts 1^—3 minutes and may be repeated with the same female
or another one at variable intervals from several minutes to a few hours. The
males fly in rapid and rather short jumps (a few inches), and all died within 72 hours
from their emergence.

200^-

100^-

Fig. 20. Trionymus newsteadi (Green), lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 131

PSEUDOCOCCUS Group

PSEUDOCOCCUS Westwood, 1840

Pseudococcus obscurus (Essig)

(Text-figs. 21, 22)

The winged forms only known ; living specimens not available. The males are narrow
and slender, of medium or large size and with comparatively short antennae and moderately
long legs. When mounted, the total length of the body 1 106-1470 (1330)11, the width at
mesothorax 280-350 (322)0. and the wing expanse 1932-2520 (2240) u..

Body setae and pores : Fleshy and hair-like setae present on the body, antennae and legs ;
all subequal in length, about 24^ long. Quadrilocular and quinquelocular disc pores about 6[i
in diameter, occurring on the head, thorax and abdomen.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression well pronounced in lateral view. Length from apex to postoccipital ridge 125-153
(137)". ; from apex to neck 153-192 (i8o)u. ; width across the genae 180-220 (207)^1. Dorsal
arm of midcranial ridge anteriorly separated from the other arms by. a short distance, and
posteriorly meeting the postoccipital ridge. Ventral and lateral arms forming together a
Y-shaped ridge. Postoccipital ridge not reaching the preocular ridges anteriorly. Dorsomedial
part of epicranium well sclerotized. Preocular + interocular ridges join the poslocular ridge
below the ocellus ; ventral rudiment of preocular ndfi;e hardly indicated. Preoral ridge slender.
Eyes : Dorsal simple eyes projecting beyond the outer margins of the head in dorsal view ;
their corneae 24-27 (25)11 in diameter and separated by 92-113 (104)11, i.e. 3-8-4-6 (43) times
their corneae apart. Ventral simple eyes 27-31 (29) (z in diameter and 24-31 (27) u. apart.
Lateral ocelli large. Cranial apophysis apically truncate. Tentorial bridge slender. Dorsal
head setae : 5-8 (6-6) f.s. and 7-1 1 (96) h.s. present on each side of the median line anterior
to the postoccipital ridge ; each gena with 13-18 (152) fleshy and 3-4 (33) hair-like genal
setae. Ventral head setae on each side : 3 4 (33) h.s. between the ventral eyes ; 5-9 (7-1)
f.s. and 2-4 (3-2) h.s. forming on both sides a transverse band in the area of the ventral preocular
depression ; ventral part of ocular sclerite with 5-1 1 (8-5) fleshy ventral ocular setae ; anteriorly,
3 _ 4 (3'3) ns - longitudinally arranged on each side of the ventral arm of the midcranial ridge.
Head disc pores : Dorsal head pores 2-3 (22) on each side near the base of the antennae.

Antennae : Filiform ; normally 10-segmented ; 616-668 (647)11 long, i.e. as long as, or
somewhat shorter than half the body length, and slightly shorter than the hind legs, the ratios
1 : 2-0-2-2 (21) and 1 : 11-12 (112) respectively. Scape 43-62 (46)11 long and 43-46 (45)11
wide at base ; with 6-8 (67) h.s. Pedicel 61-73 (70)11 long and 31-37 (34)11 wide ; with 11-18
(14-7) f.s., 9-16 (12-1) h.s., and a sensillum placodeum. Flagelhun : Segment III club-shaped,
longest of all and about 24(1 wide ; the ratio lengths of segments III to X 1-4-1-5 (142) : 1,
and the ratio width to length of segment III 1 : 3-7-4-6 (4-3). Segments IV to X cylindrical
and about 22 u. wide ; the ratio width to length of segment X being 1 : 2-4-2-9 (2-6). The
following table shows the lengths of the flagellar segments and the number of setae on each :

III IV V VI VII VIII IX X

lengths in (i

79-98

6i-73

6i-73

58-64

55-67

52-67

52-61

61-70

(av.)

(92)

(67)

(67)

(61)

(61)

(58)

(58)

(67)

f.s.

9-14

8-i 6

12-16

8-16

11-16

10-17

8-14

10-13

(av.)

(n-3)

(12-4)

(I4-I)

(n-7)

(i3-o)

(12-9)

(n-i)

(107)

h.s.

4-7

2-6

2-4

2-5

2-5

3-5

3-8

5-7

(av.)

(5-i)

(3-5)

(3-3)

(2-7)

(3-i)

(3-6)

(5-4)

(6-3)

Antennal bristles : Segments VIII and IX each with one ventral bristle ; segment X with
3 subapical bristles, a pair of capitate subapical sensory setae, and one apical hair-like seta.

132

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Imcr SC P

Fig. 21. Pseudococcus obscurus (Essig), dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

133

Thorax : 426-570 (524)^1 long. Prothorax : Pronotal ridge medially interrupted at a weak
point. Lateral pronotal sclerites and post-tergites comparatively large. Proepisternum with a
ridge-like dorsal margin. Prosternum triangular, 37-49 (43) \t long ; prosternal ridge 85-116
(ioi)jjl long. Prothoracic setae on each side of the median line : Medial pronotal setae 0-2
(09) f.s. and 0-3 (1-3) h.s. ; lateral pronotal setae 0-1 (0-3) h.s. ; post-tergital setae 3-6 (4-6) f.s. ;
antespiracular dorsal setae 0-2 (0-5) h.s. One, hair-like antespiracular ventral seta always
present. Prosternal setae 1-3 (2-1) f.s. and 1-3 (i-8) h.s. Prothoracic disc pores on each side :
Medial pronotal pores 2-4 (26) ; lateral pronotal pores 2-4 (3-1) ; antespiracular dorsal pores
2-4 (2-1). Prosternal pores 1-3 (2-1).

Mesothorax : Prescutum 73-92 (79)0. long and 113-146 (i3i)(x wide, the ratio being 1 : 15-
i-8 (1-7) ; prescutal ridge well developed and prescutal suture well defined. Scutum heavily
sclerotized antero-laterally, with a longitudinal median narrow membranous area. Scutum
101-128 (ii6)[a long, i.e. the ratio lengths of prescutum to scutum 1 : 1-3-1-6 (1-5). Scutellum
46-61 (55) [i long and 89-119 (107)^1 wide, the ratio being 1 : 1-8-2-1 (1-9), and the ratio its
length to the length of scutum 1 : 2-0-2-2 (21). Anterior and posterior postalar ridges well
separated ; postnotal apophysis well developed. Mesopleuron : Mesopleural ridge interrupted
above the articulation with coxa ; pleural apophysis, pleural wing process, basalare and
subepisternal ridge well developed ; later opleurite narrow, mesepisternum and mesepimeron
distinct. Mesosternum : Basisternum 128-171 ( 1 50) \i long and 168-271 (192)^1 wide ; marginal,
precoxal ridges and furca strongly developed. Mesothoracic spiracle 21-24 l 22 )^ wide at
opening, with a 34-43 (40) u. long supporting bar. Mesothoracic setae on each side : prescutal
setae 4-6 (4-7) h.s. ; scutal setae 4-7 (52) h.s. ; sculellar setae 2-4 (26) h.s. Tegular setae
3-5 (3-7) h.s. ; postmesostigmatal setae 1-3 (i-6) f.s. and 2-4 (2-6) h.s. occurring only laterally
behind each spiracle. Basisternal setae 7-12 (9-1) h.s. Mesothoracic disc pores : 1-2 (i-6
mesospiracular pores posteriorly associated with each spiracle.

Metathorax : Metapostnotal sclerites and metapostnotal ridge well developed. Pleural ridge,
pleural wing process, pleural apophysis, episternum and epimeron typical of the family. Precoxal
ridge and metasternal apophysis well developed. Metathoracic spiracle identical with the
mesothoracic.

Metathoracic setae on each side : Metatergal setae 1-2 (1-2) f.s. and 3-4 (3-3) h.s. in a sub-
median cluster ; metapleural setae 1-2 (i-i) f.s. behind the metathoracic spiracles. Anterior
metasternal setae 1-2 (1-5) f.s. and 1-3 (21) h.s. ; posterior metasternal setae absent. Meta-
thoracic disc pores on each side : Metaspiracular pores 0-1 (0-7) ; anterior metasternal pores
1-2 (i-6) ; posterior metasternal pores absent.

200 -^

Fig. 22. Pseudococcus obscurus (Essig), lateral view.

134 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Wings : 840-1092 (980)11 long and 392-476 (434)11 wide ; with 3 alar setae and 2 circular
sensoria. Hamulohalterae 76-85 (79) |i long and 15-21 (i8)(x wide ; with one, 55-61 (58)11
long apically hooked seta, i.e. the ratio length of seta to the length of the hamulohaltera about
1 : 14.

Legs : Moderately long ; the ratio length of the hind leg to the total length of the body
1 : 1-6-1-8 (1-7). Coxa and trochanter about 52 and 27(1. wide respectively. Femur about 43(1
wide ; that of fore leg shortest and that of hind leg longest ; the ratio width to length of the
hind femur being about 1 : 5-4. Tibia about 24^ wide ; with 2 apical stout spurs and 3-5
smaller ones ; the ratio lengths of femur to tibia in fore leg 1 : 1-1-1-2 (1-16). Tarsus about
21 (x wide ; tarsal digitules apically knobbed, about 34^ long. Claw uniformly tapering to a
pointed tip ; ungual digitules finely pointed. In the following table, the lengths of the leg
segments (in microns) and the number of setae on each are given :

Fore leg

Middle leg

Hind leg

Coxa

length

f.s.
h.s.

40-46 (43)
6-11 (9-8)
4-9 (67)

40-46 (43)
7-12 (8-5)
4-8 (6-2)

43-49 (46)
9-13 (n-8)
6-8 (6-8)

Trochanter

length

f.s.

h.s.

67-70 (68)
i-3 (2-2)
3-6 (4-5)

67-70 (68)
i-4 (2-3)
3-6 (4-2)

70-73 (71)
2-5 (3-o)
4-7 (4"8)

Femur

length

f.s.

h.s.

189-201 (192)

23-30 (28-2)

9-12 (9-8)

195-204 (198)

21-29 (26-8)

7-11 (8-2)

207-220 (211)

27-38 (32-2)

9-13 (100)

Tibia

length

f.s.

h.s.

214-235 (223)

34-48 (42-0)

4-7 (5-3)

232-253 (241)

45-57 (5o-8)
4-7 (5-2)

244-305 (281)

58-69 (64 3)

4-8 (5-8)

Tarsus

length

f.s.

h.s.

85-92 (89)
17-23 (201)
3-6 (4-2)

85-92 (89)
22-27 ( 2 4'9)
3-6 (4-3)

92-104 (98)

20-29 (26-2)

4-7 (5-3)

Claw

length

24-30 (27)

24-30 (27)

24-30 (27)

Total length

of leg

622-659 (634)

644-683 (659)

677-772 (732)

Abdomen : 403-578 (486) (i long and 251-327 (296) jx wide. Tergites of segments I and II
small ; those of segments VIII and IX + X large and distinct. Sternites of segment VIII
ill-defined. Ostioles weakly developed. Abdominal setae on each side : Dorsal setae : Fleshy
setae 0-1 on segment I, 2-3 on segment II, 1-3 on segment III, and 1-2 on segments IV to
VIII ; hair-like setae 2-4 on segment I, 3-4 on segments II to V, 2-4 on segments VI and VII,
and 1-2 on segment VIII. Pleural setae 2-4 h.s. on segment I, 4-5 on segments II to VII,
and 3-4 on segment VIII, including one slightly longer seta. Ventral setae : Fleshy setae
0-1 on segments II, III and IV ; hair-like setae usually 2-3 on segments II to VII, and
occasionally 1 or 4 ; ventral setae absent on segment VIII. Abdominal disc pores : Segments
I-VII with 6-11 (8-6), 1-2 (i-6), 1-2 (1-5), 1-2 (1-5), 1-3 (1-9). 2-4 (2-5), and 3-5 (4-3) pleural
pores, respectively.

Glandular pouches well developed ; setae of glandular pouch include two, 360-412 (387)11
long tail setae and one, 58-92 (76) [i long seta, the ratio length of the tail setae to the total length
of the body being 1 : 3-0-3-8 (3-4).

Genital segment : Subtriangular in dorsal view, with the style comparatively broad and
apically rounded ; the latter curving upwards in lateral view. Penial sheath 153-168 (162) ji
long and 76-89 (85)11 wide, i.e. the ratio its length to its width 1-8-2-0 (1-9) : 1, and its length
to the total length of the body 1 : 7-2-8-9 (8-2). Basal ridge of penial sheath with a small
projection ; process of penial sheath absent. Aedeagus tapering to a pointed tip. Setae of

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 135

genital segment : Dorsally, 3 small setae always present on each side near the base of the style ;
ventrally 4-5 (44) similar setae, and 4-5 (43) minute setal sensilla occur on each side of the
penial sheath.

Material : 10 specimens examined, obtained by K. Boratynski in a laboratory
culture on Potato, in London during vi.1961.

Remarks : Wilkey and McKenzie (1961) attempted to reveal the long puzzling
identity of P. maritimus and whether more than one species of mealybug is involved
under the so-called P. maritimus-malacearum complex. They carried out a
morphological investigation on the females primarily based on the shape of the
hind legs and the number of the translucent dots on the leg segments ; this investiga-
tion led to the conclusion that P. maritimus and P. obscurus are distinct species,
P. bakeri Essig and P. omniverae Hollinger being synonyms of the former, and
P. capensis Brain, P. longispinus var. latipes Green and P. malacearum Ferris of the
latter. Accordingly, Beardsley (1963) realized that the species long believed to
be P. maritimus in Hawaii, whose males were described as such by him in an earlier
paper (i960), was in fact obscurus.

Beardsley's description of this species indicates that it is identical with P. obscurus
here studied, and both entirely agree with " P. maritimus Ehrhorn, type B "
described by Giliomee (1961) ; therefore, "P. maritimus, type A" at present is
uncertain and may perhaps be the true P. maritimus.

Pseudococcus citriculus Green

(Text-figs. 23, 24)

Winged forms only known ; living specimens not available. The males are comparatively
slender, of medium or large size, with short antennae and moderately long legs. When
mounted, total body length 1232-1400 (1302)0, width at mesothorax 280-322 (294)1* ar, d
wing expanse 2016-2338 (2128)0.

Body setae and pores : The body antennae and legs with fleshy and hair-like setae, about
27|jl long. Quadrilocular and quinquelocular disc pores present on the head, thorax and
abdomen, 6-90 in diameter.

Head : Subtetrahedron ; triangular in dorsal and front views ; ventral preocular depression
well pronounced in lateral view. Length from apex to postoccipital ridge 122-140 (128)0 ;
from apex to neck 171-214 (186)0 ; width across the genae 201-229 ( 2I 4) U - Dorsal arm of
midcranial ridge anteriorly detached from other arms, and posteriorly meeting or almost
meeting the postoccipital ridge. Ventral and lateral arms forming together a Y-shaped ridge.
Postoccipital ridge not reaching the preocular ridges anteriorly. Preocular -f- interocidar
ridges well developed ; ventral rudiment of the former hardly indicated by a weak sclerotization.
Preoral ridge slender. Eyes : Dorsal simple eyes not projecting beyond the outer margins of
the head in dorsal view ; their corneae 21-24 ( 2 3)M- m diameter, and separated by 92-119
(104)0., i.e. 3-8-5-0 (4-3) times their corneae apart. Ventral simple eyes 24-27 (26)0. in diameter,
and 18-37 l 2 ?)^ apart. Lateral ocelli well developed. Cranial apophysis with truncate apex ;
tentorial bridge slender. Dorsal head setae : 8-1 1 (9-3) f.s. and 9-12 (10-5) h.s. on each side
anterior to the postoccipital ridge ; each gena with 16-23 ( I 9'°) fleshy and 2-4 (3-0) hair-like
genal setae. Ventral head setae on each side : 2-3 (1-3) h.s. between the ventral eyes ; 7-10
(8-8) f.s. and 2-5 (3-3) h.s. forming on both sides a transverse band in the area of the ventral
preocular depression ; ventral part of ocular sclerites with 4-7 (3-8) fleshy ventral ocular setae ;

136

MORPHOLOGY AND TAXONOMY OF ADULT MALES

anteriorly, o-i (0-2) f.s. and 3-4 (3-5) h.s. on each side of the ventral arm of the midcranial
ridge. Head disc pores : 1-3 (i-6) dorsal head pores occur on each side near the base of the
antennae.

Antennae : Filiform ; normally 10-segmented ; 540-604 (570)^ long, i.e. slightly shorter
than half the body length, the ratio being 1 : 2-2-2-3 (2-28), and shorter than the hind legs.

.. dhs l^cr
ohp , / vprd

Fig. 23. Pseudococcus citriculus Green, dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

137

the ratio 1 : 1-1-1-3 (113). Scape 46-52 (49)|x long and 40-43 (42)0. wide at base ; with
6-9 (73) h.s. Pedicel 58-64 (6i)u. long and 34-37 (35)(x wide ; with 17-22 (200) f.s., 10-14
(12-0) h.s. and a sensilhim placodeum. Flagellutn : Segment III club-shaped, longest of all
and about 24^ wide ; the ratio lengths of third and terminal segments being 1-3-1-6 (1-4) : 1,
and the ratio width to length of segment III 1 : 3-3-4-3 (3-7)- Segments IV-X cylindrical
and approximately 2 2|j. wide ; the ratio width to length of segment IX 1 : 2-0-2-3 (2-1). The
following table shows the lengths of the flagellar segments and the number of setae on each :

III IV V VI VII VIII IX X

length in u.

70-92

52-55

52-58

52-58

52-58

49-58

49-55

55-58

(av.)

(79)

(54)

(55)

(56)

(56)

(54)

(53)

(57)

f.s.

11-14

15-19

12-16

12-15

10-16

6-12

8-10

6-8

(av.)

(12-3)

(17-0)

(14-0)

(13-5)

(12-5)

(io-3)

(8-8)

(7-0)

h.s.

4-7

3-4

2-4

3-4

5-7

6-7

7-9

7-8

(av.)

(5-3)

(3-3)

(3-o)

(3-5)

(55)

(6-8)

(8-o)

(7-3)

Antennal bristles easily distinguishable ; segments VIII and IX each with a ventral bristle.
Terminal segment with 3 preapical bristles, a pair of capitate subapical sensory setae and one
apical hair-like seta.

Thorax : 494-608 (547) y. long. Prothorax : Pronotal ridge medially interrupted as usual.
Lateral pronotal sclerites and post-tergites large. Dorsal margin of proe pi sternum ridge-like.
Prosternum triangular, 31-40 (34) [z long ; posteriorly bounded by the transverse, 92-101
(98) [i long prosternal ridge. Prothoracic setae on each side : Medial pronotal setae 2-4 (33) f.s.
and 1-3 (27) h.s. ; lateral pronotal setae 0-1 (09) h.s. ; post-lergital setae 10-13 ( IX "5) IS - anc >
1-2 (1-7) h.s. ; antespiracular dorsal setae 0-2 (1-3) h.s.
h.s. Prosternal setae 3-5 (43) f.s. and 1-2 (13) h.s.
Medial pronotal pores 2-4 (2-5) ; lateral pronotal pores 4-5 (4-3)
1-3 (2-0). Prosternal pores 2-4 (3-3).

Mesothorax : Prescutum 67-82 (73)^ long and 122-137 ( I2 8)(i wide, i.e. the ratio 1 : i-6-
1-9 (17) ; prescutal ridge strong and prescutal suture distinct. Scutum heavily sclerotized
antero-laterally, and with a longitudinal median narrow membranous area. Scutum 1 10-128
(ng)n long, the ratio lengths of prescutum to scutum being 1 : 1-5-1-8 (i-6). Prealare, prealar
ridge and triangular plate well developed. Sculellum 52-58 (55) (x long and 101-116 (ioj)\l
wide, the ratio being 1 : 1 -9-2-0 (194), and the ratio its length to the length of scutum 1 : 2— 1

Antespiracular ventral setae 1-2 (1-3)
Prothoracic disc pores on each side :
antespiracular dorsal pores

200^

Fig. 24. Pseudococcus citriculus Green, lateral view.

i3»

MORPHOLOGY AND TAXONOMY OF ADULT MALES

2-2 (2-17). Anterior and posterior postalar ridges well separated. Mesopleuron : Mesopleural
ridge interrupted above the coxal articulation ; mesopleural apophysis and pleural wing process
well developed ; basalare strong. Subepisternal ridge, mesepisternum, mesepimeron and latero-
pleurite typical of the family. Mesosternum : Basisternum 131-156 (143)11 long and 168-198
(177)1* wide ! marginal and precoxal ridges well developed. Mesothoracic spiracle 18-21 (20)0. wide
at opening, with a 34-40 (37)11 long supporting bar. Mesothoracic setae on each side : Prescutal
setae 5-7 (6-o) h.s. ; scutal setae usually in two groups, the anterior including 3-5 (3-3) h.s. and
the posterior with 4-5 (4-3) slightly longer setae ; scutellar setae 2-3 (2-7) h.s. Tegular setae
3 — 4 (3'3) ns - '• postmesostigmatal setae 13-19 (16-8) f.s. and 1-4 (2-8) h.s. forming on both sides
a transverse band. Basisternal setae 10-13 ( II- 5) ns - Mesothoracic disc pores : Mesospiracular
pores 0-1 (0-5) behind each spiracle ; one postmesostigmatal pore always present medially.

Metathorax : Metapostnotal sclerites and metapostnotal ridge well developed ; pleural ridge,
pleural wing process, mesepisternum and mesepimeron well developed. Precoxal ridge and
metasternal apophysis strong. Metathoracic spiracle identical with the mesothoracic. Meta-
thoracic setae on each side : Metatergal setae 2-5 (3-8) f.s. and 3-4 (3-7) h.s. ; metapleural setae
8-10 (8-8) f.s. and 0-2 (0-5) h.s. ; postmetastigmatal setae 3-4 (3-5) f.s. Anterior metasternal
setae 7-12 (9-3) f.s. and 2-3 (2-6) h.s. ; posterior metasternal setae 0-1 (o-8) f.s. and 0-1 (0-5)
h.s. Metathoracic disc pores : Metaspiracular pores absent ; anterior and posterior metasternal
pores 0-1 (o-8) and 0-1 (0-5) on each side, respectively.

Wings : 882-1022 (938) (x long and width 350-420 (378) (i wide ; with 3-4 (3-2) alar setae
and a compact row of 3-4 (3-2) minute circular sensoria. Hamulohalterae 67-82 (76)11 long
and 15-18 (i7)u. wide ; with one, 43-49 (46) apically hooked seta, i.e. the ratio lengths of the
seta to the hamulohaltera 1 : 1-4-1-8 (1-7).

Legs : Moderately long ; the ratio length of the hind leg to the total length of the body
1 : 2-0-2-1 (2-02). Coxa and trochanter about 48 and 24(1 wide respectively. Femur about
400. wide ; that of the fore leg usually shortest but sometimes equal to that of the middle leg ;
the ratio width to length of the hind femur about 1 : 47. Tibia about 24^ wide ; with 2
apical stout spurs and 4-6 smaller spines ; the ratio lengths of femur to tibia of the fore leg
1 : i-i-i (1-03). Tarsus about 2ifi wide ; with a pair of apically knobbed tarsal digitules.
Claw gradually tapering to a pointed end ; with a pair of finely pointed ungual digitules. The
lengths of the leg segments (in microns) and the number of setae on each are given in the
following table :

Fore leg

Middle leg

Hind leg

length

40-46 (43)

40-46 (43)

43-49 (4 6 )

Coxa

f.s.

8-16 (12-0)

9-15 ("-3)

10-17 (i3-o)

h.s.

6-10 (7-8)

7-10 (8-3)

7-9 (8-3)

length

58-61 (59)

58-61 (59)

61-67 (64)

Trochanter

f.s.

4-5 (4-3)

4-5 (4-3)

4-5 (4-8)

h.s.

4-6 (5-0)

4-7 (5-3)

4-6 (4-8)

length

168-183 (174)

171-186 (177)

177-195 (186)

Femur

f.s.

29-34 (3i-o)

28-35 (32-o)

36-48 (40-0)

h.s.

8-13 (100)

8-12 (9-3)

7-14 (9-8)

length

183-192 (189)

195-207 (201)

229-244 (238)

Tibia

f.s.

33-43 (39-o)

38-46 (41 -8)

48-60 (53 0)

h.s.

6-8 (7-0)

6-10 (8-3)

8-10 (9-0)

length

73-79 (76)

73-79 (76)

76-82 (79)

Tarsus

f.s.

10-16 (140)

10-16 (13-3)

12-18 (15-0)

h.s.

8-12 (9-8)

8-13 (10-5)

8-14 (n-3)

Claw

length

24-30 (27)

24-30 (27)

24-30 (27)

Total length

of leg

558-577 (567)

567-595 (583)

628-656 (644)

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 139

Abdomen : 403-433 (4i8)[i long and 266-304 (281)11 wide. Tergite of segment I small ;
those of segments VIII and IX + X large. Sternites of segment VIII ill-defined. Ostiohs
large and prominent, 43-49 (45) \jl long at orifice. Abdominal setae on each side : Dorsal setae :
Fleshy setae 2-3 on segments I to III, 3-4 on segments IV to VI, and 2-4 on segments VII
and VIII ; hair-like setae 2-3 on segment I, 3-4 on segments II to V, 2-3 on segments VI and
VII, and 2-4 on segment VIII. Pleural setae : Fleshy setae 2-5 on segment I, usually 1-2
on segments II to VII (sometimes 3 on segments II and V), and 4-6 on segment VIII ; hair-
like setae 1-3 on segment I, 4 on segment II, usually 5 on segments III to VII (sometimes
6 on segment VI), and 3-5 on segment VIII, including one conspicuously longer seta. Ventral
setae : Fleshy setae on segments II and III, 3-5 on segment IV, 3-4 on segment V, 2-3 on
segments VI and VII, and absent on segment VIII. Abdominal pores : Segments I to VII
with 3-6 (4-3), 1-2 (1-3), 1-2 (1-3), 1-2 (1-5), 1-2 (1-5), 1-3 (i-8), and 3-4 (3-8) pleural pores
respectively.

Glandular pouch well developed ; seta of glandular pouch comprise a pair of 336-366 (354)^1
long tail setae and one 64-79 (73) \i long seta ; the ratio length of the tail setae to the total
length of the body 1 : 34-40 (37).

Genital segment : Comparatively large ; triangular in dorso-ventral view. Style more or
less straight in lateral view ; apically rounded. Penial sheath 183-214 (ig8)(i. long and 104-
107 (io5)(jl wide, i.e. the ratio length to width 17-21 (19) : 1, and the ratio its length to the
total length of the body 1 : 6-3-6-9 (6-6). Basal ridge of penial sheath and its projection well
developed ; process of penial sheath absent. Aedeagus comparatively long and slender,
anteriorly reaching the cavity of abdominal segment VII and tapering to a sharply pointed tip.
Setae of genital segment on each side : Dorsally, 3 small setae always present near the base of
the style ; ventrally, 10-15 ( I2 '3) much finer setae scattered on the penial sheath.

Material : 4 specimens only were available, collected by C. R. Wallace, on Citrus,
in Sarawak, xi.1963 (received from Dr. D. J. Williams).

DYSMICOCCUS Ferris, 1950

Dysmicoccus alazon Williams

(Text-figs. 25, 26)

Only the winged forms known. These are greenish brown with dark red eyes. A relatively
long and slender species, with short antennae and moderately long legs. . Mounted specimens
1274 — 1470 (1358)11 long, 280-322 (294) [x wide at mesothorax and 2632-3094 (2800) \i wing
expanse.

Body setae and pores : Body and appendages with numerous fleshy and hair-like setae,
about 24JJ. long. Cjuadrilocular and quinquelocular disc pores occur on the head, thorax and
abdomen, 6-9 (x in diameter.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression well pronounced in lateral view. Length from apex to postoccipital ridge 116-134
(i25)(x ; from apex to neck 168-198 (i8o)(x ; width across the genae 198-229 (214)^. Dorsal
arm of midcranial ridge anteriorly separated from other arms, and posteriorly fading out before
reaching the postoccipital ridge. Ventral and lateral arms forming a Y-shaped ridge. Post-
occipital ridge anteriorly confluent with preocular ridges. Dorsomedial part of epicranium
slightly raised. Preocular -\- interocular ridges join the postocular ridge below the ocellus ;
ventral rudiment of the preocular rige indicated by a weak sclerotization. Preoral ridge slender.
Eyes : Dorsal simple eyes projecting beyond the outer margins of the head in dorsal view ;
their corneae 21-27 ( 2 4)^ m diameter and separated by 101-116 (iio)|a, i.e. 3-9-5-3 (4-5) times
their diameter apart. Ventral eyes 24-31 (27)^ in diameter, and 21-34 ( 2 &)\>- apart. Lateral

140

MORPHOLOGY AND TAXONOMY OF ADULT MALES

ocelli large. Ocular sclerites well sclerotized. Cranial apophysis with truncate apex. Tentorial
bridge slender. Dorsal head setae on each side : 5-10 (7-3) f.s. and 6-12 (9-4) h.s. anterior to
postoccipital ridge ; dorsal ocular setae 0-2 (i-i) f.s. ; genal setae 7-15 (10-9) f.s. and 3-7 (4-0)

Fig. 25. Dysmicoccus alazon Williams, dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

Mi

h.s. Ventral head setae on each side : 3-5 (41) h.s. between the ventral eyes ; 10-16 (42-2)
f.s. and 1-3 (2-3) h.s. forming with their partners of the other side a transverse band in the area
of the ventral preocular depression ; ventral ocular setae 3-6 (44) f.s. and 0-2 (09) h.s. ; a
row of 0-1 (01) f.s. and 2-4 (28) h.s. present on each side of the ventral arm of the midcranial
ridge. Head disc pores : 1-2 (i-i) dorsal head pores occur near the base of each antenna ;
ventral pores absent.

Antennae : Filiform ; normally io-segmented ; 586-699 (647)^ long, i.e. about as long as
half the body length (ratio 1 : 20-22, av. 21), and shorter than the hind legs (ratio 1 : i-i-
13, av. 1-2). Scape 46-50 (48) \jl long and 43-46 (44) [i wide at base ; with 5-8 (59) h.s.
Pedicel 61-70 (67)0. long and 34-37 (35)(i wide ; with 14-20 (173) f.s., 10-17 (*3'3) h.s. and a
sensillum placodeum. Flagellum : Segment III club-shaped and longest of all, the ratio
lengths of third to terminal segments being 1-3-1-5 (14 : 1 ; segment III about 24(1 wide,
the ratio its width to its length 1 : 3-5-5-0 (46). Segments IV to X cylindrical and about
22 (x wide, the ratio width to length of segment IX being 1 : 2-5-2-9 (27). The lengths of
the flagellar segments and the number of setae on each are given in the following table :

Ill

IV

V

VI

VII

VIII

IX

X

length in ^

85-107

55-76

58-70

52-67

52-64

52-64

55-64

64-76

(av.)

(98)

(67)

(64)

(61)

(58)

(58)

(58)

(70)

f.s.

10-17

12-19

13-16

1 1-16

11-14

13-16

9-13

8-12

(av.)

(13-5)

(14-8)

(14-0)

(138)

(12-7)

(14-5)

(112)

(io-i)

h.s.

2-6

1-4

1-4

2-6

2-6

2-5

2-6

2-6

(av.)

(4-3)

(3o)

(3-2)

(4-3)

(37)

(37)

(4-o)

(3-2)

Antennal bristles easily distinguishable ; segments VIII and IX each with a ventral bristle.
Terminal segment with 3 preapical bristles, a pair of capitate subapical sensory setae and one
apical hair-like seta.

Thorax : 532-616 (570)^ long. Prothorax : Pronotal ridge medially interrupted by a weak
sclerotization. Lateral pronotal sclerites and post-tergites distinct. Proepisternum with a
ridge-like dorsal margin. Prosternum triangular, 31-40 (37) [i long ; prosternal ridge 85-95
(92) (x long. Prothoracic setae on each side : Medial pronotal setae 1-5 (19) f.s. and 1-3 (i-6)
h.s. ; lateral pronotal setae 1-2 (13) h.s. ; post-tergital setae 3-9 (59) f.s. and 1-2 (h.s. ; ante-
spiracular dorsal setae 1-2 (1-2) h.s. One, hair-like antespiracular ventral seta always present.
Prosternal setae 1-4 (i-8) f.s. and 1-3 (i-6) h.s. Prothoracic disc pores on each side : Medial
pronotal pores 2-4 (2-7) ; lateral pronotal pores 4-6 (5-1) ; antespiracular dorsal pores 1-3 (1-9).
Prosternal pores 1-3 (2-0).

Mesothorax : Prescutum 79-98 (85)0. long and 116-143 ( 131) [j. wide, the ratio being 1 : 14-
i-6 (1-5) ; laterally bounded by the prescutal ridge and posteriorly by the prescutal suture.

o> «> °

200

Fig. 26. Dysmicoccus alazon Williams, lateral view.

142

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Scutum heavily sclerotized antero-laterally, with a median longitudinal narrow membranous
area ; scutum 107-128 (n6)|i long, i.e. the ratio lengths of prescutum to scutum 1 : 1-3-1-5
(1-4). Prealare, prealar ridge and triangular plate well developed. Scutellum 52-70 (61) a.
long and 92-110 (ioi)jj. wide (ratio 1 : 1-6-1-7, av - l '^5> an< 3 the ratio its length to the length
of scutum 1 : 1-8-2-1, av. 1-9). A nterior and posterior postalar ridges separated. Mesopleuron :
Mesopleural ridge interrupted above the coxal articulation ; mesopleural apophysis, meso-
pleural wing process, subepisternal ridge well developed ; basalare stout. Mesepisternum and
mesepimeron distinct ; lateropleurite narrow. Mesosternum : Basisternum 143-186 (156) \i
long and 177-192 (186) \l wide ; bounded by the marginal and the precoxal ridges. Meso-
thoracic spiracle 18-21 (20) jz wide at opening, with a 37-46 (40) u. long supporting bar. Meso-
thoracic setae on each side : Prescutal setae 3-5 (4-1) h.s. ; scutal setae 4-6 (4-8) h.s. ; scutellar
setae 4-7 (5-1) h.s. Tegular setae 1-2 (1-4) h.s. Postmesostigmatal setae in two groups, 3-7
(4-6) f.s. and 1-4 (21) h.s. behind each spiracle, and 3-9 (6-i) f.s. and 1-3 (19) h.s. separated
medially. Basisternal setae 9-16 (11-9) h.s. Mesothoracic disc pores : Mesospiracular pores
0-1 (0-2).

Metathorax : Metapostnotal sclerites, metapostnotal ridge and metapleural structures typical
of the family. Metathoracic spiracles identical with the mesothoracic. Metathoracic setae on
each side : Metatergal setae 1-5 (2-7) f.s. and 2-6 (4-7) h.s. ; metapleural setae 2-8 (4-8) f.s. and
1-3 (1-4) h.s. Anterior metasternal setae 3-7 (5-3) f.s. and 2-6 (3-4) h.s. ; posterior metasternal
setae (pmss) 0-2 (1-3) f.s. and 1-4 (i-8) h.s. Metathoracic disc pores : Metaspiracular pores 0-2
(0-9) ; anterior metasternal pores 0-1 (0-4).

Wings : 1190-1400 (i274)jx long and 406-532 (448)0. wide ; with 1-4 (2-2) alar setae and
2 minute circular sensoria. Hamulohalterae 73-82 (76) jj. long and 15-18 (16) ji wide ; with a
49-61 (55) [x long apically hooked seta, i e. the ratio length of seta to the length of hamulohaltera
1 : 1-3-1-5 (l'4)-

Legs : Moderately long and slender ; the ratio length of the hind leg to the total length
of the body 1 : 1-7-2-0 (i-8). Coxa and trochanter about 52 and 241X wide respectively. Femur
about 37 (i. wide ; that of fore leg shortest and that of hind leg longest ; the ratio width to
length of the hind femur 1 : 5-8-6-4 (6-i). Tibia about 22u. wide, with 2 apical spurs and
3-5 smaller spines ; the ratio lengths of femur to tibia of fore leg 1 : 1-0-1-2 (i-i). Tarsus
about 2 1 (A wide ; with a pair of apically knobbed tarsal digitules. Claw uniformly tapering
to a pointed end ; with a pair of fine ungual digitules. The following table shows the lengths
of leg segments (in microns) and the number of setae on each :

Fore leg

Middle leg

Hind leg

length

40-46 (44)

40-46 (44)

43-49 (45)

Coxa

f.s.

9-12 (10-4)

9-13 (n-i)

10-14 ( II- 2)

h.s.

7-10 (8-8)

8-14 (10-2)

10-15 (12-4)

length

61-64 (63)

61-64 (63)

64-70 (68)

Trochanter

f.s.

i-5 (2-8)

2-5 (3-2)

3-5 (3-9)

h.s.

2-4 (3-i)

2-4 (3-6)

2-5 (4-i)

length

186-211 (198)

189-217 (204)

214-235 (226)

Femur

f.s.

13-19 (15-6)

19-26 (21-7)

22-31 (25-4)

h.s.

14-21 (17-3)

13-25 (20-6)

18-31 (26-1)

length

198-238 (214)

229-259 (241)

284-320 (299)

Tibia

f.s.

20-31 (24-0)

23-36 (29-4)

32-43 (377)

h.s.

22-35 (26-2)

22-39 (32-2)

28-49 (40-3)

length

85-92 (88)

89-95 (92)

98-107 (104)

Tarsus

f.s.

8-12 (10-4)

8-12 (io-6)

9-14 ("7)

h.s.

12-17 (147)

14-19 (15-5)

14-20 (17-4)

Claw

length

27-34 (3o)

27-34 (30)

27-34 (30)

Total length of leg

616-680 (637)

662-711 (683)

741-808 (769)

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 143

Abdomen : 464-585 (502)0. long and 274-304 (289) y. wide. Tergites of segments I and II
small ; those of segments VIII and IX + X large and well sclerotized. Sternites of segment
VIII ill-denned. Ostioles well developed, 40-46 (43) n long at orifice. Abdominal setae on each
side : Dorsal setae : Fleshy setae 1-3 on segment I, usually 2-5 on segments II to VII (some-
times i on segment II and 6 on segments V and VII), and 1-3 on se ment VIII ; hair-like
setae 0-2 on segment I, 2-4 on segments II to VII, and 0-3 on segmeni VIII. Pleural setae :
Fleshy setae 0-4 on segments I to VIII ; hair-like setae 2-4 on segment 1, 3-5 on segments
II to VII, and 2-5 on segment VIII, including one slightly longer seta. Ventral setae : Fleshy
setae 1-2 on segment II, and usually 2-4 on segments III to VIII (sometimes 5 or 6 on segments
VI and VII, and only 1 on segment VIII) ; hair-like setae usually 2-4 on segments II to VII
(sometimes 3 or 5 on segment III), and 0-1 on segment VIII. Abdominal pores : Segments I
to VII with 6-10 (7-5), 0-2 (03), 0-2 (05), 0-2 (0-5), 0-1 (08), 1-3 (15), and 2-4 (28) pleural
pores. One dorsal pore may also occur on segments I and II, and one ventral pore on segments
II and III.

Setae of glandular pouch consist of a pair of 397-488 (424)^ long tail setae and one seta about
40^1 long, i.e. the ratio length of the tail setae to the total length of the body 1 : 27-37 (3-2).

Genital segment comparatively small ; subtriangular in dorsal view ; style slightly curving
upwards in lateral view. Penial sheath 140-153 (150)1* long and 82-795 (89)1* wide, i.e. the
ratio length to width 1-5-1 -9 (17) : 1, and the ratio its length to the total length of the body
1 : 87-10-3 (9-1). Basal ridge of penial sheath and its projection well developed ; process of
penial sheath absent. Aedeagus relatively wide at its basal rod, becoming narrower towards
its pointed apex. Setae of genital segment on each side : 3 setae always occur dorsally near
the base of the style ; 3-4 (3-8) smaller setae and 3-4 (32) setal sensilla present ventrally on
the penial sheath.

Material : 8 specimens examined, collected by myself, 20. i v. 1964, on a bunch of
bananas bought in a London fruit shop, imported from the Canary Islands ; the
females were identified by K. Boratynski and confirmed by Dr. D. J. Williams.

SACCHARICOCCUS Group

SACCHARICOCCUS Ferris, 1950

Saccharicoccus sacchari (Cockerell)

(Text-figs. 27-30)

Both macropterous and apterous forms of this species were available.

(A) The macropterous form (Text-figs. 27 and 28).

Living specimens light brown. A narrow and slender species, comparatively large, with
short antennae and short legs. When mounted, total length of body 1260-1400 (1316)^1,
width at mesothorax 252-280 (226) (i and wing expanse 1764-2072 (1932) ji.

Body setae and pores : Antennae and legs with numerous fleshy, 2 1-27 \i long setae, and a
few hair-like, slightly longer ones. The body itself with fine hair-like setae only. Quadri-
locular and occasionally quinquelocular disc pores always present on head, thorax and abdomen,
about 6 [A in diameter.

Head : Subtetrahedron ; subtriangular in dorsal view ; dorso-ventrally flattened in front
and lateral views ; ventral preocular depression entirely absent. Length from apex to post-
occipital ridge 122-137 ( I 3 I )i A '< from apex to neck 159-180 (171)^ ; width across the genae
180-198 (192) (i. Dorsal arm of midcranial ridge marked only by slender sclerotization, some-

i 4 4

MORPHOLOGY AND TAXONOMY OF ADULT MALES

dhp

vhs
/ scp

**™$p&

Fig. 27. Saccharicoccus sacchari (Cockerell), macropterous form, dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

145

times short or reaching the level of the dorsal eyes. Lateral arms forming together with the
ventral arm a Y-shaped ridge. Postoccipital ridge V-shaped and medially interrupted at a
weak point ; the ridge not reaching the preocular ridges anteriorly. Dorsomedial part of
epicranium slightly raised. Preocular and interocular ridges well developed ; ventral rudiment
of the former absent. The dorsal part of the postocular ridge reduced to an atrophied arm.
Preoral ridge slender. Eyes : Dorsal simple eyes not projecting beyond the outer margins of
the head in dorsal view ; their corneae 15-21 (i8)(j. in diameter and separated by 104-116
(iio)fi, i.e. 5-0-7-6 (60) times their diameter apart. Ventral simple eyes 18-21 (20)^ in
diameter and 21-31 (24)^ apart. Lateral ocelli well developed. Cranial apophysis apically
truncate. Tentorial bridge slender. Dorsal head setae 10-14 ( IX °6) on each side in front of
the postoccipital ridge ; each gena always with 3 genal setae. Ventral head setae on each side :
2-3 (2-4) between the ventral eyes ; 8-1 1 (9-3) just anterior to the ventral eyes, forming on both
sides a transverse band. Head pores : 4-7 (5-1) dorsal head pores present near the base of
each antenna ; ventral pores absent.

Antennae : Filiform ; normally 10-segmented but fusion between two adjacent segments
of the flagellum seems to be common ; 412-482 (455) (x long, i.e. shorter than half the length
of the body, the ratio being 1 : 2-7-3-1 (2-9), and shorter than the hind leg, the ratio 1 : 1-1-1-3
(1-2). Scape 34-37 (36)11 long and 40-43 (4i)(x wide at base ; with 5-7 (6-2) h.s. Pedicel
46-49 (48) (x long and 31-34 (33) (x wide ; with 8-16 (12-2) f.s., 6-10 (8-4) h.s. and a sensillitm
placodeum. Flagellar segments : Segment III club-shaped, subequal in length to segment X,
the ratio their lengths being o-8-i-i (09) : 1. The flagellar segments about 2ijx wide, i.e. the
ratio width to length of segments III and IX 1 : 2-3-2-9 (2-6) and 1 : 1-9-2-3 (2-1), respectively.
The following table shows the lengths of the flagellar segments and the number of setae on
each :

III

IV

V

VI

VII

VIII

IX

X

length in |x

49-61

31-40

34-40

37-46

43-49

43-55

43-49

55-6i

(av.)

(55)

(36)

(38)

(43)

(47)

(49)

(48)

(58)

f.s.

5-9

5-10

5-10

6-13

9-17

8-16

8-13

10-15

(av.)

(7-2)

(7-4)

(7-6)

(io- 4 )

(13-4)

(13-2)

(n-6)

(12-6)

h.s.

i-3

1-2

1-2

2-3

3-5

2-5

2-4

4-6

(av.)

(i-8)

(1-6)

(1-2)

(2-2)

(4-o)

(3-8)

(3-4)

(4-8)

200-^

Fig. 28. Saccharicoccus sacchari (Cockerell), macropterous form, lateral view.

146 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Antennal bristles distinct ; segments VIII and IX each with one ventral bristle. Terminal
segment with 3 preapical bristles, 2 capitate subapical sensory setae and 1 apical hair-like seta.

Thorax : 517-570 (540)^ long. Prothorax : Pronotal ridges medially interrupted as usual.
Lateral pronotal sclerites small, and post-tergites comparatively large. Proepisternum with a
ridge-like dorsal margin. Prosternum subtriangular, 49-58 (52)0 long and 89—98 (92) (x wide ;
prosternal ridge absent, and its position only indicated by a narrow sclerotization. Prothoracic
setae on each side : Medial pronotal setae 3-5 (3-9) ; lateral pronotal setae 3-5 (3-4) ; post-
tergital setae 2-4 (3-0) ; antespiracular dorsal setae usually 5-7, and occasionally 9 (average 6-i).
Antespiracular ventral setae usually 2, and occasionally 3 (av. 2-1) ; prosternal setae 2-4 (2-9).
Prothoracic disc pores on each side : Medial pronotal pores 0-1 (0-3) ; lateral pronotal pores
2-4 (2-6) ; post-tergital pores absent ; antespiracular dorsal pores 2-3 (2-1). One prosternal
pore may occur on either side (av. 0-4).

Mesothorax : Prescutum 85-95 (92) u. long (it is difficult to determine the actual length of the
scutum due to the absence of the prescutal suture), and 125-140 (134)0 wide, the ratio length
to width being 1 : 1-4-1-6 (1-5) ; prescutal ridge well developed and prescutal suture absent.
Scutum heavily sclerotized anterolaterally, with a median membranous longitudinal narrow
area ; 92-116 (no)a long, i.e. the ratio lengths of prescutum to scutum 1 : 1-1-1-3 (1-2).
Prealare, prealar ridge, triangular plate, anterior and posterior notal wing processes well developed.
Scutellum 46-52 (49) (i long and 92-104 (95)0. wide, i.e. the ratio its length to its width 1 : 1-9-
2-1 (1-94), and the ratio its length to the length of scutum 1 : 2-0-2-4 (2-3). Postalare with
well separated anterior and posterior ridges. Postnotal apophysis comparatively small. Meso-
pleuron : Mesopleural ridge interrupted above the coxal articulation ; basalare stout. Sub-
episternal ridge slender. Mesepisternum and mesepimeron distinct ; lateropleurite moderately
large. Mesosternum : Basisternum 134-159 (153) u. long and 137-159 (146)0 wide ; the
antero-median part of the marginal ridge absent ; precoxal ridge well developed. Furca
comparatively small. Mesothoracic spiracle 18-21 (20) o wide at opening, with a 34-40 (37) o
long supporting bar. Mesothoracic setae on each side : Prescutal setae 3-5 (3-6) ; scutal setae
5-7 (5-3) ; scutellar setae 1-3 (2-1). Tegular setae 2-4 (2-6). Postmesostigmatal setae 6-10 (7-7)
in a latero-ventral group. Basisternal setae 5-7 (6-i). Mesothoracic pores : 1-2 .(1-4) meso-
spiracular pores present behind each spiracle ; postmesostigmatal pores absent.

Metathorax : Metapostnotal sclerites and metapostnotal ridge well developed. Metapleural
ridge with usual attenuation near the middle at the point of origin of the metapleural apophysis.
Precoxal ridge slender and metasternal apophyses small. Metathoracic spiracle identical with
the mesothoracic one. Metathoracic setae on one side : Metatergal setae 4-7 (5-1) ; metapleural
setae 1-3 (2-4). Anterior metasternal setae 3-6 (4-3) and posterior metasternal setae 2-3 (2-2).
Metathoracic pores on each side : Metaspiracular pores 1-3 (1-9) ; anterior metasternal pores
usually 1-2 and occasionally 3 or 4 (av. 1-9) ; posterior metasternal pores absent.

Wings : 770-910 (854)0. long and 280-350 (322) o wide ; with 2-4 (3-1) alar setae and 2
minute circular sensoria. Hamulohalterae 64-70 (69)0 long and 12-15 ( I 4)l jl wide ; with one
apically hooked, 40-46 (43)0 long seta, i.e. the ratio lengths of seta to hamulohaltera 1 : 1-4-
1-7 (i-6).

Legs : Short and slender ; the ratio length of hind leg to the total body length 1 : 2-3-
2*4 (2-38). Coxa and trochanter about 49 and 240 wide respectively. Femur about 40a wide ;
that of the hind leg longest and those of the front and middle legs of about the same length ;
the ratio width to length of the hind femur 1 : 3-8-4-2 (4-1). Tibia about 190 wide ; with
two apical strong spurs and 2-4 smaller spines ; the ratio lengths of femur to tibia in front
leg i-i-i (1-06) : 1. Tarsus about 180 wide ; tarsal digitules apically knobbed. Claw
uniformly tapering to a sharply pointed tip ; ungual digitules extremely fine. The lengths of
the leg segments (in microns) and the number of setae on each are given below :

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

M7

Coxa

Trochanter
Femur
Tibia

Tarsus

Claw

Total length of leg

length

f.s.

h.s.

length

f.s.

h.s.

length

f.s.

h.s.

length

f.s.

h.s.

length

f.s.

h.s.

length

Middle leg

40-43 (42)

i-3 (i-8)

5-8 (7-2)

52-58 (55)
1-2 (1-4)

3-5 (4-2)

143-159 (153)

10-16 (n-6)

5-9 (7-2)

146-171 (159)

12-18 (15-2)

4-9 (6-2)

67-70 (68)

2-4 (3-2)

6-9 (7-4)

24-30 (27) .

476-525 (500)

Hind leg

43-46 (44)

1-2 (i-6)

6-8 (6-4)

55-6i (58)

i-3 (2-2)

4-5 (4-4)

153-168 (162)

9-14 (120)

6-8 (7-4)

177-192 (186)

12-19 (15-6)

5-8 (6-4)

73-76 (74)

3-2 (4-4)

7-9 (7-8)

24-30 (27)

528-573 (552)

Fore leg

40-43 (42)

1-2 (i-6)

5-7 (6-2)

52-58 (55)
1-2 (1-4)
3-6 (4-6)

M3-I59 (153)

10-17 (12-4)

6-9 (7-4)

134-159 (143)

14-17 (15-4)

4-7 (5-8)

67-70 (68)

3-6 (46)

7-9 (8-2)

24-30 (27)

464-512 (485)

Abdomen : 448-562 (502)0. long and 243-274 (258)0. wide. Tergites of segments 1 and II
small ; those of segments VIII and IX + X large and distinct. Sternites of segment VIII
weakly sclerotized. Ostioles well developed, 40-46 (43)11 long at orifice. Abdominal setae
on each side : Dorsal setae 3-5 on segments I to VI, and 2-4 on segments VII and VIII.
Pleural setae 4-6 on segment I, 5-7 on segments II to VII, and 3-4 on segment VIII, including
a slightly longer seta. Ventral setae usually 2-3 on segments II to VII (sometimes 4 on
segments III and IV), and absent on segment VIII. Abdominal pores : Segments I to VII
with 3-5 (3-9), 1-3 (1-4), 1-3 (1-7), 1-3 (17), 1-3 (17), 1-2 (1-3), and 1 pleural pore, respectively.
Segments II to VII also with 0-1 (01), 0-2 (i-i), 0-2 (11), 0-2 (10), 0-1 (07), and 0-2 (09)
ventral pores, respectively.

Glandular pouch small ; setae of glandular pouch include a pair of 305-336 (320)0. long tail
setae and one seta, 40-55 (46)51 long, i.e. the ratio length of the tail setae to the total length of
the body 1 : 4-0-4-2 (41).

Genital segment subtriangular in dorsal view. Style apically rounded, rather straight in
lateral view. Penial sheath 131-150 (137)0. long and 70-79 (73)11 wide, the ratio being i-8-
1-9 (1-87) : 1, and the ratio its length to the total body length 1 : 9-3-10-1 (96). Basal ridge
of penial sheath with a small projection. Process of penial sheath well pronounced. Aedeagus
comparatively slender, with a large internal genital aperture. Setae of genital segment :
Dorsally, 3 setae always present on each side near the base of the style ; ventrally, 5-6 (5-2)
smaller setae occur on each side of the penial sheath ; process of the penial sheath with 2-3
setal sensilla.

Material : 7 specimens examined, collected by Mr. A. I. Ezz and myself, on
sugar cane, in Giza, Cairo, U.A.R., during the second and third weeks of August,
1964.

(B) The apterous form (Text-figs. 29 and 30).

Living specimens not available. The males are moderately robust, of medium size, with
very short antennae and comparatively short legs. Mounted specimens 1120-1190 (ii48)(i
long and 308-350 (322) u. wide.

Body setae and pores : Antennae and legs with comparatively few, about 15(1 long fleshy
setae, and about 27^ long hair-like ones. The body itself only with hair-like setae, 31-49(1
long ; each of abdominal segments VI, VII and VIII also with a rather long and stout seta on

MORPHOLOGY AND TAXONOMY OF ADULT MALES

, dhp vhs scp
spl r n I

setscla h

Fig. 29. Saccharicoccus sacchari (Cockerell), apterous form, dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

149

each side. Numerous quadrilocular, and occasionally trilocular pores, about 6u. in diameter,
present on the head, thorax and abdomen.

Head with a narrow truncate apex, becoming broader posteriorly as it fuses with the thorax.
Length from apex to postoccipital ridge 85-101 (95) n and width across the genae 174-189
(180) [jl. Midcranial ridge with its dorsal, lateral and ventral arms entirely absent. Postoccipital
ridge V-shaped. Dorsomedial part of epicranium well sclerotized. Ocular ridges usually
present, but sometimes atrophied or absent altogether. If present, the preocular and inter-
ocular ridges join the postocular ridge just below the ocellus. Preoral ridge weak. Eyes :
Dorsal and ventral simple eyes absent ; lateral ocelli small. Cranial apophysis comparatively
small, apically truncate. Tentorial bridge slender. Dorsal head setae 9-12 (105) on each side
of the median line ; each gena always with 3 genal setae. Ventral head setae on each side :
7-9 (8-3) just anterior to the level of ocelli ; posteriorly, 0-1 (o-6) occur medially. Head pores :
Dorsal head pores 2-4 (2-8) near the base of each antenna ; ventral pores absent.

Antennae : Filiform ; 8-segmented ; 256-293 (275) ji. long, i.e. the ratio its length to the
total body length 1 : 3-9-4-6 (4-2), and the ratio its length to the length of the hind leg 1 : 1-7-
2-0 (i-8). Scape 40-46 (43) u. long and 37-43 (40) (a wide at base ; with 4-5 (4-2) h.s. Pedicel
37-40 (39)u. long and 31-34 (33)(x wide ; with 4-5 (4-5) f.s., 4-8 (5-8) h.s. and a sensillum
placodeum. Flagellar segments : Segment III club-shaped and subequal in length to segment
VIII (ratio 0-9-1-2, av. i-i : 1). Flagellar segments about 24^ wide, the ratio width to length
of segments III and VII being 1 : 1-4-1-8 (i-8) and 1 : 1-2-1-3 ( I-2 5)> respectively. The
lengths of the flagellar segments and the number of setae on each are given in the following
table :

Length in jx

(av.)

f.s.

(av.)

h.s.

(av.)

Ill
34-36
(40)
4-8
(5-8)
i-3
(2-3)

IV
24-40
(34)
i-5
(3-o)
3-5
(4-2)

V
21-27
(24)
i-3
(2-3)
4-5
(4'8)

VI

27-34
(3o)
i-4

(2-1)

3-5

(4-5)

VII

27-31
(29)
1-2

(i-5)
4-6

(4-8)

Antennal bristles much stouter and longer than the antennal fleshy setae

VIII

27-40
(39)
i-3
(2-3)
6-8
(6-6)

segments VI and

o.

Q.

/
/

l/l

/
/

CL

1
1
l/l

c

E

1/1

1/1

Q.

I

Q.
1/1

\ \

- -ads

avp

200

Fig. 30. Saccharicoccus sacchari (Cockerell), apterous form, lateral view.

15°

MORPHOLOGY AND TAXONOMY OF ADULT MALES

VII each with a ventral bristle. Segment VIII with 3 preapical bristles, 2 capitate subapical
sensory setae and 1 apical hair-like seta.

Thorax : Length from postoccipital ridge to first abdominal segment 418-441 (433)^.
Prothorax : Pronotal ridges usually weak or atrophied but sometimes well developed ; pronotal
sclerites small. Post-tergites comparatively large. Dorsal margin of the proepisternum ridge-
like. Prosternum small, triangular, 9-15 (12) jjl long and 61-70 (65) u. wide ; prosiernal ridge
absent, and its position only marked by sclerotization. Prothoracic setae on each side : Medial
pronotal setae 3-4 (3-3) ; lateral pronotal setae 7-9 (8-3) ; post-tergital setae 3-4 (3-8) ; ante-
spiracular dorsal setae 5-7 (5-8). Antespiracular ventral setae 3-5 (3-4) ; prosiernal setae 2-4
(2-8). Prothoracic disc pores on one side : Medial pronotal pores 3-4 (3-8) ; lateral pronotal
pores 4-7 (5-3) ; post-tergital pores absent ; antespiracular disc pores 3-4 (3-3). Prosiernal
pores 2-3 (2-2).

Mesothorax : Largely reduced ; mesotergum represented by a weakly sclerotized sclerite.
Mesopleural ridge atrophied ; mesepisternum distinct and mesepimeron small. Basisternum
weakly sclerotized and hard to detect ; marginal and precoxal ridges absent ; furca vestigial.
Mesothoracic spiracle 18-21 (20) \i wide at opening, with 49-55 (52) pt long supporting bar.
Mesothoracic setae on each side : Dorsal setae 5-7 (6-2) forming on both sides a transverse band ;
pleural setae 4-5 (4-5) ; postmesostigmatal setae 9-13 (11-3) behind each spiracle. Basisternal
setae 3-5 (4-2). Mesothoracic pores : Mesospiracular pores 1-2 (1-5).

Metathorax : Metapleural ridge dorsally reduced ; precoxal ridge absent. Metasternal
apophyses vestigial. Melathoracic spiracle similar to the mesothoracic. Metathoracic setae
on each side : Dorsal setae 5-7 (5-8) ; metapleural setae 5-9 (6-8) ; anterior metasternal setae
2_ 4 (3 '3) '< posterior metasternal setae always 2. Metathoracic pores on each side : Dorsal
pores 1-2 (i-8) ; metaspiracular pores 5-8 (6-2) ; anterior metasternal pores 2-4 (3-1) and
posterior metasternal pores 1-2 (1-3).

Legs : Comparatively short ; the ratio length of hind leg to total body length 1 : 2-3-2-4
(2-32). Coxa about 46 and trochanter about 24^ wide. Femur about ^j\l wide ; that of the
hind leg longest and those of the front and middle legs of about the same length ; the ratio
width to length of the hind femur 1 : 3-8-4-0 (3-9). Tibia about 20\l wide ; with 2 apical
spurs (smaller spines absent) ; the femur of fore leg always longer than the tibia, the ratio
being 1-2-1-3 ( I-21 ) : *■ Tarsus about i8[x wide ; tarsal digitules apically knobbed. Claw
gradually tapering to a pointed end ; ungual digitules fine. The following table shows the
lengths of the leg segments (in microns) and the number of setae on each :

Fore leg

Middle leg

Hind leg

length

37-43 (40)

37-43 (40)

40-46 (43)

Coxa

f.s.

i-3 (i-8)

i-3 (2-1)

1-2 (1-5)

h.s.

4-7 (5-6)

5-8 (6-8)

5-8 (7-i)

length

58-61 (60)

58-61 (60)

61-64 (63)

Trochanter

f.s.

0-1 (o-8)

0-1 (0-9)

1-2 (i-3)

h.s.

3-5 (4-i)

3-5 (3-8)

4-6 (4-8)

length

134-140 (137)

134-140 (137)

140-146 (143)

Femur

f.s.

2-3 (2-8)

i-3 (2-6)

2-4 (3-i)

h.s.

5-8 (6-5)

5-8 (6-9)

6-9 (7-6)

length

110-116 (113)

116-128 (122)

137-146 (140)

Tibia

f.s.

2-4 (3-2)

2-4 (3-4)

2-4 (3-3)

h.s.

6-8 (6-9)

6-9 (7-5)

7-9 (8-2)

length

64-70 (67)

64-70 (67)

70-76 (73)

Tarsus

f.s.

i-3 (2-3)

i-3 (2-6)

2-3 (2-4)

h.s.

4-6 (5-3)

4-7 (6-i)

5-7 (6-4)

Claw

length

27-31 (29)

27-31 (29)

27-31 (29)

Total length

of leg

436-461 (445)

445-473 (455)

482-503 (494)

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 151

Abdomen : Total length from first abdominal segment to projection of basal ridge of penial
sheath 448-509 (479) [x, and width at third segment 289-342 (312)^1. Tergites and sternites
of segments I— VII occurring as narrow median sclerotized areas ; tergite of segment VIII
large and that of segments IX + X distinct ; sternites of segment VIII ill-defined. Ostioles
well developed, 34-40 (37) (i. long at orifice. Abdominal setae on each side : Dorsal setae 3-5
on segments I to VII and 2-4 on segment VIII. Pleural setae on segments I to VII and 3-4
on segment VII I, including one considerably long and stout seta on each of segments VI, VII
and VIII. Ventral setae 1-3 on segment II, 2-4 on segments III to VII, and absent on segment
VIII. Abdominal pores : Segments I to VII with 3-5 (4-2), 2-5 (2-9), 2-5 (2-8), 2-4 (2-7),
1-3 (2-1), 1-2 (i-8), and 0-1 (05) pleural pores, respectively. The segments also with 0-2
(o-8), 0-2 (0-4), 0-2 (0-5), 0-2 (0-5), 0-2 (1-2), 0-2 (0-9), 0-1 (0-3), and 0-1 (0-3) dorsal pores.
One or two ventral pores present at least on one side of segments II to VI and sometimes also
VII (averages 15, 13, 11, 13, 06, and 03, respectively).

Glandular pouch small ; setae 0/ glandular pouch include a pair of 153-229 (180) ji long tail
setae, and one seta, 61-76 (70) |j. long, i.e. the ratio length of the tail setae to the total length
of the body 1 : 5-7-8 (6-4). Genital segment identical with that of the macropterous males.
Penial sheath 128-143 (140)^ long and 79-89 (&2)[i wide, ratio 1-6-1-7 (168) : 1, and the ratio
its length to the total body length 1 : 7-8-8-9 (8-2).

Material : 12 specimens examined, collected by B. S. Chandel, on sugar cane,
in U.P. Bichpuri farm, Balawant Rajput College, Agra, United Provinces, India,
during the summer of 1964 (received from Dr. D. J. Williams).

OCTOCOCCUS Group

OCTOCOCCUS Hall, 1939

Octococcus africanus (Brain)

(Text-figs. 31, 32)

Only winged forms known ; moderately robust, small, with very long antennae and long
legs. Mounted specimens 882-1064 (966)1* long, 308-336 (322)0. wide and 2212-2534 (2366) jx
wing expanse.

Body setae and pores : Antennae with numerous fleshy setae and a few hair-like ones, about
24(x long. The body itself and the legs with hair-like setae only. Quadrilocular and
occasionally quinquelocular disc pores present only on the thorax and the first abdominal
segment, about 6u in diameter.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression well pronounced in lateral view. Length from apex to postoccipital ridge 89-110
(101)0 ; from apex to neck 137-153 (146)0 ; width across the genae 165-192 (177)0. Dorsal
arm of midcranial ridge anteriorly separated from other arms and posteriorly fading out before
reaching the postoccipital ridge. Lateral and ventral arms forming together a Y-shaped ridge.
Postoccipital ridge not reaching the preocular ridges. Preocular and interocular ridges joined ;
the former with a hardly indicated ventral rudiment below the articular process. Postocular
ridge well developed and preoral ridge slender. Eyes : Dorsal simple eyes slightly projecting
beyond the outer margins of the head in dorsal view ; their corneae 18-21 (20) o in diameter
and separated by 92-119 (104)0, i.e. 4-4-6-0 (5-1) times their diameter apart. Ventral simple
eyes 15-21 (18)0 in diameter and 24-31 (27)0 apart. Lateral ocelli large. Cranial apophysis
comparatively short and apically truncate. Tentorial bridge slender. Dorsal head setae 5-7
(5-9) on each side anterior to the postoccipital ridge ; each gena always with 3 genal setae.
Ventral head setae on each side : Setae between the ventral eyes absent ; 3-4 (3-6) setae in the

152 MORPHOLOGY AND TAXONOMY OF ADULT MALES

area of the ventral preocular depression, forming on both sides a transverse irregular row ;
2-3 (2-2) setae on each side of the ventral arm of the midcranial ridge. Head disc pores entirely
absent.

Antennae : Filiform ; normally 10-segmented ; 668-769 (729)^ long, i.e. much longer
than half the total body length (ratio 1 : 1-2-1-6, av. 1-3), and longer than the hind legs (ratio
1 • 1-1-3, av. 1-2 : 1). Scape 43-49 (46) \i long and 46-49 (47) jx wide at base ; almost always
with 4, and occasionally 5 h.s. (av. 4-1). Pedicel 58-64 (6i)|a long and 37-40 (38)11 wide ;
with 10-18 (13-1) f.s., 3-8 (5-3) h.s. and a sensillum placodeum. Flagellar segments with a
few apically knobbed, about 400. long capitate sensory setae. Segment III club-shaped, longest
of all and about 27^ wide, the ratio its length to the length of segment X 1-2-1-4 ( I- 3) : *• an( i
the ratio its width to its length 1 : 3-1-3-7 (3-3). Segments IV to X slightly narrower, the
ratio width to length of segment IX being 1 : 2-7-3-3 (3-0). The lengths of the flagellar
segments and the number of setae on each are given in the following table :

III IV V VI VII VIII IX X

length in (x

85-101

67-82

70-82

70-85

67-82

70-85

64-79

70-73

(av.)

(92)

(76)

(79)

(82)

(76)

(79)

(73)

(7i)

f.s.

12-19

10-20

10-19

13-20

13-20

12-21

14-20

13-16

(av.)

(15-6)

(14-6)

(I4-I)

(16-3)

(17-8)

(18-4)

(i7-9)

(147)

h.s.

2-4

(av.)

(3-4)

set. ca.

1-2

2-3

1-2

3-4

4-5

5-6

4-6

4

(av.)

(i-6)

(2-7)

(i-3)

(37)

(4-4)

(5-3)

(5-i)

(40)

Antennal bristles inconsiderably stouter than the fleshy setae ; segments VIII and IX each
with a ventral bristle. Terminal segment with 3 preapical bristles and 1-2 apical sensilla
basiconica.

Thorax : 418-479 (448) u. long. Prothorax : Pronotal ridges medially interrupted by a weak
sclerotization. Lateral pronotal sclerites and post-lergites distinct. Dorsal margin of pro-
episternum ridge-like. Prosternum triangular, 12-18 (15) [a long ; posteriorly bounded by the
transverse, 82-98 (92) [i long prosternal ridge. Prothoracic setae on each side : Medial pronotal
setae absent ; lateral pronotal seta 0-1 (0-3) ; post-tergital and antespiracular dorsal setae absent.
One antespiracular ventral seta always present ; prosternal setae almost always absent although
one seta may occur on one side (av. 0-2). Prothoracic disc pores on each side : Medial and
lateral pronotal pores 0-1 (o-8) and 0-2 (0-9) respectively ; antespiracular dorsal pores absent.
Prosternal pores 1-2 (1-3).

Mesothorax : Prescutum 67-79 {73)11 long and 116-137 (i28)u. wide, the ratio being 1 : i-6-
1-9 (i-8) ; laterally and posteriorly bounded by the prescutal ridge and the prescutal suture.
Scutum well sclerotized antero-laterally and with a median longitudinal membranous narrow
area ; 73-89 (76)(x long, i.e. the ratio lengths of prescutum to scutum 1 : 1-0-1-2 (i-i).
Prealare, prealar ridge, triangular plate, anterior and posterior notal wing processes well developed.
Scutellum 49-61 (55)0. long and 82-104 (92)^1 wide, i.e. the ratio 1 : 1 -6-1-8 (1-7), and the ratio
its length to the length of the scutum 1 : 1-3-1-5 (1-4). Postalare with well separated anterior
and posterior postalar ridges. Postnotal apophysis strong. Mesopleuron : Mesopleural ridge
interrupted above the coxal articulation ; basalare comparatively slender. Mesepisternum
and mesepimeron distinct ; later opleurite narrow. Mesosternum : Basisternum 1 13-134 (i22)[A
long and 189-232 (204) }x wide ; bounded by the marginal and the precoxal ridges ; furca well
developed. Mesothoracic spiracle 21-24 (23) [z wide at opening, with a 40-46 (43) (x long
supporting bar. Mesothoracic setae on each side : Prescutal setae 2-4 (3-1) ; scutal setae 4-6
(4-9) ; scutellar setae 2-3 (2-4). One tegular seta always present. Postmesostigmatal setae
2—4 (3-1) behind the spiracles, and one medially separated. Basisternal setae 5-7 (5-3). Meso-
thoracic disc pores : Mesospiracular pores 1-2 (i-i) ; postmesostigmatal pores absent.

Metathorax : Metapostnotal sclerites well defined and metapostnotal ridge weakly developed.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

153

Metapleural ridge attenuated at the point of origin of the metapleural apophysis. Precoxal
ridge well developed and metasternal apophysis distinct. Metathoracic spiracle similar to the
mesothoracic. Metathoracic setae on each side : Metatergal setae always 3 ; metapleural setae
0-1(0-4). Anterior metasternal setae usually 1 and occasionally 2 (av. 1 -i) ; posterior metasternal
setae 0-1 (01). Metathoracic pores : One metaspiracular pore always present behind each
spiracle.

Wings : 966-1120 (1036)^ long and 392-462 (420) u. wide ; with 2 alar setae and 2 minute
circular sensoria. Hamulohalterae 79-85 (82) jjl long and 18-21 (ig)(x wide ; with a 52-61 (55)(i
long apically hooked seta, i.e. the ratio lengths of seta to hamulohaltera 1 : 1-4-1-6 (1-5).

dhs vhs

Fig. 31. Octococcus africanus (Brain), dorsal and ventral view.

J54

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Legs : Comparatively long and stout ; the ratio length of the hind leg to the total body
length i : 1-5-1-7 (i-6). Coxa about 55 and trochanter about 27 \i wide ; the latter with a
differentiated long apical seta. Femur about 4051 wide ; that of the fore leg longest and that
of the middle leg shortest ; the ratio width to length of the hind femur 1 : 3-7-4-2 (4-0). Tibia
about 24 (x wide ; with 2 apical strong spurs and 2-3 smaller spines ; in fore legs the femur
usually shorter and sometimes as long as the tibia, the ratio their lengths being 1 : i-i-i (1-06).
Tarsus about 21 \l wide ; tarsal digitules apically knobbed, about 37(1 long. Claw gradually
tapering to a pointed end ; ungual digitules fine. The following table shows the lengths of the
leg segments (in microns) and the number of setae on each :

Fore leg Middle leg Hind leg

43-49 (46) 43-49 (4 6 ) 46-52 (49)

8-12 (9-3) 8-13 (10-4) 9-12 (10-5)

58-67 (61) 58-67 (61) 61-70 (66)

4-6 (4-8) 4-7 (5-2) 5~7 (5'4)

162-174 ( l6 8) 143-159 (153) 150-165 (159)

19-25 (21-3) 14-20 (17-2) 15-21 (17-8)

162-189 (177) 177-211 (189) 198-232 (214)

20-27 (23-5) 21-29 (247) 23-30 (27-1)

76-82 (79) 76-82 (79) 79-85 (82)

13-18 (154) 15-18 (16-9) 16-20 (18-1)

00-00 (24) 00-00 (24) 00-00 (24)

522-580 (555) 525-586 (555) 561-619 (592)

Abdomen : 266-334 ( 2 96)[^ long and 289-327 (312) \l wide. Tergites of segments I and II
small ; those of segments VIII and IX + X large and distinct. Sternites of segment VIII
ill-defined. Ostioles entirely absent. Abdominal setae on each side : Dorsal setae 2-3 on
segment I, 3-4 on segments II to VI, 2-3 on segment VII, and 1-2 on segment VIII. Pleural
setae 2-3 on segment I, 3-4 on segments II to VII, and always 1 on segment VIII. Ventral
setae 1 on segment II, 1-2 on segment III, 2-3 on segments IV to VII, and absent on segment
VIII. Abdominal pores : Segment with 1-3 (2-6) pleural pores on each side ; other abdominal
pores entirely absent.

Coxa

length
h.s.

Trochanter

length
h.s.

Femur

length
h.s.

Tibia

length
h.s.

Tarsus

length
h.s.

Claw length
Total length of leg

200 ^

100^-

Fig. 32. Octococcus africanus (Brain), lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCI DA E 155

Glandular pouches well developed ; setae of glandular pouch consist of a pair of 229—275 (256) y.
long tail setae, and a pair of shorter setae of unequal lengths, 159-183 (171) and 70-116 (82) ^a
long respectively, and a short one, subequal in length to the abdominal setae ; the ratio length
of tail setae to total body length being 1 : 3-4-4-2 (3-8). Genital segment small, subtriangular
in dorsal view. Style rather straight in lateral view, with a rounded apex. Penial sheath
104-116 (no)(ji long and 76-92 (85)0. wide, the ratio being 1-1-1-5 ( I- 3) : l anc l the ratio its
length to total body length 1 : 8-1-10-0 (89). Basal ridge of the penial sheath and its projection
well developed. Process of penial sheath hardly indicated. Aedeagus broad gradually tapering
dorso-posteriorly. Setae of genital segment on each side : Dorsally 4 setae always present near
the base of the style ; ventrally 2-4 (26) setae occur on the penial sheath and 3-4 small setal
sensilla on its process.

Material : 10 specimens examined, collected by O. W. Richards, on Nolletia sp.,
in Colesberg bridge (over Orange River), Cape Province, South Africa, on 26. ix. 1952.

CEROPUTO Group

CEROPUTO Sulc, 1898

Ceroputo pi lose I lav Sulc

(Text-figs. 33, 34)

Macropterous forms only known ; narrow and slender, of medium size, with comparatively
very long antennae and moderately long legs. Mounted specimens 938-1260 (no6)u long,
266-322 (294) (jl wide at mesothorax and 2254-2506 (2366)0 wing expanse.

Body setae and pores : Body and appendages with numerous hair-like setae ; fleshy setae
entirely absent. Antennal setae about 61 o long, body setae about 241* long and leg setae of
intermediate length. Ouadrilocular and occasionally quinquelocular disc pores present on
head, thorax and abdomen, about 60 in diameter.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression well pronounced in lateral view. Length from apex to postoccipital ridge 98-122
(104)0 ; from apex to neck 143-165 (153) o ; width across the genae 159-198 (180) o. Dorsal
arm of midcranial ridge absent, though its position marked by a sclerotization ; anteriorly
continuous with the ventral arm and posteriorly fading out before reaching the postoccipital
ridge. Lateral and ventral arms well developed. Postoccipital ridge indicated by a slender,
U-shaped sclerotization, anteriorly confluent with the preocular ridges. Dorsomedial part
of epicranium slightly raised. Preocular ridge widely separated from postocular ridge ; inter-
ocular ridge absent ; preoral ridge slender. Eyes : Dorsal simple eyes projecting beyond the
outer margins of the head in dorsal view ; the diameter of their corneae 15-21 (17)0, and
separated by 107-137 (122)0., i.e. 5-4-90 (6-9) times their diameter apart. Ventral eyes also
15-21 (19) o in diameter and 31-46 (37)0 apart. Lateral ocelli well developed, attached to the
postocular ridges by means of a short sclerotized arm. Ocular scleriles large. Cranial
apophysis apically truncate ; tentorial bridge slender. Dorsal head setae on each side 8-1 1
(9-7) anterior to the postoccipital ridge ; each gena almost always with 3 and occasionally 2
(av. 2-8) genal setae. Ventral head setae on each side : 1-2 (1-5) between the ventral eyes ;
6-9 (7-0) in the area of the ventral preocular depression forming on both sides a transverse
band ; 2-3 (2-8) on each side of the ventral arm of the midcranial ridge. Head disc pores :
Dorsal pores absent ; one ventral pore always present on each side.

Antennae : Filiform ; 10-segmented ; 760-970 (878)0. long, i.e. much longer than half the
body length (ratio 1 : 1-2-1-3, av - I-2 5) an d longer than the hind legs (ratio 1-2-1-4, av - *"3 : J )-
Scape 46-52 (49)0 long and 43-47 (46)0 wide at base ; always with 4 h.s. Pedicel 58-64 (61)0

156 MORPHOLOGY AND TAXONOMY OF ADULT MALES

long and 31-37 (34)fx wide ; with 8-16 (n - 3) h.s. and a sensillum placodeum. Flagellum :
Flagellar segments cylindrical and about 2i;x wide. Segment III longest of all, the ratio its
length to the length of segment X 1 -6-2-1 (i-8) : 1, and the ratio width to length of segments
III and IX 1 : 5-4-7-1 (6-4) and 1 : 3-6-4-5 (42), respectively. The following table shows
the lengths of the flagellar segments and the number of setae on each:

III IV V VI VII VIII IX X

length in u.

"6-153

95-131

92-119

76-113

73-95

67-85

70-82

61-85

(av.)

(137)

(113)

(107)

(98)

(85)

(76)

(76)

(76)

h.s.

17-28

IO-I8

10-15

9-14

9-13

9-12

8-15

12-18

(av.)

(22-8)

(i4-i)

(12-5)

(no)

(107)

(io-i)

(n-i)

(15-6)

Antennal bristles inconsiderably stouter than the antennal setae and slightly shorter ;
segments VIII and IX each with one ventral bristle. Segment X with three preapical bristles
and 2-3 preapical sensilla basiconica. Capitate, apically knobbed setae entirely absent.

Thorax : 388-479 (433) fx long. Prothorax : Pronotal ridges medially interrupted. Lateral
pronotal sclerites and post-tergites small. Dorsal and ventral margins of the proepisternum
with ridge-like sclerotizations (fig. 31). Prosternum triangular, 24-31 (27) ;x long ; posteriorly
supported by the transverse, 82-98 (89) [x long prosternal ridge. Prothoracic setae : Medial
pronotal, lateral pronotal, post-lergital and antespiracular dorsal setae all absent ; one ante-
spiracular ventral seta always present on each side. Prosternal setae usually absent, but one
seta was found on one side of one specimen. Prothoracic pores on each side : Medial pronotal
pores 2-4 (3-1) ; lateral pronotal pores 1-2 (1-3) ; antespiracular dorsal pores absent. Prosternal
pores usually 1-2, and sometimes absent on one side (av. 11).

Mesothorax : Prescutum triangular in dorsal view ; 76-98 (85)0. long and 92-119 (104) u.
wide, the ratio being 1 : 1-1-1-3 i 1 " 2 )- Presculal ridges continuous postero-medially replacing
the prescutal suture. Scutum very short and evenly sclerotized ; 27-46 (37) (x long, i.e. the
ratio lengths of prescutum to scutum 2-0-3-2 (2-3) : 1. Prealare, prealar ridge, triangular
plate, tegula, anterior and posterior notal wing processes well developed. Scutellum 58-73 (64) jx
long and 79-110 (92) jx wide, the ratio being 1 : 1-3-1-5 (1-4), and the ratio its length to the
length of scutum 1-4-2-2 (i-8) : 1. Postalare with well separated anterior and posterior postalar
ridges. Postnotal apophysis strong. Mesopleuron : Mesopleural ridge interrupted above the
coxal articulation ; basalare comparatively slender. M esepisternum and mesepimeron well
defined ; later opleurite moderately large. Mesosternum : Basisternum 101-128 (ii6)jx long
and 162-198 (180) (x wide ; marginal, precoxal ridges and ) 'urea well developed. Mesothoracic
spiracle 21-24 (23) fx wide at opening, with a 37-43 (40) [x long supporting bar. Mesothoracic
setae on each side : Prescutal setae usually 2 and occasionally 3 (av. 2-2) ; scutal setae 4-5 (4-5) ;
scutellar setae 1-2 (1-3). Tegular setae 1-4 (25). Postmesostigmatal setae 3-5 (3-7) in a latero-
ventral group. Basisternal setae 6-8 (7-2). Mesothoracic disc pores on each side ; Meso-
spiracular pores 1-2 (i-6) ; postmesostigmatal pores 1-2 (1-2).

Metathorax : Metapostnotal sclerites distinct and metapostnotal ridge weakly developed.
Metapleural ridge attenuating near the middle ; with a small metapleural wing process. Precoxal
ridge weak ; metepisternum, metepimeron and metasternal apophysis distinct. Metathoracic
spiracle identical with the mesothoracic. Metathoracic setae on each side : Metatergal setae
2-3 (2-5) ; metapleural setae 0-1 (o-i). Anterior metasternal setae always one, and posterior
metasternal setae 0-2 (0-9). Metathoracic disc pores on each side : Metatergal pores 1-2 (1-3) ;
metaspiracular pores 1-2 (i-i). Anterior metasternal pores 1-2 (1-3) and posterior metasternal
pores 0-1 (0-4).

Wings : 1008-1134 (1078) u. long and 378-448 (406) (x wide ; with 2-3 (28) alar setae and a
compact row of 2 minute circular sensoria. Hamulohalterae 79-98 (85) (x long and 21-24 (23)^
wide ; with one, 49-55 (52) jx long apically hooked seta, i.e. the ratio lengths of seta to hamulo-
haltera 1 : 1-5-1-9 (1-7).

OF I'SKIDOCOCCIDAE & ERIOCOCCIDAE

'57

Legs : Well developed and moderately long ; the middle legs shortest and the hind legs
longest ; the ratio length of the hind leg to the total body length i : 1-5-17 (i-6). Coxa
about 49 and trochanter about 24 ^ wide. Femur about 34 jz wide ; that of the fore leg longest
and that of the middle leg shortest ; the ratio width to length of the hind femur 1 : 4-7-5-8 (5-1).
Tibia about 210. wide ; with 2 apical spurs ; in fore leg the femur shorter than the tibia, the
ratio their lengths 1 : 1-2-1-4 ( I- 3) Tarsus about i8u. wide ; tarsal digitules absent. Claw

dmcr vhp

Fig. 33. Ceroputo pilosellae Sulc, dorsal and ventral view.

158

MORPHOLOGY AND TAXONOMY OF ADULT MALES

uniformly tapering to a sharply pointed apex ; ungual digitules fine, about i2(x long. The
following table shows the lengths of the leg segments (in microns) and the number of setae on
each:

Fore leg

Middle leg

Hind leg

Coxa

length
h.s.

40-46 (43)
11-14 (12-8)

40-46 (43)
11-15 (13-2)

43-49 (46)
13-16 (13-8)

Trochanter

length
h.s.

55-6i (58)
5-8 (67)

55-61 (58)
5-8 (6-8)

58-64 (61)
6-9 (7-3)

Femur

length
h.s.

174-217 ( x 98)
34-42 (36-8)

153-201 (180)
26-35 (30-9)

159-21 1 (186)
27-38 (30-4)

Tibia

length
h.s.

235-290 (256)
35-46 (41-1)

226-284 ( 2 47)
36-44 (39-5)

238-299 (265)
37-49 (42-4)

Tarsus

length
h.s.

89-107 (98)
26-31 (28-8)

89-107 (98)
26-33 (29-5)

92-110 (101)
28-35 (30-5)

Claw

length

27-34 (3i)

27-34 (3i)

27-34 (3i)

Total length

of leg

619-750 (680)

589-729 (653)

622-766 (692)

Abdomen : 357-562 (464)^ long and 258-304 (281)0. wide. Tergites of segments I, II and
sometimes III small ; those of segments VIII and IX + X distinct. Ostioles ill-defined.
Abdominal setae on each side : Dorsal setae 2-4 on segment I, 3-5 on segments II and III,
3-4 on segments IV to VII, and 2 on segment VIII. Pleural setae 1-2 on segment I, 2-3 on
segment II, 3-5 on segments III to VII, and always 3 on segment VIII. Ventral setae 1 on
segment II, 3-4 on segments III and IV, 2-3 on segments V to VII, and absent on segment
VIII. Abdominal pores : Pleural pores 3-5 (4-1), 1-2 (it), (i-o), 1-2 (it), (i-o), and (i-o) on
segments I to VI, respectively. Segments I to VII also with 0-1 (o-6), 1-2 (1-2), 1-2 (it),
0-2 (0-9), 0-1 (0-9), and 0-1 (o-8) dorsal pores, and segments II to VI with 0-2 (o-6), 0-2 (0-7),
0-1 (0-5), 0-1 (0-3), and 0-1 (o-i) ventral pores, respectively.

Glandular pouches well developed, occurring on each side of segments VII and VIII. Setae
of glandular pouch of segment VII include a pair of 168-207 (177) H- long tail setae ; those of
segment VIII include a pair of 189-223 (204)0 long tail setae, and one seta about as long as the
abdominal setae, i.e. the ratio length of the tail setae of segment VIII to the total body length
1 : 4-7-6-4 (5-4).

200

Fig. 34. Ceroputo pilosellae Sulc, lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 159

Genital segment comparatively small ; style rather straight in lateral view, with rounded
apex. Penial sheath 92-107 (98) |x long and 49-61 (55) (x wide, i.e. the ratio length to width
1-6-2-1 (i-8) : 1, and the ratio its length to the total body length 1 : 10-2-12-6 (11-3). Basal
ridge of penial sheath slender and its projection small. Process of penial sheath absent. Aedeagns
comparatively stout and short. Setae of genital segment : Dorsally 3 setae always present
on each side near the base of the style ; ventral setae absent, but 0-3 (i-i) setal sensilla occur on
each side of the penial sheath.

Material : 10 specimens examined, collected by N. Mitic-Muzina, on Fragaria
vesca, in Belgrade, Yugoslavia, 30.ix.1961.

CENTROCOCCUS Borchsenius, 1948

Centrococcus insolitus (Green)

(Text-figs. 35, 36)

Macropterous forms only known ; moderately robust, large, with comparatively very long
antennae and long and slender legs. When mounted, the total length of the body 1302-1512
(1386)^, width at mesothorax 350-392 (378)0. and wing expanse 3136-3556 (3374)1*.

Body setae and pores : Antennae with numerous, about 6ojx long fleshy setae and a few,
about 24 (x long, hair-like ones ; the legs also with fleshy (about 40^ long) and hair-like setae.
The body itself with hair-like setae only, yuadrilocular and occasionally trilocular disc pores
present on the head, thorax and abdomen, about 6[x in diameter.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression well pronounced in lateral view. Length from apex to postoccipital ridge 128-146
(137) u. ; from apex to neck 183-201 (192) \l ; width across the genae 211-238 (220)^. Dorsal
arm of midcranial ridge anteriorly continuous with the short ventral arm, and posteriorly meeting
the postoccipital ridge. Lateral arms only indicated by short sclerotizations. Postoccipital
ridge anteriorly confluent with the preocular ridges. Preocular ridge completely separated
from the postocular ridge. Interocular ridge replaced by a short arm attached to the postocular
ridge, and dorsally supporting the ocellus. Preoral ridge slender. Eyes : Dorsal simple eyes
projecting beyond the outer margins of the head in dorsal view ; their corneae 27-34 (31) jx
in diameter and separated by 122-156 (i^j)\t, i.e. 3-7-5-7 (45) times their diameter apart.
Ventral simple eyes 31-43 (37) (x in diameter and 27-43 (34)^ apart. Lateral ocelli well
developed, dorsally supported by the ridge-like arm as mentioned. Cranial apophysis apically
truncate. Tentorial bridge slender. Dorsal head setae 7-12 (9-8) on each side anterior to the
postoccipital ridge ; each gena usually with 1-3 genal setae, but in few specimens the setae
of either gena were absent (av. i-6). Ventral head setae on each side : Setae between the
ventral eyes absent ; 5-8 (6-2) in the area of the ventral preocular depression, forming on both
sides a transverse band ; 2-3 (2-3) on each side of the ventral arm of the midcranial ridge.
Head disc pores : One dorsal and one ventral head pore usually present a least on one side of
the dorsal and ventral arms of the midcranial ridge (averages 06 and 09 respectively).

Antennae : Filiform ; 10-segmented ; 875-1254 (1095) [x long, i.e. longer than half the
total length of the body (ratio 1 : i»i— 1-5, av. 1-3), and longer than the hind legs (ratio 1-1-1-3,
av. 1-2 : 1). Scape 61-64 (63)^ long and 52-58 (55) (x wide at base ; always with 4 h.s.
Pedicel 61-73 (67) u. long and 37-40 (39) u. wide ; with 6-1 1 (7-3) f.s., 4-8 (5-3) h.s. and a
sensillum placodeum. Flagellar segments cylindrical, about 2ijx wide. Segment III longest of
all, the ratio lengths of segments III to X being 1-4-1-9 (i-6) : 1, and the ratio width to length

160 MORPHOLOGY AND TAXONOMY OF ADULT MALES

of segments III and IX 5-3-7-4 (6-5) : 1 and 3-7-5-1 (4-4) : 1, respectively. In the following
table the lengths of the flagellar segments and the number of setae on each are given :

III IV V VI VII VIII IX X

length in [o

128-180

101-165

98-168

92-153

85-!34

82-122

79-110

85-H6

(av.)

(159)

(137)

(137)

(128)

(116)

(107)

(95)

(98)

f.s.

15-21

1 1-19

13-21

1 1-2 1

12-16

13-17

12-14

IO-I5

(av.)

(18-3)

(14-3)

(17-5)

(16-1)

(14-6)

(14-3)

(13-2)

(I3-0)

h.s.

5-8

i-3

i-3

1-2

1-2

1-2

1-2

1-2

(av.)

(7-i)

(1-6)

(1-9)

(1-2)

(i-3)

(i-i)

(1-2)

(1.4)

Antennal bristles inconsiderably stouter than the fleshy setae. Segments VIII and IX each
with a ventral bristle, about 3i[x long. Segment X with 3 preapical bristles, about 55(1 long
and two much shorter ventral ones, about i8[o long, at a greater distance from the segment's
apex ; terminal segment also with a preapical sensilla basiconica. Capitate, apically knobbed
sensory setae entirely absent.

Thorax : 532-623 (562)10 long. Pvothorax : Pronotal ridges medially interrupted as usual.
Lateral pronotal sclerites and post-tergites well defined. Dorsal and ventral margins of the
proepisternum ridge-like. Prosternum represented by a small plate on each side of the median
line ; prosternal ridge 107-128 (ii6)jo long. Prothoracic setae on each side : Medial and lateral
pronotal setae usually absent, but one seta of each was observed on one side of one specimen ;
post-tergital setae 0—2 (0-5) ; antespiracular dorsal setae absent. One antespiracular ventral seta
always present ; prosternal setae 0-1 (0-3). Prothoracic disc pores on each side : Medial and
lateral pronotal pores 1-3 (averages 1-4 and 2-1, respectively) ; antespiracular dorsal pores
absent. Prosternal pores 1-3 (1-7).

Mesothorax : Prescutum 122-162 (140)}! long and 131-146 (137)^ wide, the ratio being
1 : 0-9-1-1 (i-o). Prescutal ridges well developed and continuous medially, replacing the
prescutal suture. Scutum comparatively very short and evenly sclerotized ; 21-31 (27)10. long,
i.e. the ratio lengths of prescutum to scutum 4-3-6-4 (51) : 1. Prealare, prealar ridge,
triangular plate, tegular, anterior and posterior notal wing processes well developed. Scutellum
laterally supported by a strong additional ridge ; 85-101 (92)10 long and 131-143 (137) [0 wide,
i.e. the ratio length to width 1 : 1-4-1-7 (1-5), and the ratio lengths of scutellum to scutum 30-
4-0(3-3) : 1. Postalare with well separated anterior and posterior postalar ridges. Mesopleuron :
Mesopleural ridge interrupted above the coxal articulation ; basalare comparatively slender.
Mesepisternum and mesepimeron well defined ; later opleurite moderately large, with a distinct
oval semi-membranous area. Mesosternum : Basisternum 146-183 (165) \x long and 226-250
(235)10 wide ; marginal, precoxal ridges and furca well developed. Mesothoracic spiracle 21-24
(23) (x wide at opening, with a 43-49 (46) u. long supporting bar. Mesothoracic setae on each
side : Prescutal setae 6-9 (7-2) ; scutal setae 2-4 (2-9) ; scutellar setae 1-2 (i-8). Tegular setae
3-6 (4-7). Postmesostigmatal setae 1-3 (1-3) occurring sublaterally. Basisternal setae 8-16
(11 -6). Mesothoracic disc pores : Mesospiracular pores 2-4 (2-6) ; postmesostigmatal pores
absent.

Metathorax : Metapostnotal sclerites well defined ; metapostnotal ridge weakly developed.
Metapleural ridge attenuated near the middle as usual. Precoxal ridge weakly developed, and
metasternal apophysis distinct. Metathoracic spiracle identical with the mesothoracic. Meta-
thoracic setae on each side : Metatergal setae 3-4 (3-2) ; metapleural setae 1-2 (i-i). Anterior
metasternal setae 0-1 (o-8) ; posterior metasternal setae 0-2 (0-9). Metathoracic pores on each
side: Metaspiracular pores 1-2 (1-3) ; anterior metasternal pores 0-2 (1-2) ; posterior metasternal
pores 0-1 (0-3).

Wings : 1400-1610 (1484) jo long and 462-546 (504)^1 wide ; with 2-3 (2-3) alar setae and a
compact row of 2-3 (2-2) minute circular sensoria. Hamulohalterae 1 13-122 (ii"9)[o long and
21-24 (23) H wide ; with one, 52-61 (55)10. long apically hooked seta, i.e. the ratio lengths of
seta to hamulohaltera 1 : 1-9-2-3 (2-2).

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

161

Legs : Comparatively long and slender ; the middle legs shortest and the hind legs longest ;
the ratio length of the hind leg to the total length of the body i : i -2-1-6 (1-4). Coxa and
trochanter about 58 and 31^ wide respectively. Femur about 400. wide ; that of the fore leg

Fig. 35. Centrococcus insolitus (Green), dorsal and ventral view.

162

MORPHOLOGY AND TAXONOMY OF ADULT MALES

longest and those of middle and hind legs subequal in length ; the ratio width to length of the
hind femur i : 6-5-7-5 (6-9). Tibia about 2i(x wide ; with 2 apical spurs and usually 2 smaller
spines ; the fore leg the femur shorter than the tibia, the ratio their lengths being 1 : 1-3-1-4
(1-33). Tarsus about i8jx wide ; tarsal digitules entirely absent. Claw gradually tapering to
a sharply pointed tip ; ungual digitules fine. The lengths of the leg segments (in microns) and
the number of setae on each are given in the following table :

Coxa

length
f.s.

h.s.

Trochanter

length
f.s.

h.s.

Femur

length
f.s.

h.s.

Tibia

length
f.s.

h.s.

Tarsus

length
f.s.

h.s.

Claw

length

Total length of leg

Fore leg

49-55 (52)

9-13 (n-6)

2-5 (3-8)
70-76 (73)

4-6 (4-8)

2-3 (2-4)

268-308 (287)

28-39 (33-4)

6-9 (yi)

35I-42I (381)

47-56 (51-8)

6-8 (6-7)

113-131 (119)

23-27 (24-6)

6-8 (6-9)

37-43 (40)
888-1025 (949)

Middle leg

49-55 (52)
10-13 (12-5)

2-4 (3-i)

70-76 (73)

4-6 (4-5)

2-3 (2-6)

259-296 (275)

27-36 (31-2)

6-8 (6-9)

35I-4I5 (372)

43-53 (48-2)

6-10 (8-3)

II3-I34 (122)
23-29 (25-1)

6-9 (7-6)
37-43 (40)

878-1016 (933)

Hind leg

52-58 (55)
10-14 (12-7)

3-5 (4-2)
73-79 (76)

4-7 (4-5)

2-4 (27)

259-296 (275)

28-38 (32-6)

7-10 (7-8)

387-451 (409)

48-62 (57-7)

8-12 (104)

128-150 (134)
27-35 (32-8)

7-i 1 (8-3)
37-43 (40)

939-1074 (988)

Abdomen : 494-661 (555) \l long and 327-365 (342)^ wide. Tergites of segments I, II, III
and sometimes IV small ; those of segments VIII and IX + X well sclerotized. Sternites
of segment VIII ill-defined. Ostioles well developed, 37-43 (40) \i long at orifice. Abdominal
setae on each side : Dorsal setae 3-4 on segment I, 2-3 on segments II to VI, 3-4 on segment
VII, and 1-3 on segment VIII. Pleural setae 3-4 on segment I, 4-6 on segment II, 5-6 on

200^-

100.

Fig. 36. Centrococcus insolitus (Green), lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 163

segments III to VI, and 3-4 on segments VII and VIII. Ventral setae 1-2 on segment II, 2-3
on segment III, 2 on segments IV to VII, and absent on segment VIII. Abdominal pores :
Pleural pores 2-3 (2-4) on segment I, 0-1 (0-2) on segment IV, 1-2 (13) on segment VI, and
absent on other segments. Dorsal pores 1-3 (1-9). 1-2 (1-2), 1-2 (11), 0-2 (0-9), 0-1 (07),
0-2 (08), and 0-1 (07) on segments I to VII respectively. Ventral pores also sometimes
present, 0-2 (07), 0-1 (03), and 0-1 (02) on segments III, IV and V respectively.

Glandular pouches occur on each side of segments VII and VIII. Setae of the glandular
pouch of segment VII include a pair of 168-204 (183)0- long tail setae only ; those of segment
VIII include a pair of 198-229 (214)0. long tail setae and a short one, subequal to the abdominal
setae in length, i.e. the ratio length of the tail setae of segment VIII to the total body length
1 : 5-6-71 (6-5).

Genital segment comparatively small ; style rather straight in lateral view, with a rounded
apex. Penial sheath 107-116 (113)0 long and 73-82 (76)0 wide, the ratio length to width
being 1-3-1 -6 (15) : I, and the ratio its length to the total body length 1 : 11 -8-13-4 (12 -3).
Basal ridge of penial sheath slender and its projection small. Process of penial sheath hardly
pronounced. Aedeagus comparatively stout. Setae of genital segment on each side : Dorsal
setae 5-7 (6-4) near the base of the style ; ventral setae 2-4 (2-8) on the penial sheath.

Material : 10 specimens examined, collected by H. K. Patel, on Cajanus indicus,
in India (Anand), 14.xii.1955 (received from Dr. D. J. Williams).

NAIROBI A Group

NAIROBI A De Lotto, 1964

Nairobia bifrons De Lotto

(Text-figs. 37, 38)

Winged forms only known ; narrow and slender, comparatively large, with moderately long
antennae and legs. The mounted males 1260-1596 (1442)0 long, 308-350 (336)0 wide at
mesothorax and 2814-3528 (3248)0 wing expanse.

Body setae and pores : Fleshy setae entirely absent. Hair-like setae present on the antennae,
legs and body, about 210 long. Disc pores absent altogether.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression hardly indicated in lateral view. Length from apex to postoccipital ridge 128-140
(131)0 ; from apex to neck 177-195 (186)0 ; width across the genae 220-241 (232)0. Dorsal
arm of midcranial ridge marked by a narrow sclerotization ; this sclerotization continuous
with those indicating the lateral and the ventral arms anteriorly, and almost meeting the post-
ocular ridge posteriorly. Postoccipital ridge confluent with preocular ridges. Preocular ridge
well developed and completely separated from postocular ridge ; interocular ridge absent ;
preoral ridge slender. Eyes : Dorsal simple eyes projecting beyond the outer margins of the
head in dorsal view ; their corneae 18-24 ( 2I )i jL m diameter and separated by 125-162 (146)0,
i.e. 5-5-7-6 (6-9) times their diameter apart. Ventral simple eyes 21-24 (23)0 in diameter and
34-55 (46) o apart. Lateral ocelli vestigial. Ocular scleriles large. Cranial apophysis truncate ;
tentorial bridge slender. Dorsal head setae on each side : Setae in front of the postoccipital ridge
in two separable groups, 3-5 (4-1) anteriorly and 4-6 (4-4) posteriorly ; each gena usually with
2 genal setae but 1 or 3 setae may occur on one side (av. 2-1). Ventral head setae on each side :
One seta may occur between the ventral eyes (av. 0-4) ; 8-1 1 (8-8) forming on both sides a
transverse band in the area of the ventral preocular depression ; 2-3 (2-2) in a longitudinal
row anteriorly.

Antennae : Filiform ; normally 10-segmented but fusion between two or more adjacent
flagellar segments on one or both antennae very common ; 799-933 (894)0 long, i.e. much

164 MORPHOLOGY AND TAXONOMY OF ADULT MALES

longer than half the body length, the ratio i : 1-4-17 (i-6), and subequal in length to the hind
legs, the ratio i-o-i-i (1-08) : 1. Scape 58-64 (6i)pi long and 52-58 (55)^ wide at base ; always
with 4 h.s. Pedicel 76-85 (79)0. long and 43-46 (45) (i. wide ; with 10-13 ( ri '6) h.s. and a
sensillum placodeum. Flagellum : Flagellar segments cylindrical and 21-24JJ. wide ; segment
III approximately as long as segment X, both being usually longest, the ratio their lengths
0-9-1-1 (1-06) : 1, and the ratio width to length of segments III and IX 1 : 3-4-4-2 (3-9) and
1 : 3-5-4-0 (3-7) respectively. Apart from the hair-like setae, the flagellar segments also with
numerous apically knobbed, capitate sensory setae. The lengths of the flagellar segments and
the number of setae on each are given below :

Ill

IV

V

VI

VII

VIII

IX

X

length in u.

89-116

73-io4

89-107

95-116

85-107

79-95

79-92

85-98

(av.)

(98)

(92)

(95)

(107)

(101)

(89)

(85)

(92)

h.s.

6-9

3-6

i-3

3-4

2-4

i-3

1-2

1-2

(av.)

(7-3)

(4-4)

(i-9)

(37)

(2-7)

(i-7)

(i-i)

(i-6)

set. ca.

2-5

2-5

7-1 1

8-13

7-12

6-12

7-i5

9-13

(av.)

(3-6)

(4-3)

(87)

(ii-i)

(IO-2)

(9-4)

(ii-i)

(10-9)

Antennal bristles well defined ; segments VIII and IX each with one ventral bristle. Segment
X with 3 preapical bristles, the dorsal of which is slightly shorter ; terminal segment also with
one apical hair-like seta.

Thorax : 502-578 (540) u. long. Prothorax : Pronotal ridges medially interrupted as usual ;
lateral pronotal sclerites and post-tergites well defined. Proepisternum with a ridge-like dorsal
margin. Prosternum very short, triangular, 12-18 (15) jx long ; prosternal ridge 101-113 (107) y.
long. Prothoracic setae : Medial pronotal, lateral pronotal and post-tergital setae absent.
Antespiracular dorsal setae 2-4 (3-1) ; antespiracular ventral seta always 1 ; prosternal setae
2-3 (2-4)-

Mesothorax : Prescutum triangular in dorsal view ; 95-113 (107)0. long and 1 16-128 (125) ^
wide, the ratio being 1 : 1-1-1-3 ( I-2 )- Prescutal ridge well developed and prescutal suture
distinct. Scutum comparatively short and evenly sclerotized ; 55-67 (6i)jjl long, i.e. the ratio
lengths of prescutum to scutum 1-6-2-1 (i-8) : 1. Prealare, prealar ridge, triangular plate,
tegula, anterior and posterior notal wing processes well developed. Scutellum 82-98 (92)0. long
and 107-119 (113)^ wide, ratio 1 : 1-2-1-3 (1-23), and the ratio its length to the length of
scutum 1-4-1-7 (1-5) : 1. Postalare with well separated anterior and posterior postalar ridges.
Mesopleuron : Mesopleural ridge interrupted above the coxal articulation ; basalare slender.
Mesepisternum and mesepimeron well defined ; later opleurite moderately large, with a distinct
oval membranous area. Mesosternam : Basisternum 153-168 (162) fj. long and 198-232 (223)11
wide ; marginal, precoxal ridges and j r urca well developed. Mesothoracic spiracle 24-27 (26) u.
wide at opening, with a 43-49 (46) jx long supporting bar. Mesothoracic setae on each side :
Prescutal setae 1-2 (1-7) ; scutal setae 1-3 (i-8) ; scutellar setae usually 1 and occasionally 2
(i-i). Tegular setae 3-4 (3-7). Postmesostigmatal setae absent. Basisternal setae (stn25) 2-4
(3-6).

Metathorax : Metapostnotal sclerites distinct but the metapostnotal ridge absent. Metapleural
ridge with the usual attenuation near the middle. Precoxal ridge weakly developed ; meta-
sternal apophysis distinct. Metathoracic spiracle similar to the mesothoracic. Metathoracic
setae on each side : Metatergal setae always 2 ; metapleural setae usually 1-2 but sometimes
missing on one side (av. o-8). Anterior and posterior metasternal setae 1-2 (averages 1-3 and
i-i respectively).

Wings : 1260-1610 (1470) [i. long and 420-532 (490)^ wide ; alar setae absent and circular
sensoria could not be detected and presumably also absent. Hamulohalterae 1 10-125 ( II 9)( 1
long and 24-31 (28) u. wide ; with one apically hooked seta, 40-49 (46)^ long, the ratio lengths
of seta to the hamulohaltera being 1 : 2-4-3-0 (2-6).

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

165

Legs : Moderately long ; fore legs shortest and hind legs longest ; the ratio length of the
hind leg to the total body length 1 : 1-5-20 (i-8). Coxa and trochanter about 55 and 31^
wide. Femur about 40^ wide ; that of the fore leg shortest and that of the hind leg longest ;
the ratio width to length of the hind femur 1 : 5-4-6-2 (5-8). Tibia about 2iu. wide ; with 2
apical spurs ; in front leg, the femur slightly shorter than the tibia, the ratio their lengths being

dmcr

vhs

y^~

'■■■^^

,-vse

i ,

y ocs

*)

/ pocr

por

fV

J pepcv

Fig. 37. Nairobia bifrons De Lotto, dorsal and ventral view.

1 66

MORPHOLOGY AND TAXONOMY OF ADULT MALES

i : I-I-I-2 (1-17). Tarsus about i8(x wide ; tarsal digitules absent. Claw gradually tapering
to a sharply pointed tip ; ungual digitules fine. The lengths of the leg segments (in microns)
and the number of setae on each are given in the following table :

Fore leg

Middle leg

Hind leg

Coxa

length
h.s.

49-55 52)
6-9 (7-1)

49-55 (52)
6-10 (7-9)

52-58 (55)
8-1 1 (9-4)

Trochanter

length
h.s.

64-73 (69)
4-6 (5-2)

64-73 (69)
4-6 (5-o)

67-76 (73)
4-7 (5-6)

Femur

length
h.s.

207-241 (226)
15-23 (19-2)

214-244 (229)
18-25 (20-6)

214-247 (232)
16-24 ( x 9'8)

Tibia

length
h.s.

244-281 (265)
16-22 (198)

250-293 (278)
17-26 (21-4)

268-311 (296)
19-30 (26-2)

Tarsus

length
h.s.

107-122 (116)
"-15 (13-3)

107-122 (119)
11-14 (12-8)

113-128 (122)
12-16 (14-5)

Claw

length

31-37 (34)

31-37 (34)

31-37 (34)

Total length

of leg

702-796 (744)

720-821 (784)

747-857 (814)

Abdomen : 464-707 (623)^ long and 319-365 (350) (x wide. Tergites of segments I and II small ;
those of segments VIII and IX + X well defined. Sternites of segment VIII small. Ostioles
entirely absent. Abdominal setae on each side : Dorsal setae 2 on segment I, 2-3 on segments
II to VII, and absent on segment VIII. Pleural setae 2 on segment I, 2-3 on segment II, 3-4
on segments III to V, 3-5 on segments VI and VII, and 2-3 on segment VIII. Ventral setae
2 on segment II, 3-5 on segments III to V, 2-3 on segments VI and VII, and absent on segment
VIII.

Setae of glandular pouch include a pair of 122-153 (137)^ long tail setae and one seta about as
long as the abdominal setae, i.e. the ratio length of tail setae to the total body length 1 : 8-2-
n-9 (10-5).

Genital segment comparatively small ; style rather straight in lateral view, with a pointed
apex. Penial sheath 134-146 (140)^ long and 92-98 (95)10. wide, the ratio being 1-4-1-6 (1-5) : 1,
and the ratio its length to the total length of the body 1 : 9-2-11-5 (10-3). Basal ridges of

Fig. 38. Nairobia bifrons De Lotto, lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 167

penial sheath medially continuous ; their projections as well as the processes of penial sheath
absent. Aedeagns with a large internal genital aperture. Setae of genital segment : Dor sally
3 setae always present on each side near the base of the style ; ventrally 2-3 (2-1) smaller setae
occur on each side of the penial sheath. Style also usually with 3 latero-dorsal minute sensilla
on each side.

Material : 10 specimens examined, collected by G. De Lotto, on Lonicera
battiscombei Hutch., in Nairobi, Kenya, 8.xi.io,62.

ERIOCOCCIDAE

ERIOCOCCUS Targioni-Tozetti, 1868

Eriococcus araucariae Maskell

(Text-figs. 39, 40)

Macropterous forms only known ; living specimens light brown in colour. A moderately
robust species, moderately large, with comparatively short antennae and moderately long
legs. Mounted specimens 1078-1344 (1218)0 long, 308-364 (336) (j. wide at mesothorax and
2016-2324 (2170)0. wing expanse.

Body setae : Antennae and legs with numerous, approximately 15a long fleshy setae and
slightly longer hair-like ones ; the body with hair-like setae only.

Head : Subtetrahedron ; subtriangular in dorsal view and more or less rounded in front
view ; ventral preocular depression hardly indicated in lateral view. Length from apex to
postoccipital ridge 420-518 (462)0 from apex to neck 756-826 (798)0 ; width across the genae
980-1106 (1036)0. Midcranial ridge cruciform ; dorsal arm short, jointed to the ventral and
the lateral arms at the apex of the head (Text-fig. 39, B). Postoccipital ridge strongly developed,
with the lateral extremities sharply bent posteriorly and giving off a short sclerotization
anteriorly. Dorsomedial part of epicranium narrow and slightly raised. Preocular and post-
ocular ridges widely separated ; the latter ventrally articulates with a small additional ridge ;
preoral ridge slender. Eyes : Dorsal simple eyes projecting well beyond the outer margins of
the head in dorsal view ; their corneae 34-40 (38)0 in diameter and separated by 1 10-125
(116)0, i.e. 2-9-3-4 (3' 2 ) times their diameter apart. Ventral simple eyes 27-34 (3 1 )" m
diameter and 21-24 ( 2 3) u apart. Lateral ocelli anteriorly attached to the postocular ridges.
Ocular sclerites weakly sclerotized. Cranial apophysis apically bifurcate. Anterior tentorial
arms join the cranial apophysis separately ; tentorial bridge comparatively stout. Dorsal
head setae 8-1 1 (9-1) on each side ; each gena with 3-6 (43) genal setae. Ventral head setae
on each side : Setae between the ventral eyes absent ; 14-22 (17-8) forming on both sides a
transverse band in the area of the ventral preocular depression ; 3-5 (41) longitudinally
arranged anteriorly. A peculiar organ of obscure homology (X) always present on each side
of the dorsal arm of the midcranial ridge near the apex of the head.

Antennae : Rather filiform ; normally 10-segmented, with occasional fusion between two
adjacent segments of the fiagellum ; 516-622 (564)0 long, i.e. as long as or slightly shorter
than half the total length of the body, the ratio being 1 : 2-0-2-3 ( 2 ' 2 )> ar >d nearly as long as
the hind legs, the ratio 1 : 1-0-1-2 (i-i). Scape 40-46 (42)0 long and 37-43 (40)0 wide at
base ; with 4-5 (4-3) h.s. Pedicel 61-70 (67)0 long and 34-40 (37)0 wide ; with 13-24 (168)
f.s., 20-28 (26-0) h.s. and a sensillum placodeum. Fiagellum : Flagellar segments about 240
wide, with the preterminal segments somewhat wider. Segment III club-shaped and longest
of all, the ratio its length to the length of segment X 1-4-1-7 (1-5) : 1, and the ratio its width
to its length 1 : 3-4-4-0 (3*8). Segments IV to IX barrel-shaped, the ratio width to length

168 MORPHOLOGY AND TAXONOMY OF ADULT MALES

of segment IX being i : 1-4-1-9 (1-7) ; terminal segment pear-shaped. Segments V or VI to
X always with a few apically knobbed capitate sensory setae. The following table shows the
lengths of the flagellar segments and the number of setae on each :

III IV V VI VII VIII IX X

Length in fx

82-98

55-73

49-58

49-55

43-52

40-49

40-46

52-61

(av.)

(89)

(64)

(55)

(52)

(49)

(45)

(45)

(58)

f.s.

14-19

11-22

12-19

13-18

12-20

IO-I7

6-IO

i-4

(av.)

(17-0)

(167)

(15-4)

(157)

(14-5)

(12-7)

(8-i)

(2-7)

h.s.

9-12

7-10

7-10

7-10

6-1 1

6-IO

4-10

5-8

(av.)

(107)

(87)

(8-4)

(9-1)

(9-2)

(8-i)

(7-i)

(6-3)

Setc. ca.

0-2

i-3

1-4

2-4

2-5

3-6

(av.)

(0-9)

(i-5)

(2-1)

(2-5)

(3-o)

(4-1)

Antennal bristles stouter and longer than the fleshy setae ; segments VIII and IX each with
a ventral bristle. Terminal segment with 3 bristles, the dorsal of which slightly shorter and
nearer to the apex ; the segment also with a lateral pair of much smaller bristles and one
apical hair-like seta. Jancke (1955) recorded 2 bristles only occurring on the terminal segment.

Thorax : 517-585 (547) u. long. Prothorax : Comparatively short ; pronotal ridges medially
interrupted at a weakly sclerotized point ; lateral pronotal sclerites and post-tergites ill-defined.
Proepisternum without any ridge-like sclerotization ; propleural ridge short and propleural
apophysis small. Prosternum triangular, 58-76 (67) [x long ; prosternal ridge 92-107 (ioi)(x
long, with a small, crescent-like extension antero-medially. Prothoracic setae : Medial
pronotal setae usually absent although one seta was found in one specimen ; lateral pronotal,
post-tergital and antespiracidar dorsal setae absent. Antespiracular ventral setae always one
on each side ; prosternal setae 0-1 (o-8).

Mesothorax : Prescutum 70-79 (76)^1 long and 131-156 (i43)(x wide (ratio 1 : 1-8-2-0, av.
1-9) ; laterally and posteriorly bounded by the prescutal ridges and the prescutal suture
respectively. Scutum short and evenly sclerotized ; 40-49 (43) jx long, i.e. the ratio lengths of
prescutum to scutum 1-6-1-9 (i*8) : I. Prealare, prealar ridge, triangular plate, tegula, anterior
and posterior notal wing processes well developed. Scutellum transverse, rectangular in dorsal
view ; 52-61 (58)^ long and 113-140 (i28)fx wide, i.e. the ratios its length to its width 1 : 2-0-
2-3 (2-2), and its length to the length of scutum 1-3-1-5 (1-4) : 1 ; the scutellum obliquely
transversed on each side by a well developed scutellar ridge dividing it into 3 triangular parts.
Anterior and posterior postalar ridges anteriorly joined. Postnotal apophysis strong. Meso-
pleuron : Mesopleural ridge continuous above the coxal articulation ; mesopleural apophysis
and mesopleural wing process well developed ; basalare slender. Subepisternal ridge distinct.
Mesepisternum weakly sclerotized and mesepimeron small ; later opleuvite comparatively large.
Mesosternum : Basisternum 125-156 (140) |x long and 168-214 (i8g)(x wide ; marginal, precoxal
ridges and furca well developed. Mesothoracic spiracle 21-24 (22) (x wide at opening, with a
37-43 (40) |x long supporting bar. Mesothoracic setae on each side : Prescutal setae 1-2 (i-6) ;
scutal setae 2-4 (2-9) ; scutellar setae 1-2 (i-i). Tegular setae 4-5 (4-2). Postmesostigmatal
and basisternal setae absent.

Metathorax : Metapostnotal sclerites well defined ; rnetapostnotal ridge absent. Metapleural
ridge attenuated towards the small metapleural apophysis and dorsally supporting a small
pleural wing process. Metepisternum and metepimeron distinct. Precoxal ridge and metasternal
apophysis well developed. Metathoracic spiracle identical with the mesothoracic. Meta-
thoracic setae on each side : Metatergal setae always 3, one of which separated submedially ;
metapleural setae absent ; anterior metasternal setae 0-2 (i-i) and posterior metasternal setae
missing.

Wings : 868-994 (9 2 4)[x long and 378-434 (406)10. wide ; always with 2 alar setae. Hamulo-
halterae 67-76 (73) jx long and 18-21 (19) jx wide ; with one apically hooked, 67-79 (76) [x long
seta, i.e. the ratio lengths of seta to hamulohaltera i-o-i-i (1-04) : 1.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

169

Legs : Moderately long ; the fore and the middle legs subequal, and the hind legs longest ;
the ratio length of the hind leg to the total body length 1 : 1-8-2-1 (20). Coxa about 46 and
trochanter about 24 |x wide. Femur about 370. wide ; that of the fore leg usually longest and
that of tne middle leg shortest ; the ratio width to length of the hind femur 1 : 4-2-4-8 (4-6).
Tibia about 2i[x wide ; with 2 apical strong spurs and 2-4 smaller spines ; in fore leg the femur
shorter than the tibia, the ratio their lengths being 1 : 1-29-1-33 (1-3). Tarsus about i8u.

pdc dhs

Fig. 39. Eriococcus araucariae Maskell, dorsal and ventral view

170

MORPHOLOGY AND TAXONOMY OF ADULT MALES

wide ; tarsal digitules apically knobbed, 24-27 (26) (i long. Claw broad at base, abruptly
tapering to a pointed tip ; with a pair of apically knobbed ungual digitules. The following
table shows the length of the leg segments (in microns) and the number of setae on each :

Fore leg

Middle leg

Hind leg

Coxa

length

f.s.

h.s.

40-43 (41)
8-12 (9-6)
10-15 (12-4)

40-46 (42)
9-16 (n-8)
11-14 (12-2)

46-49 (47)
11-19 (136)
10-13 ("-8)

Trochanter

length

f.s.

h.s.

58-64 (61)

3-5 (4-2)
6-9 (8-8)

61-70 (65)
4-8 (6-i)
6-11 (8-2)

64-73 (70)
5-9 (6-8)
6-1 1 (8-o)

Femur

length

f.s.

h.s.

159-183 (168)
22-31 (26-2)
29-39 (35-i)

137-165 (15°)
19-28 (24-6)
26-31 (29-0)

153-177 (162)
23-29 (25-9)
26-33 (302)

Tibia

length

f.s.

h.s.

207-244 (217)

35-49 (440)
42-61 (526)

214-250 (229)
40-52 (48-2)
44-63 (55-2)

226-256 (238)
42-55 (496)
46-67 (57-4)

Tarsus

length

f.s.

h.s.

73-82 (76)
10-13 (n-8)
16-27 (22-2)

79-85 (82)
10-13 (n-4)
17-27 (22-4)

79-92 (83)

IO-I4 ( II-2 )
22-28 (24-4)

Claw

length

15-18 (17)

15-18 (17)

I5-I8 (l 7 )

Total length of leg

549-631 (573)

546-634 (593)

586-653 (619)

Abdomen : 327-479 (410)11 long and 281-334 (3 I2 )[i wide. Tergites of segments I and II
small ; that of segment VIII large. Sternites of segment VIII ill-defined. Ostioles absent.
Abdominal setae on each side : Dorsal setae 2-3 on segment I, 2 on segments II to VII, and
one slightly longer seta on segment VIII. Pleural setae 2 on segment I, 3-4 on segments II
to VI, and 4-5 on segments VII and VIII, including one slightly longer seta on the latter.
Ventral setae 2-3 on segments II to IV, 3-4 on segments V to VII, and 1-2 on segment VIII.

Glandular pouches small ; setae of glandular pouch include a pair of 198-217 (207)^ long tail
setae, the ratio their length to the total length of the body being 1 : 5-4-6-3 (5-9).

200^-

Fig. 40. Eriococcus araucariae Maskell, lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 171

Genital segment : Genital capsule 137-156 (143)^ long and 73-79 (76)1/ wide, i.e. the ratio
its length to its width 1-8-2-0 (1-9) : 1, and its length to total body length 1 : 7-9-8-9 (8-5) ;
style undulating in lateral view. Anal opening 18-21 (20) [x in diameter. Ventrally, a median,
longitudinal slender ridge present, giving off two lateral branches extending towards the base
of the aedeagus. Aedeagus strongly developed and ventrally sclerotized ; also with latero-
ventral heavy sclerotizations and a pair of small processes on each side. Setae of genital
segment : Dorsally two setae, about 61 jx long always present on each side near the base of the
style ; ventrally 5-6 (5-2) setae occur on each side of the genital capsule.

Material : 10 specimens examined, collected by myself on Araucaria excelsa,
in Alexandria, Egypt, U.A.R., during the last week of June and the first week of
July, 1961.

Eriococcus orariensis Hoy

(Text-figs. 41, 42)

Winged forms only known ; living specimens not available. The males moderately robust,
comparatively small, with short antennae and short legs. Mounted specimens 812-938 (882) ;x
long, 224-266 (25o)(x wide at mesothorax and 1652-1848 (1736)^ wing expanse.

Body setae : Antennae and legs with fleshy and hair-like setae, about 12 and i8(x long
respectively ; the body itself with slightly smaller hair-like setae only.

Head : Subtetrahedron ; subtriangular in dorsal view and rather rounded in front view ;
ventral preocular depression hardly indicated in lateral view. Length from apex to post-
occipital ridge 70-85 (76)^ ; from apex to neck 1 16-125 i 122 )^ '• width across the genae 156-
1 ^>5 ( I 59)M- Dorsal arm of midcranial ridge short, joined to the lateral and the ventral arms
at the apex of the head. Postoccipital ridge strongly developed, with the lateral extremities
bending posteriorly and giving off a short heavy sclerotization anteriorly. Dorsomedial part
of epicranium narrow. Preocular and postocular ridges widely separated ; the latter ventrally
articulates with a small additional ridge. Preoral ridge slender. Eyes : Dorsal simple eyes
slightly projecting beyond the lateral margins of the head ; their corneae 15-18 (16) ;x in
diameter and separated by 85-96 (89) u., i.e. 4-7-6-0 (5-4) times their diameter apart. Ventral
simple eyes subequal in diameter and 18-24 t 21 )^ apart. Lateral ocelli anteriorly attached to
postocular ridges. Ocular sclerites large. Cranial apophysis apically bifurcate. Tentorial
bridge stout. Dorsal head setae 8-10 (9-8) on each side ; each gena with 2-3 (2-7) genal setae.
Ventral head setae on each side : 3-5 (4-1) in the area of the ventral preocular depression,
forming on both sides a transverse band ; anteriorly a row of 2-3 (2-7) h.s. present on each
side of the ventral arm of the midcranial ridge.

Antennae : Rather moniliform ; normally 10-segmented ; 329-354 (339) [x long, i.e. shorter
than half the total length of the body, the ratio being 1 : 2-4-2-8), and about as long as the
hind legs, the ratio 1 : 1-1-1-2 (1-15). Scape 31-34 (32) jx long and 34^ wide at base ; always
with 4 h.s. Pedicel 43-46 (45)fx long and 27-31 (28) jx wide ; with 4-10 (6-6 f.s., 8-13 (11-3)
h.s. and a sensillum placodeum. Flagellum : Flagellar segments 18-2 ijx wide, with the pre-
terminal segments becoming slightly wider. Segment III club-shaped and longest of all, the
ratio its length to the length of the terminal segment 1-40-1-54 (1-5) : 1, and the ratio its width
to its length 1 : 2-9-3-3 (3 -2 )- Segments IV to IX barrel-shaped, the ratio width to length of
segment IX being 1 : 1-0-1-4 ( I-2 ) > terminal segment pear-shaped. Segments VIII, IX, X
and sometimes VII carry a number of apically knobbed capitate sensory setae. In the following
table the lengths of the flagellar segments and the number of setae on each are given :

172 MORPHOLOGY AND TAXONOMY OF ADULT MALES

III IV V VI VII VIII IX X

length in u.

58-61

27-31

24-31

21-27

24-31

24-31

24-31

40

(av.)

(60)

(29)

(27)

(24)

(27)

(27)

(27)

(40)

f.s.

6-10

2-5

3-8

4-8

3-8

3-6

2-5

2-3

(av.)

(77)

(3-i)

(6-o)

(5-8)

(5-8)

(3-9)

(3-3)

(2-1)

h.s.

3-6

3-5

3-5

2-4

3-7

3-8

5-8

2-5

(av.)

(4-8)

(3-3)

(4-i)

(3-2)

(5-2)

(5-6)

(6-1)

(3-3)

Set. ca.

O-I

1-2

2-3

5-7

(av.) 0000 (0-2) (1-3) (2-3) (5-6)

A ntennal bristles well denned ; segments VIII and IX each with one ventral bristle. Terminal
segment with 3 slightly longer bristles, the dorsal of which nearer to the apex, and one apical
hair-like seta.

Thorax : 350-403 (388) jx long. Prothorax : Pronotal ridges with the usual weak interruption
at the middle ; lateral pronotal sclerite and post-tergites small. Proepisternum without any
ridge-like sclerotization. Prosternum triangular, 18-31 (24)0. long ; prosternal ridge with a
small crescent-like antero-median extension, 67-79 (73)^ long. Prothoracic setae : Medial,
lateral pronotal, post-tergital and antespiracular dorsal setae absent. One antespiracular ventral
seta may be present (av. 0-4), and one prosternal seta always present on each side of the median
line.

Mesothorax : Prescutum 49-58 (55) [x long and 70-89 (85)0. wide, the ratio being 1 : 1-4-1-7
(i-6) ; prescutal ridge well developed and prescntal suture distinct. Scutum 46-52 (49) u. long,
i.e. the ratio lengths of prescutum to scutum 1 -0-1-3 (i-i) : 1. Prealare, prealar ridge, tegula,
anterior and posterior notal wing processes well developed. Scutellum 31-34 (33) |x long and
73-82 (79) [x wide, i.e. the ratio 1 : 2-3-2-7 (2-4), and its length to the length of the scutum
1 : 1-4-1-6 (1-5) ; scutellar ridge distinct. Postalare with the anterior and posterior postalar
ridges joined. Mesopleuron : Mesopleural ridge continuous above the coxal articulation ;
mesopleural apophysis, mesopleural wing process and subepisternal ridge well developed ; basalare
slender. Mesepisternum and mesepimeron w-ell defined ; lateropleurite comparatively large.
Mesosternum : Basisternum 85-98 (92) [x long and 134-159 (150) jx wide ; bounded by the
marginal and the precoxal ridges. Furca well developed. Mesothoracic spiracle 12-15 (14)1/
wide at opening, with a 27-31 (29) jx long supporting bar. Mesothoracic setae on each side :
Prescutal setae 1-2 (1-5) ; scutal setae 2-3 (2-2) ; scutellar seta always one. Tegular setae 2-3
(2-6). Each lateropleurite always with a small later opleurital seta. Postmesostigmatal and
basisternal setae absent. Metathorax : Metapostnotal sclerite small ; metapostnotal ridge
absent. Metapleural ridge attenuated towards the small metapleural apophysis. Metepisternum
and metepimeron distinct. Precoxal ridge and metasternal apophysis well developed. Meta-
thoracic spiracles similar to the mesothoracic. Metathoracic setae on each side : Metatergal
setae 1-2 (1-7) ; metapleural setae absent ; one anterior metasternal seta always present, and
posterior metasternal setae absent.

Wings : 714-798 (756) jx long and 266-308 (280) (x wide ; with one alar seta. Hamulohalterae
49-58 (52) |x long and 15-18 (17) (x wide ; with one apically hooked, 49-67 (61) jx long seta, i.e.
the ratio lengths of seta to hamulohaltera 1 : 0-9-1-1 (10).

Legs : Comparatively short and stout ; the fore and middle legs of subequal length and
shorter than the hind leg ; the ratio length of the hind leg to the total body length 1 : 21— 2-4
(2-3). Coxa and trochanter about 37 and 21 [x wide respectively. Femur about 31 [x wide ;
that of the middle leg shortest, and those of the fore and hind legs subequal ; the ratio width
to length of the hind femur 1 : 3-4-3-8 (3-6). Tibia about 18 a. wide ; with 2 apical spurs and
2-3 smaller spines ; in the fore leg the femur shorter than the tibia, the ratio their lengths
being 1 : 1-1-1-3 (1-2). Tarsus about i5jx wide ; tarsal digitules apically knobbed, about
27(x long. Claw broad at base and abruptly tapering towards a pointed tip ; ungual digitules

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

173

dmep scp

dmcr dmep

Fig. 41. Eriococcas orariensis Hoy, dorsal and ventral view.

174

MORPHOLOGY AND TAXONOMY OF ADULT MALES

apically knobbed. The lengths of the leg segments (in microns) and the number of setae on
each are given in the following table :

Fore leg

Middle leg

Hind leg

Coxa

length

f.s.

h.s.

27-31 (29)

i-3 (2-1)
6-10 (7-5)

27-31 (29)
i-5 (2-9)
5-8 (6-4)

31-34 (32)

i-5 (3-i)
5-8 (6-2)

Trochanter

length

f.s.

h.s.

40-46 (43)
0-1 (o-i)
3-6 (3-9)

43-49 (4 6 )
0-1 (o-i)

3-6 (4-3)

46-49 (48)
0-1 (0-2)
4-5 (4-5)

Femur

length

f.s.

h.s.

104-110 (107)

6-10 (7-8)

13-19 (15-8)

92-104 (98)

5-10 (8-i)

11-15 (13-5)

104-116 (no)

4-9 (6-2)

10-14 ( II- 8)

Tibia

length

f.s.

h.s.

116-137 (128)

6-9 (7-9)
15-22 (18-8)

116-137 (128)

7-10 (8-2)

14-24 (20-4)

137-153 (143)

7-13 (9-i)

18-25 (2i'3)

Tarsus

length

f.s.

h.s.

49-52 (50)
3-5 (3-5)
10-13 (n-6)

49-52 (50J
2-5 (3-4)
10-14 ( I2-1 )

49-55 (52)
2-5 (3-5)
10-14 ( IX '8)

Claw

length

12-15 (14)

12-15 (14)

12-15 (14)

Total length

of leg

348-384 (366)

342-384 (363)

384-421 (394)

Abdomen : 228-319 (28i)(i long and 205-220 (213)^1. wide. Tergites of segments I and II
small ; that of segment VIII large. Sternites of segment VIII bounded along their outer
margins by a longitudinal heavily sclerotized strip. Ostioles absent. Abdominal setae on
each side : Dorsal setae usually 2 on segments I to VII (sometimes 3 on segments IV and VI),
and one slightly longer seta on segment VIII. Pleural setae 2 on segment I, 2-3 on segment II,
and 3 on segments III to VIII, including a longer one on the latter segment. Ventral setae
1 on segment II, 2-3 on segment III, 4-5 on segments IV to VII, and 1 on segment VIII.

Glandular pouches small ; setae of glandular pouch include a pair of 119-137 (128)^ long tail
setae, i.e. the ratio their length to the total body length 1 : 6-0-7-9 (6-9).

100.

100 ^

Fig. 42. Eriococcus orariensis Hoy, lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 175

Genital segment : Genital capsule elongate, 119-134 (128)51 long and 43-49 (46) ;x wide, the
ratio being 2-6-3-0 (2-8) : 1, and the ratio its length to the total length of body 1 : 6-2-7-8 (6-9).
Style rather straight in lateral view, with a pointed apex. Anal opening about i2y. in diameter.
Aedeagus with a well sclerotized ventral margin and a sharply pointed tip. Setae of genital
segment : Dorsally 2 comparatively long setae always present on each side of the style at about
half its length. Ventrally 4-5 (4-6) similar setae occur on each side of the genital capsule.

Material : 10 specimens from two lots examined, all of which entirely agreed
with each other. The specimens of one lot were collected by J. M. Kelsey, on
Leptospermum scoparium, in Ashburton, New Zealand, 10. vi. 1948 ; the other
specimens were collected by J. M. Hoy, on the same host plant, in Palmerston
North, New Zealand, 21.iii.1956.

Eriococcus buxi (Fonscolombe)
(Text-figs. 43, 44)

Macropterous males only known ; living specimens not available. The males moderately
robust, comparatively small, with short antennae and moderately long legs. When mounted,
942-1050 (980) [i long, 266-294 (280) u. wide at mesothorax and 1862-2100 (2002) u. wing
expanse.

Body setae : Antennae and legs with about i8(ji long fleshy setae and somewhat longer hair-
like ones ; both types of setae hard to separate. Body itself with hair-like setae only.

Head : Subtetrahedron ; subtriangular in dorsal and front views ; ventral preocular
depression hardly indicated in lateral view. Length from apex to postoccipital ridge 67-76
(70)51 ; from apex to neck 140-150 (144)1,1 ; width across the genae 174-183 (180)51. Dorsal
arm of midcranial ridge short, joining lateral and ventral arms at the apex of the head. Post-
occipital ridge well developed, laterally giving off two branches, one anteriorly and the other
posteriorly. Dorsomedial part of epicranium slightly raised. Preocular and postocular ridges
well separated ; ventral part of the former atrophied. Preoral ridge slender. Eyes : Dorsal
simple eyes not projecting beyond the outer margins of the head ; their corneae 18-24 ( 2I )( Jl
in diameter and separated by 58-61 (59)51, i.e. 2-4-3-3 ( 2 "7) times their diameter apart. Ventral
simple eyes 21-24 ( 22 )i JL an ^ 18-21 (20)51 apart. Lateral ocelli closely attached to postocular
ridges. Ocular sclerites large. Cranial apophysis apically truncate. Tentorial bridge stout.
Dorsal head setae 11-15 (12-8) on each side ; each gena with 9-15 (n-8) genal setae ; setae of
ocular sclerites 10-13 ( II- 6)- Ventral head setae on each side : 2-3 (2-6) between the ventral
eyes ; 11-15 (13-2) in the area of the ventral preocular depression, forming on both sides a
transverse band ; 4-5 (4-2) on each side of the ventral arm of the midcranial ridge.

Antennae : Rather filiform ; 9-segmented ; 458-506 (476)51 long, i.e. about as long as half
the total body length, the ratio being 1 : 2-0-2-1 (2-05), and slightly shorter than the hind legs,
the ratio 1 : 1-1-1-3 ( I-2 )- Scape 37-40 (39)51 long and 34-37 (36)51 wide at base ; always
with 4 h.s. Pedicel 43-49 (46) ji long and 31-37 (34) 51 wide ; with 2-5 (3-5) f.s., 10-16 (13-3)
h.s. and a sensillum placodeum. Flagellum : Flagellar segments 21-2451, wide, with segment
III somewhat wider and segment IX somewhat narrower. Segment III club-shaped and
longest of all, the ratio its length to the length of terminal segment 2-6-2-9 (2-8) : 1, and the
ratio its width to its length 1 : 3-2-3-6 (3-5). Segments IV to VIII rather barrel-shaped, the
ratio width to length of segment VIII being 1 : 1-5-1-9 (i-6) ; terminal segment pear-shaped,
shortest of all and always with 5-6 apically knobbed capitate sensory setae. The following table
shows the lengths of the flagellar segments and the number of setae on each :

176

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Ill

IV

V

VI

VII

VIII

IX

length in ja

79-89

73-82

49-64

61-67

37-43

37-43

31

(av.)

(85)

(76)

(58)

(64)

(40)

(40)

(3i)

f.s.

17-26

18-26

17-26

22-28

11-20

10-16

(av.)

(22-1)

(22-5)

(20-5)

(24-5)

(16-3)

(13-3)

h.s.

5-9

5-8

3-5

4-8

3-6

4-6

(av.)

(7-5)

(6-5)

(3-8)

(5-8)

(4-5)

(4-8)

set. ca.

5-6

(av.)

(5-8)

,. scWrctiza'"" 1

Fig. 43. Eriococcus buxi (Fonscolombe) , dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

*77

Antennal bristles well defined ; segments VII and VIII each with a ventral bristle. Segment
IX with three bristles near its base and one apical hair-like seta.

Thorax : 380-418 (403) ;x long. Prothorax : Pronotal ridges medially interrupted by a weak
sclerotization. Lateral pronotal sclerites and post-tergites small. Proepisternum without any
ridge-like sclerotization. Prosternum triangular, 24-34 (27)10. long ; prosternal ridge 92-98
(95) [x long. Prothoracic setae : Medial, lateral pronotal, posl-tergital and antespiracular dorsal
setae absent. One antespiracular ventral seta always present and one prosternal seta usually
present (av. 08) on each side.

Mesothorax : Presculum 67-82 (76) jx long and 1 10-122 (i 16) jj. wide, the ratio being 1 : 15-
1-7 (i-6) ; laterally bounded by the prescutal ridges and posteriorly by the prescutal suture.
Scutum 43-52 (46) u. long, i.e. the ratio lengths of prescutum to scutum 1-6-1-9 ( r '7) : *■
Prealare, prealar ridge, tegula, anterior and posterior notal wing processes well developed.
Scutellum 37-46 (40)1* long and 107-122 (t 16) jj. wide, i.e. the ratio 1 : 2-7-3-0 (2-9), and its
length to the length of the scutum 1 : 1-1-1-2 (1-15) ; scutellar ridge distinct. Postalare with
the anterior and posterior ridges joined. Postnotal apophysis strong. Mesopleuron : Meso-
pleural ridge continuous above the coxal articulation ; basalare slender. Subepisternal ridge
well developed. M esepisternum distinct and mesepimeron small. Lateropleurite comparatively
large, with an oval membranous area. Mesosternum : Basisternum 92-110 (ioi)u. long and
174-189 (180) (i wide ; antero-laterally and latero-posteriorly bounded by the marginal and
precoxal ridges respectively. Furca well developed. Mesolhoracic spiracle 15-18 (17)0. wide
at opening, with a 27-31 (29) (i. long supporting bar. Mesothoracic setae on each side : Prescutal
setae 6-9 (76) ; scutal setae 5-7 (6-i) ; scutellar setae 2-3 (28). Tegular setae 7-9 (79).
Anepisternum with 2-3 (2-8) mesepisternal setae. Postmesostigmatal setae in two groups, 3-4
(3-4) latero-ventrally and 2-3 (2-6) medially ; basisternal setae 10-15 (n-6). Metathorax :
Metapostnotal sclerites small ; metapostnotal ridge absent. Metapleural ridge attenuated as
usual towards the small pleural apophysis. Metepisternum and metepinurtni distinct. Precoxal
ridge weak and metasternal apophysis small Metathoracic spiracle identical with the meso-
thoracic. Metathoracic setae on each side : Metatergal setae 4-5 (4-2) ; metapleural setae 1-2
(i-6) ; postmetastigmatal setae 2-3 (2-6). Anterior metasternal setae 2-3 (2-4) and posterior
metasternal setae usually absent, although one seta was found in one specimen.

Wings : 812-924 (868) \i long and 294-350 (322)0. wide ; alar setae almost always absent
and only one seta may occur (av. o-i). Hamulohalterae 58-67 (64)^ long and 15-18 (i6)(x wide ;
with one apically hooked, 52-67 (58)^ long seta, i.e. the ratio lengths of seta to hamulohaltera
1 : i-o-i-i (108).

ads_

^ aps ^

.— *.

200-*-

100 ^

Fig. 44. Eriococcus buxi (Fonscolombe), lateral view.

178 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Legs : Moderately long and stout ; the fore and the middle legs of subequal length and
shorter than the hind legs ; the ratio length of the hind leg to the total length of the body
1 : 1 -8-i -9 (1-85). Coxa and trochanter about 40 and 24(1. wide respectively. Femur about
31 (x wide ; that of the middle leg shortest and that of the hind leg longest ; the ratio width to
length of the hind femur 1 : 4-9-5-2 (50). Tibia about i8jj. wide ; with 2 apical spurs and
2-3 smaller spines ; in the fore leg the femur slightly shorter than the tibia, the ratio their
lengths 1 : 1-1-1-2 (1-17). Tarsus about i8(i wide ; tarsal digitules apically knobbed, about
29 (i. long. Claw broad at base and abruptly tapering to a sharply pointed tip ; ungual
digitules apically knobbed. The following table shows the lengths of the leg segments (in
microns) and the number of setae on each :

Fore leg Middle leg Hind leg

length 34-40 (37) 34-40 (37) 37-43 (40)

Coxa f.s. 3-6 (4-2) 3-5 ( 4 -6) 4-6 (5-1)

h.s. 4-6 (5-2) 4-7 (5-5) 5-8 (6-6)

length 52-55 (54) 52-55 (54) 55-61 (58)

Trochanter f.s. 1-3 (i-8) 1-3 (i-6) 1-4 (2-2)

h.s. 2-4 (2-6) 2-5 (2-9) 2-5 (3-2)

length 134-150 (143) 122-134 (128) H3-I59 (i53)

Femur f.s. 7-12 (9-5) 6-10 (8-i) 7-13 (io-8)

h.s. 13-19 (15-7) 10-17 (13-3) 12-21 (16-5)

length 162-177 ( l68 ) 174-186 (180) 186-195 U92)

Tibia f.s. 17-25 (217) 20-26 (22-4) 22-30 (27-1)

h.s. 19-34 (26-6) 23-39 (31-8) 27-41 (35-4)

length 70-73 (72) 70-73 (72) 73-76 (75)

Tarsus f.s. n-15 (12-6) 11-17 (13-1) 13-18 (16-2)

h.s. 13-18 (15-5) 12-19 (15-9) 16-22 (18-8)

Claw length 12-15 (*4) 12-15 (*4) 12-15 ( J 4)

Total length of leg 467-509 (49i) 467-506 (488) 5Q9-549 (528)

Abdomen : 304-380 (357) ^ long and 251-289 (266)}! wide. Tergites of segments I and II
small ; that of segment VIII large, anteriorly bounded by a transverse stripe of heavy
sclerotization. Sternites of segment VIII also bounded laterally by a longitudinal, similarly
heavy sclerotization. Ostioles absent. Abdominal setae on each side : Dorsal setae 4-6 on
segment I, 4-5 on segments II to V, 2-4 on segments VI and VII, and one slightly longer seta
on segment VIII. Pleural setae 5-7 on segments I to VII and 2-3 on segment VIII, including
one slightly longer seta. Ventral setae 1-2 on segment II, 2-3 on segments III to VII, and
always 2 on segment VIII.

Glandular pouches small ; setae of glandular pouch include a pair of 107-122 (ii3)[x long tail
setae, i.e. the ratio their length to the total body length 1 : 7-7-9-8 (8-7).

Genital segment : Genital capsule comparatively large, 140-153 (146)^ long and 82-92 (85) [x
wide, the ratio being 1-6-1-9 (1-7) : 1, and the ratio its length to the total length of the body
1 : 6-6-6-9 (6-7). Style rather straight in lateral view, with a pointed apex. Anal opening
about i8jx in diameter. Aedeagus with the ventral margin well sclerotized ; internal genital
aperture large. Setae of genital segment : Dor sally 2 long setae always present on each side of
the style near its apex ; ventrally 3 smaller setae occur on each side of the genital capsule.

Material : 7 specimens examined, collected by N. S. Borchsensius, on Buxus
sempervirens, in Abhasia, U.S.S.R., in 1932 (no exact date given).

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 179

OVATICOCCUS Kloet, 1944

Ovaticoccus agavium Douglas

(Text-figs. 45, 46)

Winged forms only known ; these dark reddish brown, with the thorax slightly lighter.
The males moderately robust, comparatively small, with moderately short antennae and
moderately long legs. Mounted specimens 1050-1204 (ii2o)u. long, 308-350 (336) u. wide at
mesothorax and 2128-2422 (2296) jo. wing expanse.

Body setae : Antennae with about i2u. long fleshy setae and about twice as long hair-like
ones ; body and legs with hair-like setae only, much smaller than those of the antennae.

Head : Subtetrahedron ; subtriangular in dorsal view and more or less rounded in front
view ; ventral preocular depression hardly indicated in lateral view. Length from apex to
postoccipital ridge 107-125 (n6)u. ; from apex to neck 159-177 {i6j)[l ; width across the
genae 198-207 (201) \x. Dorsal arm of midcranial ridge anteriorly continuous with the ventral
arm and posteriorly fading out before reaching the postoccipital ridge ; lateral arms indicated
by short, weak sclerotizations. Postoccipital ridge laterally giving off two small branches,
one anteriorly and the other slightly bent posteriorly. Preocular and postocular ridges well
separated ; preoral ridge slender. Eyes : Dorsal simple eyes not projecting beyond the outer
margins of the head in dorsal view ; their corneae 15-18 (17)10. in diameter and both separated
by 98-113 (107)0., i.e. 5-3-7-4 (60) times their diameter apart. Ventral simple eyes 15-21
(i8)|x in diameter and 43-55 (49) [x apart. Lateral ocellus vestigial and only represented by a
small sclerotized spot on the ocular sclerite, attached to the postocular ridge by means of a short
sclerotized arm. Cranial apophysis apically bifurcate. Tentorial bridge stout. Mouth
opening posteriorly situated on a small mouth tubercle. Dorsal head setae 6-9 (7-1) on each
side ; each gena with 2-3 (2-1) genal setae. Ventral head setae on each side : Setae between
the ventral eyes absent ; 2-4 (3-1) in the area of the ventral preocular depression, forming
on both sides transverse single or double rows ; anteriorly 1-2 (i-8) on each side of the ventral
arm of the midcranial ridge.

Antennae : Rather filiform ; 10-segmented ; 601-677 (637)1* l° n g. i- e - as l° n g as or slightly
longer than half the body length, the ratio being 1 : 1-6-2-0 (i-8), and about as long as the
hind legs, the ratio 1 : 10-11 (1-07). Scape 43-49 (46)10. long and 49-55 (52)10. wide at base ;
always with 4 h.s. Pedicel 58-64 (6i)u. long and 37-43 (40)10. wide ; with 7-11 (8-4) h.s. and a
sensillum placodenm. Flagellum : Flagellar segment 27-31 (29) [o. wide, with segments VIII,
IX and sometimes VII inconsiderably wider. Segment III club-shaped and longest of all,
the ratio its length to the length of the terminal segment 1-7-2-3 (1-9) : 1, and its width to its
length 1 : 3-4-4-2 (4-0). Segments IV to IX rather barrel-shape ; the ratio width to length
of segment IX 1 : 1-4-1-8 (i-6). Terminal segment pear-shaped, with a narrowly constricted
or even pointed apex. Segments VI to X and sometimes V also, with a number of apically
knobbed, capitate sensory setae. In the following table, the lengths of the flagellar segments
and the number of setae on each are given :

III IV V VI VII VIII IX X

length in (o. 101-116 61-76 61-76 61-73 52-64 49-61 46-61 46-64

(av.) (no) (67) (66) (66) (58) (55) (55) (58)

f.s. 2-7 4-7 5-10 4-12 6-1 1 5-1 1 3-6 2-5

(av.) (4-2) (5-6) (6-6) (9-2) (8-4) (7-4) (4-3) (3-7)

h.s. 7 -I 4 4 - 8 5-8 2-4 0-3 0-2 0-2 1-2

(av.) (n-2) (6-o) (6-i) (3-2) (1-3) (i-o) (i-i) (2-0)

Set. ca. o o 0-1 2-4 3-4 3-5 3-6 3-5

(av.) o o (o- 4 ) (3-1) (3-6) (4-3) (4-2) (3-6)

i8o

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Antennal bristles well defined ; segments VIII and IX each with one ventral bristle. Segment
X with 3 bristles, the dorsal of which slightly shorter and nearer to the apex ; the segment
also with one apical hair-like seta.

Thorax : 388-494 (448)^1 long. Prothorax : Pronotal ridges medially interrupted at a weakly
sclerotized point. Lateral pronotal sclerites and post-tergites small. Proepisternum without
any ridge-like sclerotization. Prosternum triangular, 34-46 (40) ja long ; prosternal ridge

spl Pdc

dhs dmep

vmcr
dmcr / Sep

Fig. 45. Ovaticoccus agavium Douglas, dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

181

92-107 (95) y. long. Prothoracic setae : Medial, lateral pronotal, post-tergital and antespiracular
dorsal setae absent. One antespiracular ventral seta always present on each side ; prosternal
setae absent.

Mesothorax : Prescutum 58-70 (64)^ long and 122-140 (131)^ wide, the ratio being 1 : 19-
2-3 (2-0) ; laterally and posteriorly bounded by the prescutal ridge and the prescutal suture
respectively. Scutum 49-61 (52) ^ long, i.e. the ratio lengths of prescutum and scutum 1 -1-1-4
(1-2) : i. Prealare, prealar ridge, triangular plate and the mesopleural structures well developed.
Sculellum 61-70 (67)^ long and 113-131 (122)^ wide, i.e. the ratio 1 : 1-6-2-0 (i-8), and its
length to the length of the scutum 1-1-1-4 ( I- 3) : * '• scutellar ridge strongly developed.
Anterior and posterior postalar ridges anteriorly joined. Postnotal apophysis well developed.
Mesopleuron : Mesopleural ridge continuous above the coxal articulation ; basalare slender.
Subepisternal ridge distinct. Mesepisternum and mesepimeron well defined. Latero-pleurite
comparatively large. Mesosternum : Basisternum 116-134 (i25)[x long and 192-211 (2oi)(i.
wide ; marginal, precoxal ridges and /urea well developed. Mesothoracic spiracle 21-24 [ 22 )v>-
wide at opening, with a 43-49 (46) (x long supporting bar. Mesothoracic setae on each side :
Prescutal setae 0-1 (0-4) ; scutal setae usually 1-2 and occasionally 3 (av. i-6) ; scutellar setae
1-2 (1-9). Tegular setae 2-3 (i-8). Postmesostigmatal, and basisternal setae absent. Meta-
thorax : Metapostnotal sclerites small ; metapostnotal ridge absent. Melapleural ridge
attenuating near the middle as usual ; melapleural wing process missing. Metepisternum and
metepimeron small. Precoxal ridge weak and metasternal apophysis small. Metathoracic
spiracle similar to the mesothoracic. Metathoracic setae on each side : Metatergal setae 3-4
(3-4) ; melapleural setae absent. Anterior metasternal seta 0-1 (o-6) ; posterior metasternal
setae absent.

Wings : 910-1050 (994) \i long and 308-392 (350) (i. wide ; always with 2 alar setae. The
hamulohalterae and the suspensorial sclerites absent.

Legs : Moderately long and stout ; the fore and the middle legs of subequal length and
shorter than the hind legs ; the ratio length of hind leg to the total length of body 1 : 1-5-1-8
(i-6). Coxa about 55 and trochanter about 3i[i wide respectively ; the latter with one, about
46(x long differentiated apical seta. Femur about 401A wide ; that of the middle leg shortest
and those of the fore and hind legs subequal ; the ratio width to length of the hind femur
1 : 4-3-4-7 (4-5). Tibia about 24^1 wide ; with 2 apical spurs ; in fore leg the femur slightly
shorter than the tibia, the ratio their lengths 1 : 1-2-1-4 ( x '3)- Tarsus about i8(j. wide ; tarsal

200 ^*-

Fig. 46. Ovaticoccus agavium Douglas, lateral view.

Coxa

length
h.s.

Trochanter

length
h.s.

Femur

length
h.s.

Tibia

length
h.s.

Tarsus

length
h.s.

Claw

length

Total length of leg

182 MORPHOLOGY AND TAXONOMY OF ADULT MALES

digitules apically knobbed, 31-37 (34) (J. long. Claw broad at base, abruptly tapering towards
a sharply pointed tip ; ungual digitules apically knobbed. In the following table the lengths
of the leg segments (in microns) and the number of setae on each are given :

Fore leg Middle leg Hind leg

37-43 (39) 37-43 (39) 40-46 (42)

6-10 (7-6) 6-11 (8-i) 7-11 (9-2)

61-70 (64) 64-70 (67) 70-76 (73)

2-4 (2-8) 2-4 (3-0) 2-5 (3-4)

174-189 (183) I53-I7 1 ( l6 5) 174-186 (180)
11-18 (14-6) 10-16 (12-8) 11-16 (136)

226-253 (238) 232-259 (244) 250-293 (272)
29-38 (33-2) 30-42 (36-1) 32-46 (38-4)
82-92 (85) 85-95 (89) 89-101 (95)

17-23 (19-2) 16-23 (19-6) 18-25 (21-2)

18-24 (2l) 18-24 (2l) 18-24 (2l)

613-659 (631) 607-656 (625) 659-720 (683)

Abdomen : 342-471 (4io)u. long and 296-334 (3i9)(x wide. Tergites of segment I small ;
that of segment VIII large. Sternites of segment VIII ill-defined. Ostioles absent.
Abdominal setae on each side : Dorsal setae 3-4 on segments I to V, 2-3 on segments VI and
VII, and one slightly longer seta on segment VIII. Pleural setae 2-3 on segment I, 3-4 on
segments II to VII, and 3 on segment VIII, including a slightly longer seta. Ventral setae
1-2 on segments II and III, 2—3 on segments IV to VII, and 2 on segment VIII.

Setae of glandular pouch include a pair of 1 16-140 (128) u. long tail setae, the ratio their length
to the total body length being 1 : 7-7-10-3 (8-8).

Genital segment : Genital capsule moderately large, 128-143 (134) \i long and 85-95 (89) ji.
wide, i.e. the ratio 1-4-1-6 (1-5) : 1, and the ratio its length to the total length of body 1 : 7-8-
8-9 (8-4). Style rather straight in lateral view, apically pointed. Anal opening about i2(x in
diameter. Aedeagus tubular, with a short, strongly sclerotized basal rod. Setae of genital
segment : Dorsally 2 long setae present on each side of the style near its base ; ventrally, 4-6
(5-1) smaller setae occur on each side of the genital capsule.

Material : 10 specimens examined, obtained from the following two lots, both
of which completely agreed with each other : (A) 5 specimens collected by
K. Boratynski, on Agave decipiens, in Kew Gardens, England, 27. vi. 1957. (B) 5
specimens collected by myself, on Agave franzosinii, also in Kew Gardens, England,
during the last two weeks of June, 1964.

This species was also recorded by Dr. Boratynski on Agave parryi.

GOSSYPARIA Signoret, 1875
Gossyparia spuria (Mod.)
(Text-figs. 47-49)
Macropterous and brachypterous forms of this species were available for study.
(A) The macropterous form (Text-figs. 47 and 48)

The males moderately robust, comparatively large, with short antennae and short legs.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

183

When mounted 1442-1512 (1484)0 long, 392-420 (406)11 wide and 2520-2758 (2632)0, wing
expanse.

Body setae : Antennae with numerous fleshy and few hair-like setae, about 370 long ; body
and legs with much shorter hair-like setae only.

Head : Subtetrahedron ; subtriangular in dorsal view and more or less rounded in front
view ; ventral preocnlar depression absent in lateral view. Length from apex to postoccipital
ridge 122-131 (127)0 ; from apex to neck 189-198 (194)0 ; width across the genae 238-250

x vhs

Fig. 47. Gossyparia spuria (Mod.), macropterous form, dorsal and ventral view.

184

MORPHOLOGY AND TAXONOMY OF ADULT MALES

(244) [i. Midcranial ridge well developed and cruciform ; dorsal arm short, joining the lateral
and the ventral arms at the apex of the head. Postoccipital ridge laterally bending posteriorly
and giving off short, heavily sclerotized extensions anteriorly. Pre- and postocular ridges well
separated ; preoral ridge slender. Eyes : Dorsal simple eyes slightly projecting beyond the
outer margins of the head in dorsal view ; their corneae 24-27 (26) \l in diameter and separated
by 128-137 (134) (x, i.e. 5-0-5-6 (5-3) times their diameter apart. Ventral simple eyes equal to
the dorsal in diameter and 37-40 (39) u. apart. Lateral ocelli large, closely attached to post-
ocular ridge. Cranial apophysis apically bifurcate. Anterior tentorial arms join the cranial
apophysis separately ; tentorial bridge stout. Dorsal head setae 10-14 (11 -8) on each side ;
each gena with 4-6 (5-0) genal setae. Ventral head setae : Setae between the ventral eyes
absent ; 3-5 (4-1) just anterior to the preocular ridges, forming on both sides a transverse
irregular row ; 2-3 (2-5) on each side of the ventral arm of the midcranial ridge.

One and sometimes two (av. 1-2) small bodies of unknown homology (X) occur dorsally on
each side of the dorsal arm of the midcranial ridge, near the apex of the head.

Antennae : Rather moniliform ; 10-segmented ; 625-677 (653)^ long, i.e. somewhat
shorter than half the length of the body, the ratio being 1 : 2-2-2-4 ( 2- 3). ar *d equal or slightly
shorter than the hind legs, the ratio 1 : 1-0-1-2 (it). Scape 52-55 (54)^ long and 55-58 (57)[i
wide at base ; with 4-5 (4-5) h.s. Pedicel 67-70 (69) (j. long and 49^ wide ; with 3-5 (4-0) f.s.,
2— 4 (3-0) h.s. and a sensillum placodenm. Flagellum : Flagellar segments IV to VII 31-37 (34) \x
wide ; segments III and X slightly narrower and segments VIII and IX slightly wider. Segment
III club-shaped and longest of all, the ratio its length to the length of segment X being 1-7-
1-9 (i-8) : 1, and its width to its length 1 : 3-1-3-6 (3-4). Segments IV to IX barrel-shaped,
the ratio width to length of segment IX 1 : 1-2-1-5 (1-4). Terminal segment pear-shaped.
Segments V to X and sometimes IV also, with a few apically knobbed, capitate sensory setae.
The following table shows the lengths of the flagellar segments and the number of setae on each :

III

IV

V

VI

VII

VIII

IX

X

length in y.

95-98

82-85

61-67

61-70

58-64

49-55

49-55

52-58

(av.)

(97)

(84)

(64)

(66)

(61)

(52)

(52)

(55)

f.s.

4 -8

I5-J9

7-10

8-12

10-13

10-14

9-12

8-1 1

(av.)

(6-o)

(17-0)

(8-5)

(io-o)

("•5)

(12-0)

(io-5)

(9-5)

h.s.

3-4

3-5

2-3

i-3

1-2

1-2

1-2

1-2

(av.)

(3-5)

(4-o)

(2-5)

(2-0)

(i-5)

(i"5)

(i-5)

(i-5)

Set. ca.

O-I

3-4

2-4

3-4

2-4

3-4

3-4

(av.)

(o-5)

(3-5)

(3-o)

(3-5)

(3-o)

(3-5)

(3-5)

200-^

Fig. 48. Gossyparia spuria (Mod.), macropterous form, lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 185

Antennal bristles subequal to the fleshy setae in length ; segments VIII and IX each with
one ventral bristle. Terminal segment with one dorsal preapical bristle, two lateral ones
at a greater distance from the apex and two apical small setae.

Thorax : 585-623 (608) |x long. Pronotal ridges medially interrupted at a weakly sclerotized
point ; lateral pronotal sclerites small and post-tergites narrow. Proepisternum without any
ridge-like sclerotization. Prosternum comparatively long and narrow, 82-85 (84) fx long ;
posteriorly bounded by a 119-125 (122) y. long prosternal ridge. Prothoracic setae absent
altogether.

Mesothorax : Prescntum 89—98 (95)(x long and 171-174 (i73)[x wide, the ratio being 1 : i-8-
20 (1-9) ; prescutal ridge well developed and presculal suture distinct. Scutum 91-95 (94) (x
long, i.e. the ratio lengths of prescutum to scutum 1 : 0-9-1-1 (10). Prealare, prealar ridge,
tegular, triangular plate, anterior and posterior notal wing processes well developed. Scutellum
70-73 (72)1* long and 159-168 (165)11 wide, the ratio being 1 : 2-2-2-4 ( 2 "3). an d its length to
the length of scutum 1 : 1-2-1-4 ( I- 3) '• scutellar ridge well developed. Postalare with anterior
and posterior ridges joined. Postnotal apophysis large. Mesopleuron : Mesopleural ridge
continuous above the coxal articulation. Basalare comparatively slender. Subepisternal
ridge well developed. Mesepisternum distinct and mesepimeron small. Later o-pleurile large,
with an oval membranous area. Mesosternum : Basisternum 183-195 (189) jx long and 241-
250 (247) (x wide ; marginal, precoxal ridges and furca well developed. Mesothoracic spiracle
21-24 ( 2 3)|x wide at opening, with a 46-52 (49) \x long supporting bar. Mesothoracic setae on
each side : One prescutal setae always present ; scutal as well as scutellar setae 2-3 (25).
Tegular setae always 2. Postmesostigmatal and basisternal setae absent. Metathorax : Meta-
postnotal sclerites ill-defined ; metapostnotal ridge absent. Metapleural ridge attenuated as
usual near the middle ; dorsally supporting a small pleural wing process. Precoxal ridge
strong and metasternal apophysis distinct. Metathoracic spiracle identical with the meso-
thoracic. Metathoracic setae : 4-5 (4-5) metatergal setae and one anterior metasternal seta
present on each side ; metapleural and posterior metasternal setae absent.

Wings : 1 064-1 148 (no6)(x long and 420-490 (462) jx wide ; always with 2 alar setae.
Hamulohalterae 92 \x long and 24-27 (26) |x wide ; with one, 52-58 (55) jx long apically hooked
seta, the ratio its length to the length of the hamulohaltera being 1 : 1 -6-1-8 (17).

Legs : Well developed, short and stout ; the middle legs usually shortest and the hind legs
longest ; the ratio length of the hind leg to the total length of body 1 : 2-1-2-2 (2-15). Coxa
and trochanter about 64 and 371X wide respectively ; the latter with a differentiated long apical
seta. Femur about 46(x wide ; that of the middle leg shortest and those of the fore and hind
legs subequal ; the ratio width to length of the hind femur 1 : 3-5-40 (3-8). Tibia about 24(x
wide ; with 2 apical spurs ; in fore leg the femur slightly shorter than the tibia, the ratio their
lengths being 1 : 1-2-1-4 ( I- 3)- Tarsus about 2i(x wide ; tarsal digitules apically knobbed,
about 37 (x long. Claw broad at base, abruptly tapering apically ; ungual digitules also apically
knobbed. The following table shows the lengths of the leg segments (in microns) and the
number of setae on each :

Fore leg Middle leg Hind leg

Coxa length 55-61 (58) 55-61 (58) 58-64 (61)

h.s. 8-12 (9-4) 8-1 1 (9-1) 10-14 ( II- 6)

Trochanter length 73"70 (74) 73-7° (74) 76-79 (77)

h.s. 4-6 (5-i) 4-5 (4-6) 4-° (5"3)

Femur length 159-186 (174) 146-165 (156) 162-183 ( J 74)

h.s. 15-22 (18-5) 14-20 (168) 14-18 (15-9)

Tibia length 214-238 (229) 217-244 (232) 226-259 (247)

h.s. 18-29 (24-4) 20-34 (27-6) 22-35 (28-8)

[86 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Tarsus length

67-76 (73)

67-79 (74)

70-82 (76)

h.s.

10-12 (111)

10-13 ("-8)

11-15 (133)

Claw length

21-24 (23)

21-24 (23)

21-24 (23)

Total length of leg

589-659 (628)

580-647 (613)

613-689 (656)

Abdomen : 540-578 (560)0. long and 380-410 (395)0. wide. Tergites of segments I, II and
sometimes III weak and narrow ; that of segment VIII large. Sternites of segment VIII
ill-defined. Ostioles absent. Abdominal setae on each side : Dorsal setae 4—6 on segments I
to IV, 3-4 on segment V, 3 on segments VI and VII, and 2-3 on segment VIII, including a
slightly longer seta. Pleural setae 3-4 on segments I and II, 4-5 on segments III to VII, and
4 on segment VIII, including a considerably longer seta (about 90 [jl long). Ventral setae 1
on segment II, 2-3 on segments III to VII, and 1 on segment VIII.

Setae of glandular pouch only include a pair of 168-177 (173)0. long tail setae, the ratio their
length to the total length of body being 1 : 8-1-9-0 (8-6).

Genital segment : Genital capsule moderately large, 177-183 (180) o. long and no[z wide, i.e.
the ratio 1-6-1-7 ( I- 65) : i> and the ratio its length to the total length of body 1 : 8-1-8-3 (8 - 2).
Style undulating and apically pointed in alteral view. Anal opening about 24(1 in diameter.
Aedeagus with well sclerotized lateral margins and tapering towards a sharply pointed tip.
Setae of genital segment on each side : Dorsally two conspicuously long setae always present
near the base of the style ; ventrally 7-9 (8-o) slightly smaller setae occur on the genital capsule.

Material : 3 specimens only were available, collected by A. Dziedzicka, on
Ulmus sp., in Mogila, nr. Nowa Huta, nr. Krakow, Poland, 26. iv. 1963.

(B) The brachypterous form (Text-fig. 49)

These males rather robust, with the abdomen slightly broader than the thorax ; compara-
tively large, with short antennae and short legs. Mounted specimens 1274-1470 (1358)11 long,
336-378 (350)11 wide at mesothorax and 546-938 (686)0 wing expanse.

Body setae : Antennae with numerous fleshy and a few hair-like setae, about 31 o long.
Body and legs with much shorter hair-like setae only.

Head : Subtetrahedron ; subtriangular in dorsal view and rather rounded in front view ;
ventral preocular depression absent. Length from apex to postoccipital ridge 128-153 ( x 4 ) a '•
from apex to neck 183-214 (192)0 ; width across the genae 226-244 ( 2 35)( x - Midcranial
ridge cruciform. Postoccipital ridge laterally forked into anteriorly and posteriorly directed
branches. Dorsomedial part of epicranium slightly raised. Pre- and postocular ridges well
separated ; preoral ridge slender. Eyes : Dorsal simple eyes slightly projecting beyond the
outer margins of the head in dorsal view ; their corneae 15-21 (18)0 in diameter and separated
by 140-156 (150)0 ; i.e. 6-7-9-8 (8-2) times their diameter apart. Ventral simple eyes 12-18
(16) o in diameter and 61-82 (70) o apart. Ocellus vestigial and only represented by a sclerotized
spot, attached to the postocular ridge by means of a slender sclerotized arm. Cranial
apophysis apically bifurcate. Tentorial bridge stout ; anterior tentorial arms join the cranial
apophysis separately (Text-fig. 46, I). Mouth opening situated on a large mouth tubercle
posteriorly. Dorsal head setae 8-14 (io-i) on each side ; each gena usually with 2-4 and
occasionally 1 or 5 (av. 2-9) genal setae. Ventral head setae : Setae between the ventral eyes
absent ; 3-5 (4-2) just anterior to the preocular ridges, forming on both sides a transverse
irregular row ; 1-2 (i-6) on each side of the ventral arm of the midcranial ridge.

A small body of unknown homology (X) always present on each side of the dorsal arm of the
midcranial ridge, near the apex of the head.

Antennae : Rather moniliform ; 10-segmented ; 491-583 (534)0. long, i.e. shorter than

OF l'SElDOCOCCIDAE & ERIOCOCCIDAE 187

half the length of body, the ratio being 1 : 2-3-27 (2-5), and slightly shorter than the hind
legs, the ratio about 1 : 11. Scape 46-58 (52) u. long and 61-70 (64)0. wide at base ; almost
always with 4 and occasionally 5 h.s. (av. 42). Pedicel 58-67 (6i)ji. long and 46-52 (50) u, wide ;
with 1-4 (22) f.s., 3-7 (5-5) h.s. and a sensillnm placodeum. Flagellum : Flagellar segments
III to IX 37-43^ wide (segments VI, VII and sometimes VIII somewhat wider than the
preceding segments) ; segment X 31-34;/ wide. Segment III club-shaped and longest of all,
the ratio its length to the length of segment X 1-5-2-0 (i-8) : 1, and its width to its length
1 : 1-8-2-3 ( 21 )- Segments IV to IX barrel -shaped, the ratio width to length of segment IX
being 1 : 0-9-1-1 (i-o). Terminal segment pear-shaped. Segments VI to X and sometimes V
also, with a few apically knobbed capitate sensory setae. The lengths of the flagellar segments
and the number of setae on each are given in the following table :

III IV V VI VII VIII IX X

length in jx

73-92

46-58

46-61

46-61

43-55

37-46

37-46

43-55

(av.)

(82)

(49)

(58)

(55)

(49)

(43)

(40)

(46)

f.s.

7-15

7-1 1

8-16

12-17

13-17

7-*4

8-12

4-8

(av.)

(1 1 -9)

(9-2)

(n-8)

(14-1)

(149)

(103)

(9-2)

(.VI)

h.s.

3-6

i-3

i-3

i-3

1-2

0-2

O-I

O-I

(av.)

(5-0

(2-1)

(i"9)

(i-i)

(1-2)

"(o-8)

(o-5)

(04)

set. ca.

O

0-2

i-4

2-5

2-5

2-4

3-4

(av.)

O

(i-i)

(1-9)

(3-2)

(3-i)

(2-6)

(3-6)

A ntennal bristles well denned ; segments VI 1 1 and IX each with one ventral bristle. Segment
X with one dorsal preapical bristle, two lateral ones at a greater distance from the apex and two
apical small hair-like setae.

Thorax : 372-471 (426) y. long. Pronotal ridges medially interrupted at a weakly sclerotized
point ; lateral pronotal sclerites and post-tergites small. Proepistermim without any ridge-like
sclerotization. Prosternum narrow, 24-34 ( 2 7)|^ long ; prosternal ridge 79-92 (89) u. long.
Prothoracic setae almost always absent but one medial pronotal, one lateral pronotal and one
anlespiracular dorsal seta were found on each side of one specimen.

Mcsothorax : Prescutum 43-52 (46) [x long and 122 143 { 1 3 1 ) jjl wide, the ratio being 1 : 2-5-
3-2 (2-9) ; prescatal ridge slender and prescutal suture distinct. Scutum 31-55 (49) u. long, i.e.
the ratio lengths of prescutum to scutum 1 : 0-7-1-4 (i-i). Prealare, prealar ridge, tegula and
triangular plate conspicuously degenerate. Scutellum trapezoid in dorsal view ; 43-61 (55) t*.
long and 159-183 (168) u. wide, the ratio being 1 : 2-6-3-7 (3 1 ). anci i ts length to the length of
scutum 1-0-1-5 ( I-1 ) : l '• scutoscutellar suture weakly developed and usually partly missing ;
scutellar ridge absent. Postalare with the anterior and posterior ridges anteriorly joined.
Postnotal apophysis greatly reduced. Mesopleuron : Mesopleural ridge continuous above the
coxal articulation, dorsally supporting a small pleural wing process. Basalare extremely weak
and slender. Subepisternal ridge atrophied and sometimes absent altogether. Mesepisternum
well-defined and mesepimeron small. Later opleurite comparatively large, with a small oval
membranous area. Mesosternum : Basisternum 107-131 (ii9)|x long and 177-201 (i86)u.
wide ; marginal ridge usually partly absent anteriorly ; precoxal ridge slender and furca greatly
reduced. Mesothoracic spiracle 18-24 (21) u- wide at opening, with a 46-52 (49) u. long supporting
bar. Mesothoracic setae on each side : Prescatal setae absent ; sculal setae 1-5 (2-9) ; scutellar
setae 1-3 (i-8). Tegular setae 0-1 (0-2). Postmesostigmatal and basisternal setae absent.
Metathorax : Metapostnotal sclerites narrow ; metapostnotal ridge absent. Metapleural ridge
weak, attenuated near the middle as usual, and dorsally reduced. Precoxal ridges absent and
metasternal apophysis vestigial. Metathoracic spiracle similar to the mesothoracic. Mela-
thoracic setae on each side : Metatergal setae 3-8 (47) and anterior metasternal setae 0-1 (0-9) ;
metapleural setae absent and posterior metasternal setae 0-1 (0-3).

Wings : Greatly reduced ; 126-308 (182) u. long and 42-140 (84)0. wide ; axillary sclerites
small ; additional sclerite and alar lobe absent ; one alar seta almost always present and

MORPHOLOGY AND TAXONOMY OF ADULT MALES

set.ca.

Fig. 49. Gossyparia spuria (Mod.), brachypterous form, dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 189

occasionally absent (av. o-8). Hamulohalterae atrophied and sometimes absent altogether ;
when present, 31-52 (40)^ long and 12-18 (15)0. wide ; the hamulohalteral ridge and the seta
absent.

Legs : Well developed, short and stout ; the middle legs usually shortest and the hind legs
longest ; the ratio length of the hind leg to the total length of body 1 : 2-2-2-4 ( 2 '3)- Coxa
and trochanter about 64 and $jy. wide respectively ; the latter with a long apical seta. Femur
about 49(1 wide ; that of the middle leg shortest and those of the fore and hind legs subequal ;
the ratio width to length of the hind femur 1 : 2-9-3-4 (3 1 )- Tibia about 31(1 wide ; with 2
apical spurs ; in fore leg the femur shorter than the tibia, the ratio their lengths being 1 : 11-
1-2 (1-15)- Tarsus about 27(1 wide ; tarsal digitules apically knobbed, about 34(1 long. Claw
wide at base, abruptly tapering apically ; ungual digitules apically knobbed. The lengths of
the leg segments (in microns) and the number of setae on each are given in the following table :

Fore leg Middle leg Hind leg

Coxa

length
h.s.

52-58 (55)
7-i 1 (9-5)

52-58 (55)
7-10 (8-9)

55-61 (58)

8-13 (117)

Trochanter

length
h.s.

64-70 (67)
4-5 (4-5)

64-70 (68)
4-5 (4-4)

7°-79 (73)
4-6 (5-0

Femur

length
h.s.

143-168 (156)
11-16 (131)

134-153 (143)
8-12 (104)

140-168 (153)
9-13 (ii-8)

Tibia

length
h.s.

168-189 (180)
16-22 (185)

168-189 (180)
15-23 (i9i)

177-214 (198)
14-21 (18-2)

Tarsus

length
h.s.

70-76 (72)
9-12 (10-7)

70-79 (74)

9-13 (II-2)

73-85 (79)
11-14 (125)

Claw

length

21-24 (22)

21-24 (22)

21-24 (22)

Total length

of leg

516-583 (549)

506-57O (54O)

534-622 (583)

Abdomen : 570-646 (608) u. long and 365-426 (395) (x wide. Tergites of segments I and II
weak and narrow ; that of segment VIII large. Sternites of segment VIII ill-defined. Ostioles
absent. Abdominal setae on each side : Dorsal setae usually 3-5 on segments I to VI (some-
times 7 on segment I and 6 on segments II and IV), 2-4 on segments VII and VIII, including
a slightly longer seta on the latter. Pleural setae 3-4 and sometimes 5 on segments I to VII,
and always 4 on segment VIII, including one considerably longer seta (85-95, av. 89(1 long).
Ventral setae 1 on segment II, 1-2 on segment III, 2-3 on segments IV to VII, and 1-2 on
segment VIII.

Glandular pouches comparatively very small ; setae 0/ glandular pouch only include a pair of
comparatively short tail setae, 46-92 (73) u. long, i.e. the ratio their length to the total length of
body 1 : 15-2-26-3 (18-6).

The appearance and structure of the genital segment similar to that of the macropterous form ;
genital capsule 162-192 (177)^ long and 113-131 (i22)u. wide, the ratio 1-4-1-5 (145) : 1, and
the ratio its length to the total length of body 1 : 7-0-8-1 (7-7). Anal opening 15-21 (i8)[i
in diamter. Setae of genital segment on each side : Dorsally two long setae always present
near the base of the style ; ventrally 8-12 (9-6) slightly smaller setae occur on the genital
capsule.

Material : 10 specimens examined, received in two lots as follows :

1. 5 specimens collected by A. Dziedzicka, on Ulmus sp., in Krakow, Poland, during
May, 1964.

2. 5 specimens also collected by A. Dziedzicka, on Ulmus sp., in Moglia, nr. Krakow,
Poland, 26. iv. 1963.

igo MORPHOLOGY AND TAXONOMY OF ADULT MALES

Gossyparia salicicola Borchsenius
(Text-figs. 50, 51)

Brachypterous form only known. The males rather robust, with the abdomen slightly
broader than the thorax ; comparatively large, with short antennae and short legs. When
mounted 1302-1442 (1373) [x long, 322-350 (336) y. wide at mesothorax and 532-980 (644) [z wing
expanse.

Body setae : Antennae with numerous fleshy and a few hair-like setae, about 28 jx long.
Body and legs with much shorter hair-like setae only.

Head : Subtetrahedron ; subtriangular in dorsal view and rather rounded in front view ;
ventral preocular depression absent in lateral view. Length from apex to postoccipital ridge
131-143 (137) (^ ; from apex to neck 189-204 (195) jx ; width across the genae 223-244 (229)^1.
Midcranial ridge cruciform. Postoccipital ridge laterally forked. Preocular and postocular
ridges well separated ; preoral ridge slender. Eyes : Dorsal simple eyes slightly projecting
beyond the outer margins of the head in dorsal view ; their corneae 18-21 (20) y. in diameter
and separated by 134-165 (146) |x, i.e. 6-3-8-5 (7-0) times their corneae apart. Ventral simple
eyes 18-21 (19) [x in diameter and 73-95 (82) [x apart. Ocellus vestigial and merely represented
by a sclerotized spot, attached to the postocular ridge by means of a slender sclerotization.
Cranial apophysis apically bifurcate. Tentorial bridge stout. Mouth tubercle large. Dorsal
head setae 10-15 ( I2 '5) on each side ; each gena usually with 3-6 and occasionally 8 (av.) 4-7)
genal setae. Ventral head setae : Setae between the ventral eyes absent ; 2-4 (2-8) just
anterior to the preocular ridges, forming on both sides a transverse irregular row ; 1-2 (1-5)
on each side of the ventral arm of the midcranial ridge. The mouth tubercle with 1-2 (i-6)
minute setae.

A small body of unknown homology (X) occurs dorsally on each side near the apex of the
head.

Antennae : Rather moniliform ; 10-segmented ; 522-583 (549)^1 long, i.e. shorter than half
the body length, the ratio being 1 : 2-3-2-6 '2-5), and slightly shorter than the hind legs, the
ratio 1 : 1-1-1-2 (1-13). Scape 46-52 (49)(x long and 55-61 (58)jx wide at base ; always with
4 h.s. Pedicel 61-67 (64) (x long and 43-46 (45) (x wide ; with 1-2 (i-i) f.s., 2-3 (2-4) h.s. and a
sensillum placodeum. Flagellum : Segment III-IX 34-40^ wide and segment X 27-31 (29) jx
wide. Segment III club-shaped and longest of all, the ratio its length to the length of segment
X 1-7-2-1 (1-9) : 1, and its width to its length 1 : 2-2-2-5 ( 2 '3)- Segments IV to IX barrel-
shaped, the ratio width to length of segment IX being 1 : 1-0-1-4 ( I-2 )- Terminal segment
pear-shaped. Apically knobbed sensory setae entirely absent on the antennae. The following
table shows the lengths of the flagellar segments and the number of setae on each :

III IV V VI VII VIII IX X

length in (x

76-92

46-61

52-67

49-64

49-61

40-49

37-46

43-4<

(av.)

(85)

(50)

(61)

(58)

(52)

(46)

(43)

(45)

f.s.

3-8

3-9

5-9

7-12

8-15

5-8

4-8

2-4

(av.)

(5-i)

(6-2)

(7"4)

(9-5)

(io-8)

(7-i)

(6-2)

(2-8)

h.s.

2-6

2-4

1-2

1-2

i-3

O-I

1-2

O-I

(av.)

(3-3)

(2-9)

(i-5)

(i-3)

(i-8)

(07)

(1-2)

(o-6)

A ntennal bristles well-defined ; segments VIII and IX each with one ventral bristle. Terminal
segment with one dorsal preapical bristle, two lateral ones at a greater distance from the apex
and one or two apical small hair-like setae.

Thorax : 403-464 (433) (x long. The thoracic structures, ridges and plates similar to G.
spuria. Prosternum and prosternal ridge 21-27 (24) and 82-95 (89) y- long respectively.
Prothoracic setae on each side : Medial pronotal setae 0-3 and occasionally 5 (av. 1 -4) ; lateral

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE

191

pronotal and antespiracnlar dorsal setae usually absent but 1-2 setae occasionally occur on either
or both sides (averages 0-5 and 04 respectively). Antespiracnlar ventral and prosternal setae
absent.

gts A

Fig. 50. Gossyparia salicicola Borchsenius, dorsal and ventral view.

IQ2

MORPHOLOGY AND TAXONOMY OF ADULT MALES

Mesothorax : Prescutum 52-64 (58) (j. long and 1 19-137 (131)0. wide, the ratio being 1 : 1-9-
2-6(2-3). Scutum 24-40 (32) (x long, i.e. the ratio lengths of prescutum to scutum 1-3-2-6 (1 -8) : 1.
Scutellum 40-58 (52) y. long and 150-165 (156)^ wide, the ratio 1 : 2-7-3-8 (3-0), and its length
to the length of scutum 1-2-2-1 (i-6) : 1. Basisternum 101-122 (iio)u. long and 168-183
(177)^ wide. Mesothoracic spiracle 18-21 (2o)[x wide at opening, with a 49-55 (52)fz long
supporting bar. Mesothoracic setae on each side : Prescutal seta 0-1 (0-3) ; scutal setae 2-6
(3'3) '• scutellar setae 1-3 (i-6). Tegular setae 0-1 (0-3). Postmesostigmatal and basisternal
setae absent. Metathoracic setae on each side : Metatergal setae 6-14 (8-6) ; one anterior
metasternal seta always present ; metapleural and posterior metasternal setae absent.

Wings : Greatly reduced ; 112-336 (168) u. long and 42-98 (56) \i wide ; axillary sclerites
small ; additional sclerites and alar lobe absent ; alar seta 0-1 (o-6). Hamulohalterae atrophied,
27-49 (37) pi long and 12-15 (14) u. wide.

Legs : Short and stout ; the fore and the middle legs subequal in length and the hind legs
longest ; the ratio length of the latter to the total length of body 1 : 2-1-2-3 ( 2- 2). Coxa and
trochanter about 64 and 40^ wide respectively ; the latter with a differentiated long apical seta.
Femur about 490. wide ; that of the middle leg shortest and those of the fore and hind legs
subequal ; the ratio width to length of the hind femur 1 : 3-3-3-5 (3-4). Tibia about 310.
wide ; with 2 apical spurs ; in fore leg the ratio lengths of femur to tibia 1 : 1-17-1-24 (1-2).
Tarsus about 24^ wide ; tarsal digitules apically knobbed, about 3751 long. Ungual digitules
also apically knobbed. The following table shows the lengths of the leg segments (in microns)
and the number of setae on each :

Fore leg

Middle leg

Hind leg

Coxa

length
h.s.

49-55 (52)
5-9 (7-4)

49-55 (52)
5-io (8-2)

52-58 (55)
6-1 1 (8-5)

Trochanter

length
h.s.

64-67 (66)
3-4 (3-5)

64-70 (67)
3-4 (3-4)

67-73 (70)
3-5 (3-9)

Femur

length
h.s.

159-168 (165)

8-I5 (I2-I)

153-159 (156)
6-12 (9-5)

162-171 (168)

7-13 (II-2)

Tibia

length
h.s.

189-204 (198)
15-19 (16-8)

192-207 (201)
13-19 (17-1)

204-223 (217)
I3-20 (17-6)

200^-

Fig. 51. Gossyparia salicicola Borchsenius, lateral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 193

Tarsus length
h.s.

76-82 (79)
7-10 (8-2)

79-89 (86)
7-11 (9-1)

85-92 (89)
8-12 (99)

Claw length

18-24 ( 2I )

18-24 (21)

18-24 ( 2I )

Total length of leg

558-595 (580)

558-595 (583)

592-634 (619)

Abdomen : 555-684 (623)1* l° n g an d 388-418 (410)0. wide. Abdominal setae on each side :
Dorsal setae 4-9 on segments I and II, 4-7 on segments III and IV, 3-6 on segments V to VII,
and 2 on segment VIII, including one slightly longer seta. Pleural setae 3-4 on segment I,
4-5 on segments II to VI, and 3-4 on segments VII and VIII, including one very long seta
on the latter segment (107-137, av. n6u). Ventral setae 1-2 on segment II, usually 3 on
segments III to VII (sometimes 2 on segments III and IV, or 4 on segments IV and VII), and
1 on segment VIII.

Glandular pouches very small ; setae of glandular pouch only include a pair of comparatively
short tail setae, 40-82 (64) u. long, the ratio their length to the total length of body being
1 : 17-1-34-9 (20-8). Genital segment typical of G. spuria ; genital capsule 146-171 (162)0.
long and 113-122 (119)0. wide, the ratio being 1-3-1-5 (1-4) : 1, and the ratio its length to the
total body length 1 : 7-8-9-8 (8-5). Anal opening 18-24 ( 22 )l* m diameter. Setae of genital
segment on each side : Dorsally two relatively long setae present near the base of the style ;
ventrally 7-9 (7-8) smaller setae occur on the genital capsule.

Material : 10 specimens examined, collected by G. Matesova on Salix sp., in
Alma Ata, Kazakhstan, U.S.S.R., 2.iv.i95i.

Remarks : This species, although very closely related to the brachypterous males
of G. spuria, can be distinguished mainly by the absence of the capitate sensory
setae on the antennal segments ; other minor differences are the acquisition of
more numerous abdominal setae and longer anal lobe seta.

PSEUDOCHERMES Nitsche, 1895

Pseudochermes fraxini (Kalt.)

(Text-fig. 52)

The apterous forms only known ; these are degenerate, and appear like the nymphal forms.
The males dorso-ventrally flattened, moderately robust and spindle-shaped, i.e. the body
broadest at mesothorax and gradually becoming narrower posteriorly ; the body ridges and
sclerites absent or greatly reduced. A comparatively very small species, with very short
antennae and legs ; mounted specimens 518-658 (602)0. long and 196-238 (210)0. wide at
mesothorax.

Body setae : Antennae with very few fleshy and hair-like setae, 9-1 2 o. long ; the body with
sparse, slightly smaller hair-like setae only.

Head : Intimately fused with thorax ; head ridges entirely absent. Eyes represented by
weakly sclerotized lateral spots, 89-95 (9 2 )! 1 apart. Cranial apophysis vestigial, apically
bifurcate. Mouth opening situated on a small, double-bulging mouth tubercle posteriorly.
Dorsal head setae 3-4 (3-8) and ventral head setae 2-3 (2-8) on each side of the median line.

Antennae : Neither filiform nor moniliform, with segment VI distinctly widest of all flagellar
segments ; normally 8-segmented, but sometimes fusion between adjacent segments occurs ;
146-174 (162) u. long, i.e. much shorter than half the body length, the ratio 1 : 3-5-3-9 (3-7),
and slightly shorter than the hind legs, the ratio 1 : 1-1-1-2 (1-17). Scape 18-24 ( 2 °)v- l° n g

194

MORPHOLOGY AND TAXONOMY OF ADULT MALES

and 24-31 (28) [x wide ; with 2-3 (2-4) h.s. Pedicel 18-27 ( 2I )t x l°ng and 21-24 ( 2 3)p>- wide ; with
1-2 (i-6) h.s. Flagellum : Segment III club-shaped, with a very narrow stalk ; the ratio its
length to the length of segment VIII being 1-2-1-8 (1-5) : 1, and the ratio width to length of

dhs vhs

Pdc \ , scp

Ptr-

— sp 3

Fig. 52. Pseucochermes fraxini (Kalt.), apterous form, dorsal and ventral view.

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 195

segments III and VII 1 : 1-2-1-8 (1-4) and 1-1-2 (i-i) : 1, respectively. Segments V and VI
becoming progressively wider, and segments VII and VIII decreasing in width.

The lengths, widths of the flagellar segments and the number of setae on each are given in
the following table :

III IV V VI VII VIII

length in u. 21-27 15-21 18-24 18-21 18 15-18

(av.) (25) (17) (20) (19) (18) (17)

width in ja 15-18 15-18 18-24 21-24 18-21 15-18

(av.) (17) (17) (21) (23) (20) (16)

f.s. 001 1-2 1-2 o

(av.) o o (1) (1-2) (1-2) o

h.s. 0-1 0-1 1-2 0-1 1-2 0-1

(av.) (0-2) (0-4) (i-8) (0-2) (i-8) (o- 4 )

Antennal bristles well defined ; segments VI and VII each with one ventral bristle. Terminal
segment with 3 preapical bristles, a pair of capitate sabapical sensory setae and one apical
hair-like seta.

Thorax : The thoracic ridges and sclerites greatly degenerate. ' Pleural ridges of thorax
short and rudimentary. Spiracles 9-12 (10) u. wide at opening, with 24-31 (27)^ long supporting
bars. Prothoracic and mesothoracic setae : 2-3 (25) dorsal, one pleural, and one ventral setae
present on each side of the median line. Metathoracic setae : 2-3 (2-7) dorsal, and one pleural
setae occur on each side.

Legs : Comparatively short and very stout , the fore and the middle legs subequal in length,
the hind leg slightly longer ; the ratio length of the hind leg to the total length of body
1 : 2-9-3-4 (3 -2 )- Coxa and trochanter about 31 and i8[i wide respectively ; the latter with
a differentiated long apical seta. Femur about 28^ wide ; that of the middle leg shortest and
those of the fore and hind legs subequal ; the ratio width to length of the hind femur being
1 : 1-8-2-0 (1-9). Tibia about i8(x wide ; with 2 small spines ; in fore leg the ratio lengths of
femur to tibia 1-4-1-5 (1-45) : 1. Tarsus about i8(x wide ; tarsal digitules apically knobbed,
about 24 u. long. Claw broad at base and abruptly tapering apically ; ungual digitules also
apically knobbed. The lengths of the leg segments (in microns) and the number of setae on
each are given in the following table :

Fore leg Middle leg Hind leg

24-31 (28) 24-31 (28) 27-31 (29)

2-3 (2-2) 2-3 (2-2) 2-3 (2-6)

18-21 (19) 18-21 (19) 21-24 (22)

(1) (1) (1)

49-55 (52) 49-55 (53) 49~55 (53)

1-2 (l-2) 1-2 (l-2) 1-2 (1-4)

34-37 (35) 34-37 (35) 37-4° (39)

000

27-31 (29) 27-31 (29) 27-31 (30)

2-3 (2-4) 2-3 (2-2) 2-3 (2-2)

I5-I8 (16) I5-18 (16) I5-I8 (16)

I68-183 (180) I7I-183 (180) 177-195 (189)

Abdomen : Abdominal tergites and sternites absent. Abdominal setae on each side :
Dorsal setae usually 2 and sometimes only 1 on segments I to VIII. Pleural setae always 1.
Ventral setae 1 on segment II, 2 on segments III to VI, and 1-2 on segments VII and VIII.

Ostioles as well as glandular pouches entirely absent. Genital segment : Genital capsule

Coxa

length
h.s.

Trochanter

length
h.s.

Femur

length
h.s.

Tibia

length
h.s.

Tarsus

length
h.s.

Claw

length

Total length of

leg

i 9 6 MORPHOLOGY AND TAXONOMY OF ADULT MALES

comparatively very small, 37-40 (39) pt long and 31-34 (32) [i. wide, i.e. the ratio 1-2-1-3 ( I-2 3) : *■
and its length to the total length of body 1 : 14-1-17-0 (15-4) ; style rather straight in lateral
view, apically rounded. Aedeagus ventrally sclerotized and tapering apically. Setae of genital
segment : The genital capsule always with one dorsal and two ventral setae on each side.

Material : 10 specimens examined, collected by K. Boratynski, on ash trees
(Fraxinus excelsior), in the gardens of Buckingham Palace, London, 20.x. 1963.

Remarks : The mating behaviour of this species was observed in the laboratory.
The females, covered with copious waxy secretion, remain fixed usually deep in
the crevices of the bark with their heads at the bottom of the crevices and their
abdomina directed towards the surface. The male wandering about, somehow
is capable of locating the position of the posterior end of the female ready for
copulation, pushes its abdomen through the waxy covering and copulation takes
place. These observations may suggest that the females remain under their waxy
covering until the eggs are fertilized, then migrate to another position to start
laying the eggs ; those unmated appear to remain in their positions until they
perish.

KEYS

The following keys are intended to separate the studied species and their supra-
specific categories. Since the significance of some characters is still uncertain, the
keys will include as many characters as possible, even more than actually necessary
for this purpose. It should be noticed that the numbers of setae and disc pores
given in these keys are always on one side only.

Macropterous Males

Key to Pseudococcidae and Eriococcidae

Postoccipital ridge weak, slender, U- or V-shaped. Tentorial bridge comparatively
slender. Flagellar segments of the antennae, including the terminal, cylindrical
and elongated. Dorsal margin of the proepisternum ridge-like. Scutellum
pentagonal in dorsal view ; its length half its width or more ; scutellar ridge absent.
Anterior and posterior postalar ridges well separated. Mesopleural ridge inter-
rupted above the coxal articulation. Basal part of the trochanter conspicuously
longer than the distal. Claw gradually tapering apically ; ungual digitules not
knobbed. Dorsal setae of abdominal segment VIII subequal in length to other
abdominal setae ; setae of the glandular pouch include a pair of long tail setae and
one or more much shorter ones. Basal part of the penial sheath short, not fused
with the 9th tergite + 10th segment. Anal opening minute. Aedeagus arising
just behind the basal ridges of the penial sheath. Dorsal setae of the genital
segment similar to other abdominal setae, always 3 or more. Derm pores present
(except in Nairobia bifrons). The puparia fluffy and loosely felted Pseudococcidae (p. 93)

Postoccipital ridge strongly developed and laterally forked. Tentorial bridge compara-
tively stout. Terminal flagellar segments of the antennae barrel-shaped, often as
long as wide ; the terminal segement distinctly pear-shaped. Proepisternum with-
out any ridge-like sclerotization. Scutellum transverse, rectangular in dorsal view ;
its length less than half its width ; with a well developed scutellar ridge on each

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 197

side. Anterior and posterior postalar ridges anteriorly joined. Mesopleural ridge
continuous above the coxal articulation. Basal part of the trochanter shorter
than the distal. Claw broad at base, abruptly tapering apically ; ungual digitules
apically knobbed. One of the dorsal setae on each side of abdominal segment VIII
much longer than the others ; setae of the glandular pouch include one pair of tail
setae only. Basal part of the penial sheath long, fused with the 9th tergite + 10th
segment and forming together a compact genital capsule. Anal opening large.
Aedeagus arising at a comparatively large distance from the basal ring of the genital
capsule. Dorsal setae of the genital segment considerably longer than other
abdominal setae, always 2. Derm pores absent. The puparia compact, closely
felted and rather flattened ....... Eriococcidae (p. 167)

Pseudococcidae
Key to Groups of Genera

Dorsal, lateral and ventral arms of the midcranial ridge joined, forming
together a cruciform structure. Pre- and postocular ridges widely separated ;
interocular ridge absent. Anterior tentorial arms each apparently indivi-
dually fused with the cranial apophysis. Penultimate antennal segment
more than 37 times as long as wide. Prescutum triangular in dorsal view.
Scutum without a median longitudinal narrow membranous area, and much
shorter than either the prescutum or the scutellum. Precoxal ridge of the
metathorax weakly developed ; metapleural wing process absent. Tarsal
digitules absent. Setae of the glandular pouch include two long tail setae
and a short one, subequal in length to the other abdominal setae ; the tail
setae comparatively short, the body being 5 times as long, or longer . . 2

Dorsal arm of the midcranial ridge distinctly detached from the Y- or T-shaped
ridge formed by the lateral and the ventral arms. Pre- and postocular
ridges joined together by means of a well developed interocular ridge.
Anterior tentorial arms anteriorly fused with each other before joining the
cranial apophysis. Penultimate antennal segment less than 3-7 times as
long as wide. Prescutum transverse, rectangular in dorsal view. Scutum
with a distinct median longitudinal narrow membranous area, and longer
than either the prescutum or the scutellum. Precoxal ridge of the meta-
thorax well developed ; with a small metapleural wing process. Tarsal
digitules present. Setae of the glandular pouch include a pair of long tail
setae and one or more setae conspicuously longer than other abdominal
setae ; the tail setae comparatively long, the body being less than 5 times
as long .... ....... 3

(1) Disc pores absent. Lateral ocelli vestigial. Third and terminal antennal
segments subequal in length (third not more than 11 times as long as
terminal) ; the length of segment III less than 5 times its width. Antennal
setae subequal in length to the body setae ; capitate, apically knobbed
sensory setae present on segments III to X. Proepisternum with only the
dorsal margin ridge-like. Prescutal ridges well separated postero-medially.
Postmesostigmatal setae, alar setae and metapostnotal ridge absent. Fore
legs shortest. Ostioles absent. Dorsal setae of abdominal segment VIII
absent. Glandular pouches and their setae only on abdominal segment
VIII ; the tail setae relatively short, the body being 8 or more times as
long. Basal ridges of the penial sheath medially continuous

NAIROBIA group (p. 163)
[Only one species, N. bifrons]

198 MORPHOLOGY AND TAXONOMY OF ADULT MALES

Disc pores present. Lateral ocelli well developed. Third antennal segment
longer than the terminal (1-4 or more times as long) ; the length of segment
III more than 5 times its width. Antennal setae about twice as long as the
body setae ; capitate, apically knobbed sensory setae entirely absent.
Proepisternum with dorsal and ventral margins ridge-like. Prescutal ridges
continuous postero-medially. Postmesostigmatal setae, alar setae and
metapostnotal ridge present. Middle legs shortest. Ostioles present.
Dorsal setae of abdominal segment VIII present. Glandular pouches and
their setae on abdominal segments VII and VIII ; the tail setae of segment
VIII rather long, the body being less than 8 times as long. Basal ridges of
the penial sheath medially separated . . . CEROPUTO group (p. 155)

3 (1) Antennae comparatively long (the body 1-5 or less times as long) ; longer than

the hind legs. Few capitate, apically knobbed sensory setae present on
antennal segments III to X. Antespiracular dorsal setae and pores absent.
Basalare weak and slender. Abdominal disc pores only present on segment
I. Setae of the glandular pouch include a pair of long tail setae, two setae of
medium, unequal lengths and a short one . OCTOCOCCUS group (p. 151)

[Only one species, O. africanus.]
Antenna comparatively short (the body i-6 or more times as long) ; shorter
than the hind leg, or both subequal in length. Capitate, apically knobbed,
sensory setae only present on terminal antennal segment. Antespiracular
dorsal setae and pores present. Basalare comparatively stout. Abdominal
disc pores present on all abdominal segments. Setae of the glandular pouch
include a pair of long tail setae and one of medium length or short . . 4

4 (3) The head and the body dorso-ventrally flattened ; ventral preocular depression

entirely absent. Postoccipital ridge V-shaped. Postocular ridge dorsally
reduced. Prosternal ridge absent. Two antespiracular ventral setae
always present. Prescutal suture absent. The marginal ridge of the
basisternum anteriorly absent. Furca comparatively small. Hind legs
short, the body more than 2-2 times as long SACCHARICOCCUS group (p. 143)

[Only one species, S. sacchari.]

— The body cylindrical ; ventral preocular depression present. Postoccipital

ridge U-shaped. Postocular ridge well developed. Prosternal ridge
distinct. Only one antespiracular ventral seta present. Prescutal suture
well-defined. The marginal ridge of the basisternum well developed.
Furca comparatively large. Hind legs long, the body less than 2-2 times
as long 5

5 (4) Fleshy setae present on the body itself, giving the males a distinctive hairy

appearance. Ventral ocular setae present. Genal setae more than 7, and
prosternal setae more than 3 . . . . PSEUDOCOCCUS group (p. 131)

- Fleshy setae absent on the body itself. Ventral ocular setae absent. Genal

setae less than 7, and prosternal setae less than 3 PLANOCOCCUS group (p. 93)

Key to CEROPUTO Group of Genera

Fleshy setae present on antennae and legs. Dorsal arm of midcranial ridge distinct.
Setae between the ventral simple eyes absent. Terminal antennal segment with
3 preapical large bristles and 2 smaller ones ; the preapical bristles about twice as
long as those of the two penultimate segments. Prosternum represented by a pair
of separate, small plates connected by the prosternal ridge. Prescutum 4 times or
more as long as scutum. Scutellum laterally supported by a distinct additional
ridge. Lateropleurite with an oval membranous area. Postmesostigmatal pores

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 199

absent. Ostioles well developed. Aedeagus comparatively slender. More than 3
dorsal setae of the genital segment present . . . CENTROCOCCUS (p. 159)

[Only one species, C. insolitus.]
Fleshy setae entirely absent. Dorsal arm of the midcranial ridge only marked by weak
sclerotization. Setae between the ventral simple eyes present. Terminal antennal
segment with only three well defined preapical bristles, subequal in length to those of
the two penultimate segments. Presternum triangular. Prescutum less than 3-5
times as long as scutum. Scutellum without lateral edges. Lateropleurite without
membranous areas. Postmesostigmatal pores present at least on one side.
Ostioles ill-defined. Aedeagus rather stout. 3 dorsal setae of the genital segment

always present CEROPUTO (p. 155)

[Only one species, C. pilosellae.]

Key to PSEUDOCOCCUS Group of Genera

Dorsal arm of the midcranial ridge disappearing before meeting postoccipital ridge ;
the latter anteriorly confluent with the preocular ridge. Dorsal ocular setae
present. One dorsal head pore usually present. Tegular setae less than 4. Abdominal
disc pores occur laterally and also dorsally and ventrally . DYSMICOCCUS (p. 139)

[One species only, D. alazon.]

Dorsal arm of the midcranial ridge posteriorly meeting the postoccipital ridge ; the
latter not reaching the preocular ridges anteriorly. Dorsal ocular setae absent.
More than one dorsal head pore present. Tegular setae more than 4. Abdominal
disc pores present only laterally PSEUDOCOCCUS (p. 131)

Key to PLANOCOCCUS Group of Genera

1 Style of the genital segment more or less straight and apically rounded in lateral

view ............. 2

Style curving upwards and apically pointed in lateral view ... 3

2 (1) Dorsal arm of midcranial ridge absent. Head disc pores absent. Antennal

setae about twice as long as the body setae ; the dorsal bristle of the
terminal antennal segment slightly shorter than the lateral bristles. Post-
tergital setae present. Prosternal and postmesostigmatal pores absent ;
metatergal pores present. Abdominal disc pores only present laterally. The
body 10 times or more as long as the penial sheath ; the style comparatively
broad. Aedeagus rather short, curving dorso-posteriorly FERRISIANA (p. 122)

[Only one species, F. virgata.']
Dorsal arm of midcranial ridge distinct. Head disc pores present. Antennal
setae about as long as the body setae ; the dorsal bristle of the terminal
antennal segment much shorter than the lateral ones (about half as long).
Post-tergital setae absent. Prosternal and postmesostigmatal pores present ;
metatergal pores missing. Abdominal pores present laterally and also
dorsally and ventrally. The body 8-5 times or less as long as the penial
sheath ; the style comparatively narrow. Aedeagus rather long, anteriorly
curving from its basal rod and reaching the cavity of abdominal segment

VII MACONELLICOCCUS (p. 118)

[Only one species, M. hirsutus.]

3 (1) Dorsal arm of midcranial ridge posteriorly meeting the postoccipital ridge.

One dorsal head pore only usually present. Antennal scape almost always
with 4 hair-like setae. Postmesostigmatal pores present, at least on one
side. Ostioles ill-defined or absent ........ 4

>oo MORPHOLOGY AND TAXONOMY OF ADULT MALES

- Dorsal arm of midcranial ridge fading out well before meeting the postoccipital

ridge. Two or more dorsal head pores present. Antennal scape with more
than 4 hair-like setae. Postmesostigmatal pores absent. Ostioles well
developed ............ 5

4 (3) Lateral arms of midcranial ridge represented only by weak sclerotization.

Preocular ridge with a well-marked ventral rudiment below the articular
process. Genal setae more than 3. Prosternum long and narrow. Post-
mesostigmatal setae present in the median and the sublateral areas.
Hamulohaltera 1-9 times or more as long as its apical seta. Femur of
fore-leg shortest. All pleural setae of abdominal segment VIII subequal

in length TRIONYMUS (p. 126)

[Only one species, T. newsteadi.]

- Lateral arms of midcranial ridge well developed. Ventral rudiment of the

preocular ridge absent. Genal setae 3 or less. Prosternum triangular.
Postmesostigmatal setae only present sublaterally. Hamulohaltera 16 or
less times as long as its apical seta. Femur of middle leg shortest. Pleural
setae of abdominal segment VIII include one slightly longer seta

NIPAECOCCUS (p. no)

5 (3) Ventral rudiment of the preocular ridge entirely absent. Third antennal

segment 3-3 times or less as long as wide. The three large bristles of the
terminal antennal segment of subequal lengths. In fore leg, the tibia

shorter than the femur PLANOCOCCOIDES (p. 106)

[Only one species, P. ireneus.]

— Ventral rudiment of the preocular ridge distinct. Third antennal segment

usually more than 3-3 times as long as wide. The dorsal bristle of the
terminal antennal segment slightly shorter than the two lateral ones. In
fore leg, the tibia longer than the femur . . . PLANOCOCCUS (p. 93)

PSEUDOCOCCUS

Key to Species

Penial sheath distinctly triangular in dorsal view, with the style gradually

tapering apically ........... 2

Penial sheath subtriangular in dorsal view, the style being rather broad or

truncate posteriorly .......... 3

(1) Antennae short, the body being more than twice as long. Only one ante-
spiracular ventral seta present. One postmesostigmatal pore always
present medially. Posterior metasternal seta less than 3. Ventral setae of
abdominal segment VIII absent. Abdominal pores only present laterally.
Genital segment long, the body being less than 7 times as long. Aedeagus
long, anteriorly reaching the cavity of abdominal segment VII, and evenly
tapering posteriorly ........ P. citriculus

Antennae moderately long, the body being less than twice as long. More than
one antespiracular ventral seta present. Posterior metasternal setae more
than 5. Ventral setae of abdominal segment VIII at least 4. Abdominal
pores present laterally and dorsally. The genital segment rather short, the
body being more than 8 times as long. Aedeagus not reaching the cavity
of abdominal segment VII, with the distal part broadened in lateral view

P. fragilis

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 2o t

3 (i) Post-tergital setae 9 or more. Anterior metasternal setae more than 10.

Metaspiracular and anterior metasternal pores absent. Penial sheath
broad, its length less than 15 times its width ; process of the penial sheath
well pronounced ; style apically truncate in dorsal view . . P. adonidum

Post-tergital setae 6 or less. Anterior metasternal setae less than 9. Meta-
spiracular and anterior metasternal pores present. Penial sheath moder-
ately narrow, its length more than i-6 times its width ; process of the penial
sheath absent ; apex of style broadly rounded in dorsal view ... 4

4 (3) Postmesostigmatal setae only present sublaterally. Ventral setae of abdom-

inal segment VIII absent ...... P. obscurus (p. 131)

- Postmesostigmatal setae present sublaterally and also medially. Ventral

setae of abdominal segment VIII usually present P. maritimus 'Type A' (p. 135)

NIPAECOCCUS

Key to Species

Relatively small, 840-980 (896) pL long. Fleshy setae present on the legs. The dorsal
simple eyes smaller than the ventral ; the dorsal eyes more than 4 times their
diameter apart. With less than 3 genal setae. Antennae as long as, or slightly
longer than the hind legs. The antennal setae about twice as long as the body
setae. Terminal antennal segment with 3 bristles only. Anterior apex of pro-
sternum forming a weakly sclerotized ring. With 7 or more medial pronotal pores.
Width of prescutum less than 1-4 times its length. Scutal setae less than 4. Meta-
spiracular pores and ostioles absent. Abdominal pores present only laterally

N. nipae (p. 114)

Moderately large, 1 148-1358 (1246) [i long. Fleshy setae absent on the legs. The dorsal
simple eyes larger than the ventral ; the dorsal eyes less than 4 times their diameter
apart. Genal setae almost always 3. Antennae shorter than the hind legs. The
antennal setae about as long as the body setae. Terminal antennal segment with
3 large and 2 smaller bristles. Anterior apex of prosternum without a sclerotized
ring. With less than 5 medial pronotal pores. Width of prescutum 1 -4 or more
times its length. Scutal setae more than 5. Metaspiracular pores and ostioles
present. Abdominal pores present laterally and also ventrally on one or more
segments .......... N. vastator (p. no)

PLANOCOCCUS

Key to Species

Postoccipital ridge anteriorly confluent with the preocular ridges. Dorsal head
pores 4 or more. Prosternal ridge double-barred. Post-tergital pores
present. Metatergal setae absent . . . . .P. dioscoreae (p. 102)

Postoccipital ridge not reaching the preocular ridges anteriorly. Dorsal head
pores less than 4. Prosternal ridge singular. Post-tergital pores absent.
Metatergal setae present ......... 2

(1) Antennal segment III longer than the terminal. Middle femur slightly shorter
than the front. A ventral pore present at least on one side of abdominal
segment VII. Aedeagus comparatively broad . . .P. kenyae (p. 98)

Antennal segment III shorter, or as long as the terminal. Middle femur
slightly longer than the front. Ventral abdominal pores entirely absent.
Aedeagus comparatively narrow ...... P. citri (p. 93)

MORPHOLOGY AND TAXONOMY OF ADULT MALES

ERIOCOCCIDAE

Key to Genera

Fleshy setae present on the legs. Prosternal setae present. Hamulohaltera
subequal in length to its apically hooked seta. Trochanter without long,
differentiated apical setae ERIOCOCCUS (p. 167)

Fleshy setae absent on the legs. Prosternal setae absent. Hamulohaltera,
if present, i-6 or more times as long as its apically hooked seta. Trochanter
with a long apical seta .......... 2

(1) Lateral arms of the midcranial ridge short and weak. Ocelli vestigial. Genal
setae 3 or less. Unidentified disc-like structures absent on the head. With
a small mouth tubercle. The body not more than twice as long as the
antennae. Antespiracular ventral setae present. Scutellum slightly longer
than scutum. Hamulohalterae and suspensorial sclerites entirely absent.
The body less than twice as long as the hind leg. Pleural setae of abdominal
segment VIII include one seta slightly longer than the others. Style rather

straight in lateral view OVATICOCCUS (p. 179)

[Only one species, O. agavium.]

Lateral arms of the midcranial ridge well developed. Ocelli large. Genal
setae 4 or more. One or more disc-like structures present dorsally on each
side of the midcranial ridge. Mouth tubercle absent. The body more than
twice as long as the antennae. Antespiracular ventral setae absent.
Scutellum shorter than scutum. Hamulohalterae and suspensorial sclerites
well developed. The body more than twice as long as the hind leg. Pleural
setae of abdominal segment VIII include one seta about twice as long as
others. Style sinuated in lateral view . . . GOSSYPARIA (p. 182)

[Only one species, G. spuria.]

ERIOCOCCUS

Key to Species

Ventral part of the preocular ridge reduced. Cranial apophysis truncate. With
12 or more dorsal head setae ; setae of the ocular sclerites and the setae
between the ventral eyes present. Genal setae more than 7. Antennae 9-
segmented ; 3rd segment 2-5 or more times as long as the terminal ; capitate
sensory setae only present on the terminal segment. Lateropleurite with a
distinct membranous area. Prescutal setae 6 or more ; mesopisternal,
postmesostigmatal and basisternal setae present. Metatergal setae 4 or
more ; metapleural and postmetastigmatal setae present. Tergite of
abdominal segment VIII anteriorly bounded by differentiated heavy
sclerotization. The body 7-7 or more times as long as the tail setae of the
glandular pouches ........ E. buxi (p. 175)

Preocular ridge well developed. Cranial apophysis bifurcate. With less
than 12 dorsal head setae ; setae of the ocular sclerite and the setae between
the ventral eyes absent. Genal setae less than 7. Antennae 10-segmented ;
3rd segment less than i-8 times as long as the terminal ; capitate sensory
setae present at least on the three terminal segments. Lateropleurite
without membranous areas. Prescutal setae 2 or less ; mesepisternal,
postmesostigmatal and basisternal setae absent. Metatergal setae 3 or less ;
metapleural and postmetastigmatal setae absent. Tergite of abdominal
segment VIII without anterior heavy sclerotization. The body usually
less than 7-7 times as long as the tail setae of the glandular pouches . . 2

OF PSEUDOCOCCIDAE & ERIOCOCCIDAE 203

(1) Comparatively small species, less than 1000 [i long. Dorsal and ventral
simple eyes subequal in diameter ; the former separated by 45 or more
times their diameter. Structures of obscure homology absent on the head.
Third antennal segment less than 34 times long as wide ; terminal segment
only with 3 antennal bristles. Scutellum slightly shorter than scutum.
With a minute lateropleurital seta. Total length of the body usually more
than 21 times the length of the hind legs. Sternites of abdominal segment
VIII laterally bounded with longitudinal heavy sclerotization. Genital
capsule 2-6 or more times long as wide, and the body less than 79 times its
length; style rather straight in lateral view . . . E. orariensis (p. 171)

Comparatively large species, more than 1000 (i long. The dorsal eyes larger
than the ventral, and separated by less than 35 times their diameter. A
body of unknown homology present on each side of the midcranial ridge.
Third antennal segment 34 or more times long as wide ; terminal segment
with 3 large and 2 smaller antennal bristles. Scutellum longer than scutum.
Lateropleurital setae absent. Total body length less than 21 times the
length of the hind legs. Sternites of abdominal segment VIII without
differentiated heavy sclerotization. Genital capsule twice or less times as
long as wide, and the body 7-9 or more times its length ; style sinuating in
lateral view . . . . . . . . E. araucariae (p. 167)

Brachypterous Males
GOSSYPARIA

Key to Species

Capitate sensory setae entirely absent. Usually with 7 or more metatergal setae. The

fore and the middle legs subequal in length . . . G. salicicola (p. 190)

Capitate sensory setae present on the antennae. Usually with less than 7 metatergal

setae. The middle legs shortest ...... G. spuria (p. 186)

Apterous Males

Only two species whose apterous male forms were studied, Saccharicoccus sacchari
and Pseitdochermes fraxini. While the macropterous males of the former were
also studied, only the apterous form of the latter was available which is obviously
degenerate. The comparison therefore would not serve any useful purpose.

LETTERING USED IN TEXT-FIGURES

(A) General dorso-ventral view of the body. (F) Hind leg.

(B) Front view of the head. (G) Hind claw and distal end of tarsus
(B x ) Lateral view of the head. showing the tarsal digitules (if

(C) Terminal 3 antennal segments showing present) and the claw digitules.

the antennal bristles, few sensory setae, (H) Lateral view of the genital segment,

one fleshy and one hair-like setae. (Hi) Ventral view of the genital segment.

(D) Mesoprephragma (anterior view) and (I) Tentorium and cranial apophysis,
mesopostphragma (posterior view). (X) Structures of obscure homology.

(E) Disc pores.

204

MORPHOLOGY AND TAXONOMY OF ADULT MALES

LIST OF ABBREVIATIONS

a

ab

addr

adp

ads

aed

al

als

amsp

amss

an

anp

ap

apar

app

aps

as

asc

asdp

asds

ase

asvs

at

ata

avp

avs

axi

ax2

ax 3

axe

bas
bra
brps
bs

ca
ccx

cl
ex
cam.s

dhp

dhs

dmcr

dmep

dos

dse

eprri2
epm 3

finger-like apodeme
antennal bristles
additional ridge
abdominal dorsal pores
abdominal dorsal setae
aedeagus
alar lobe
alar setae

anterior metasternal pores
anterior metasternal setae
anus

anterior notal wing process
articular process
anterior postalar ridge
abdominal pleural pores
abdominal pleural setae
abdominal sternite
additional sclerite
antespiracular dorsal pores
antespiracular dorsal setae
differentiated apical seta
antespiracular ventral setae
abdominal tergite
anterior tentorial arms
abdominal ventral pores
abdominal ventral setae
first axillary sclerite
second axillary sclerite
third axillary sclerite
axillary cord

basalare

basal rod of aedeagus
basal ridge of penial sheath
sensilla basiconica

cranial apophysis

costal complex of wing veins

claw

coxa

campaniform sensillum

dorsal head pores

dorsal head setae

dorsal arm of midcranial ridge

dorsomedial part of epicranium

dorsal ocular setae

dorsal simple eyes

mesepimeron
metepimeron

eps 2

mesepisternum

eps2S

mesepisternal setae

eps 3

metepisternum

eps 3 s

postmetastigmatal setae

f

furca

fern

femur

fp

furcal pit

fs

fleshy setae

g

gena

gc

genital capsule

gls

setae of glandular pouch

gp

glandular pouch

gs

genal setae

gts

setae of genital segment

h

hamulohaltera

hr

hamulohalteral ridge

hs

hair-like setae

iga

internal genital aperture

ior

interocular ridge

lmcr

lateral arm of midcranial ridge

lpl

lateropleurite

lpls

lateropleurital setae

lpp

lateral pronotal pores

Ips

lateral pronotal setae

mcr

midcranial ridge

med

media

mo

mouth opening

mpnp

medial pronotal pores

mpns

medial pronotal setae

mps

metapleural setae

mr

marginal ridge

mt

mouth tubercle

mtp

metatergal pores

mts

metatergal setae

o

ocellus

ocs

ocular sclerite

ocse

setae of ocular sclerite

ost

ostiole

pa

postalare

pcr2

precoxal ridge of mesothorax

pcr3

precoxal ridge of metathorax

pdc

pedicel

pepcv

proepisternum + cervical sclerite

OF PSEUDOCOCCIDAE & ERIOCOCCIDAF

205

phri

mesoprephragma

phr 2

mesopostphragma

plai

propleural apophysis

pla 2

mesopleural apophysis

pla 3

vestigial metapleural apophysis

plri

propleural ridge

plr 2

mesopleural ridge

ph 3

metapleural ridge

pmp

postmesostigmatal pores

pms

postmesostigmatal setae

pmsp

posterior metasternal pores

pmss

posterior metasternal setae

pn 3

metapostnotal sclerite

pna

postnotal apophysis

pnp

posterior notal wing process

pn 3 r

metapostnotal ridge

pocr

postocular ridge

por

postoccipital ridge

ppar

posterior postalar ridge

P r

projection of basal ridge

pra

prealare

prar

prealar ridge

prn

lateral pronotal sclerite

prnr

pronotal ridge

pro

process of penial sheath

procr

preocular ridge

pror

preoral ridge

pros

setal sensilla of process of penial

sheath

prsc

prescutum

ps

penial sheath

pscr

prescutal ridge

pscs

prescutal suture

pscse

prescutal setae

Pt

post-tergite

pta

posterior tentorial arm

ptdp

post-tergital pores

ptp

posterior tentorial pit

pts

post-tergital setae

pwp2

mesopleural wing process

pwp3

vestigial metapleural wing

process

rad

radius

rd

posterior marginal fold of notum

sa

subalare

scl

scutellum

sclr

scutellar ridge

scls

scutellar setae

scp

scape

set

scutum

sets

scutoscutellar suture

sctse

scutal setae

sens.

circular sensoria

ser

subepisternal ridge

set. ca

capitate sensory setae

set. scla

subapical sensory setae

sp2

mesothoracic spiracle

SP3

metathoracic spiracle

spl

sensillum placodeum

S P 2 P

mesospiracular pores

SP3P

metaspiracular pores

ss

suspensorial sclerite

St

style

sta

metasternal apophysis

stni

presternum

stn 2

mesosternum (basisternum)

stnip

prosternal pores

stnxr

prosternal ridge

stnis

prosternal setae

stn 2 s

basisternal setae

sts

minute sensilla of style

tar

tarsus

tb

tentorial bridge

tdgt

tarsal digitule

teg

tegula

tegs

tegular setae

tib

tibia

tn

trochantin

tp

triangular plate

tr

trochanter

ts

tail setae

udgt

ungual digitules

vc

ventral cavity

vhp

ventral head pores

vhs

ventral head setae

vmcr

ventral arm of midcranial ridge

vos

ventral ocular setae

vprd

ventral preocular depression

vse

ventral simple eyes

structure of obscure homology

206 MORPHOLOGY AND TAXONOMY OF ADULT MALES

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* Paper not seen and only known to me from other references.

[Index overleaf]

INDEX

Centrococcus, 159
insolitus, 159

Ceroputo, 155
pilosellae, 155

Dysmicoccus, 139
alazon, 139

Eriococcus, 167
araucariae, 167
buxi, 175
orariensis, 171

Ferrisiana, 122
virgata, 122

Gossyparia, 182
salicicola, 190
spuria, 182

Maconellicoccus, nl
hirsutus, 118

Nairobia, 163
bifrons, 163

Nipaecoccus, no
nipae, 114
vastator, no

Octococcus, 151
africanus, 151

Ovaticoccus, 179
agavium, 179

Pseudochermes, 193

fraxini, 193
Planococcoides, 106

ireneus, 106
Planococcus, 93

citri, 93

dioscoreae, 102

kenyae, 98
Pseudococcus, 131

citriculus, 135

obscurus, 131

Saccharicoccus, 143
sacchari, 143

Trionymus, 126
newsteadi. 126

— <^A>

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5. Ahmad, I. The Leptocorisinae (Heteroptera : Alydidae) of the World.
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7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family
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8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the
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9. Hemming, A. F. The Generic Names of the Butterflies and their type-species
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10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera : Rho-
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12. Watson, A. The Taxonomy of the Drepaninae represented in China, with
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