Kozar Et Al - Acanthococcidae and Related Families of The Palaearctic Region

ACANTHOCOCCIDAE
AND RELATED FAMILIES

OF THE PALAEARCTIC REGION
In Memory of Jan Koteja (1932-2004), the revolutionary coccidologist of
all times.
ACANTHOCOCCIDAE
AND RELATED FAMILIES
OF THE PALAEARCTIC
REGION
FERENC KOZÁR*
M. BORA KAYDAN**,
ZSUZSANNA KONCZNÉ BENEDICTY*
ÉVA SZITA*
*Plant Protection Institute, Centre for Agricultural Research,

Hungarian Academy of Sciences, Budapest, Hungary
** Çukurova Üniversity, Imamoglu Vocational School, Adana, Turkey
Plant Protection Institute, Centre for Agricultural Research,

Hungarian Academy of Sciences
Budapest, Hungary
2013
Copyright © 2013 by Plant Protection Institute, Centre for Agricultural Research,
Hungarian Academy of Sciences
All rights reserved.
No part of this book may be reproduced in any form or by any means without permission
in writing from the author and publisher, except that the scientific data herein is a part
of the public domain.
Technical editor: Éva Szita
ISBN: 978-963-508-678-8
Printed by Akaprint Nyomdaipari Kft., Budapest, Hungary

FERENC KOZÁR
1943-2013
TABLE OF CONTENTS
INTRODUCTION...................................................................................................................9
MORPHOLOGICAL CHARACTERS OF THE ACANTHOCOCCIDAE FAMILY
GROUP.................................................................................................................................11
PHYLOGENY...........................................................................................................................24
COLLECTING AND MOUNTING METHODS.............................................................52
DEPOSITORIES OF TYPES AND STUDIED INSECTS.............................................53
ACKNOWLEDGEMENT.......................................................................................................55
SYSTEMATIC PART.............................................................................................................56
ACANTHOCOCCIDAE FAMILY GROUP Signoret, 1875 sensu Koteja, 1974......57
ACANTHOCOCCIDAE Signoret, 1875.............................................................................59
Acalyptococcus Lambdin & Kosztarab, 1977........................................................................64
Acanthococcus Signoret, 1875..................................................................................................72
Anophococcus Balachowsky, 1954 .......................................................................................180
Borchseniococcus Kaydan & Kozár, 2008...........................................................................257
Eriokermes Miller & Miller, 1993...........................................................................................260
Gossyparia Signoret, 1875.......................................................................................................261
Gossypariella Borchsenius, 1960 ...........................................................................................269
Greenisca Borchsenius, 1948..................................................................................................278
Greenoripersia Bodenheimer, 1929.......................................................................................295
Gregoporia Danzig, 1979........................................................................................................295
Hispaniococcus Kozár, 2008 .................................................................................................300
Hujinlinococcus Kozár & Wu Genus nova..........................................................................307
Kaweckia Koteja & Zak-Ogaza, 1981...................................................................................310
Kotejacoccus Kaydan & Kozár, 2008....................................................................................324
Neoacanthococcus Borchsenius, 1948..................................................................................334
Neokaweckia Tang & Hao, 1995............................................................................................351
Neotrichococcus Miller & Gimpel, 1999..............................................................................357
Noteococcus Hoy, 1962 . ........................................................................................................364
Orontesicoccus Kaydan Genus nova.....................................................................................368
Ovaticoccus Kloet, 1944 ........................................................................................................373
Proteriococcus Borchsenius, 1960..........................................................................................383
Pseudoacanthococcus Kaydan & Kozár Genus nova.........................................................386
Rhizococcus Signoret, 1875 Reestablishment in sense of Koteja (1974)........................390
Uhleria Cooke, 1881.................................................................................................................579
Balticococcus Koteja, 1988......................................................................................................587
Gedanicoccus Koteja, 1988.....................................................................................................589
Jutlandicoccus Koteja, 1988 ...................................................................................................590
Kuenowicoccus Koteja, 1988..................................................................................................592
8 Acanthococcidae and related families
CRYPTOCOCCIDAE Kosztarab, 1967.............................................................................594

Cryptococcus Douglas, 1890..................................................................................................595
Pseudochermes Nitsche, 1895................................................................................................608
ERIOCOCCIDAE Signoret, 1875.......................................................................................616
Aculeococcus Lepage, 1941....................................................................................................618
Asiacornococcus Tang & Hao, 1995 ....................................................................................621
Eriococcus Targioni Tozzetti, 1868........................................................................................629
KUWANINIDAE Kozár Family nova................................................................................639
Kuwanina Cockerell, 1903 . ....................................................................................................640
REFERENCES......................................................................................................................647
TAXONOMIC INDEX........................................................................................................666
HOST PLANT INDEX.......................................................................................................670
DISTRIBUTION INDEX..................................................................................................677
Introduction 9
INTRODUCTION
This work on the FAMILY GROUP “ACANTHOCOCCIDAE” deals with 4 families

found in the Palaearctic region. These have been treated in different ways by those who
have worked on “Eriococcidae” sensu lato but here we partly follow the ideas of Koteja
(1974a).
Family Acanthococcidae Signoret, 1875 (Acanthococcinae sensu (Koteja 1974a)
is treated here as a family): 22 valid genera: Acalyptococcus, Acanthococcus, Anophococcus,
Borchseniococcus, Gossyparia, Gossypariella, Greenisca, Gregoporia, Hispaniococcus, Huajinlinococcus,
Kaweckia, Kotejacoccus, Neoacanthococcus, Neokaweckia, Neotrichococcus, Noteococcus, Orontesicoccus,
Ovaticoccus, Proteriococcus, Pseudoacanthococcus, Rhizococcus and Uhleria.
Family Cryptococcidae Kosztarab, 1968: 2 genera: Cryptococcus and Pseudochermes.
Family Eriococcidae Cockerell, 1903 (Eriococcinae sensu (Koteja 1974a) is treated
here as a family): 3 genera: Aculeococcus, Asiacornococcus and Eriococcus sensu stricto
Family Kuwaninidae Kozár, fam. nov., 1 genus: Kuwanina.
On the other hand Koteja (1974a) treated several more groups in the “Acanthococcidae
Family Group” (Dactylopiidae, Apiomorphidae, Kermesiidae, Cerococcidae,
Calycicoccidae in his study, but here we are treating only four families which were
treated under the Eriococcidae sensu lato by Miller & Gimpel (2000) (except two families
Apiomorphidae and Calycicoccidae which are not present in the Palaearctic Region).
A further 16 species synonyms are listed. In addition, 3 new genera, 11 new species
and 1 reestablished genus and 1 reestablished species names are described in this work,
plus 52 species are redescribed and 80 new species combinations are proposed. The
present revision therefore covers a total of 188 (plus 6 fossil) species in 28 (plus 4 fossil)
genera.
Nikolaj Sergeevits Borchsenius (1949) published the last comprehensive work on
the Eriococcinae (now the Acanthococcidae FG) in the Palaearctic Region in his book
of Pseudococcidae sensu lato. The present study continues the work started by Signoret
(1875), Borchsenius (1949), Ferris (1955), Hoy (l962, 1963), Koteja (1974a, b), Kosztarab
& Kozár (1978, 1988), Danzig (1980), Gullan (1984), Kozár & Walter (1985), Miller &
Miller (1992, 1993), Tang & Hao (1995), Miller & Gimpel (2000), Cook & Gullan (2004),
Kozár (2009), Hodgson & Miller (2010) and others. The Acanthococcidae family group
(FG) is one of the peculiar branches of the Coccoidea (Hemiptera) (Downie & Gullan
2004, Koteja 1974a, b) and deserves detailed study. The subfamily was raised to family
status by Afifi (1968) as a result of his studies on the adult males. Here, we follow and test
the generic concepts of Signoret (1875), Borchsenius (1949), Kosztarab & Kozár (1988),
Tang & Hao (1995), and Köhler (1998) as working hypotheses.
The scale insects include many important pest species, several of which are regarded
as amongst the most destructive insect pests and this is also true for the Acanthococcidae
FG. They are very often intercepted by quarantine services but their identification is
difficult and needs to be done by specialists.
The literature listed in the systematic part of this publication includes the
following: original descriptions, new generic combinations, redescriptions, revisions
and monographs. Because of space limitations, the whole reference list could not be
included, but these are available in the Catalogue of Miller & Gimpel (2000), and in
ScaleNet (Miller et al., 2013, last updated in 13 June 2013).
The families, tribes, genera and species included below are presented in alphabetical
order. All invalid taxa, nomina dubia, nomina nuda, misidentifications, misspellings (with
some exceptions) have been omitted from the text as suggested by the International
Code of Zoological Nomenclature (Ride et al., 2008)
The authors are fully aware that this work is not free from inaccuracies and regret
that they have been unable to solve many of the problems and to meet everyone's
expectations. Colleagues are kindly requested to send remarks and modifications to the
authors in order to enable them to make the necessary modifications.
Introduction 11
MORPHOLOGICAL CHARACTERS
OF THE ACANTHOCOCCIDAE FAMILY GROUP
Unmounted adult female

Body of adult female in life generally elongate, but oval or rotund in Cryptococcidae and
Kuwaninidae; anal lobes well-developed, long and generally slightly to heavily sclerotized;
these occasionally undeveloped (e.g. in Cryptococcidae and Kuwaninidae). Body usually
covered with a felted wax ovisac, but median part of dorsum sometimes bare, not covered
by wax (Fig. 1).
Mounted adult female

Body elongated, mostly 0.5–3.75 mm long, with well-developed, long, sometimes
sclerotized anal lobes, but anal lobes occasionally undeveloped, and body oval or rotund
as in Cryptococcidae and Kuwaninidae.
Antennae with 1-7 segments (Fig. 2), with 3-7 in Acanthococcidae and Eriococcidae
s.s. but only 1-4 in Cryptococcidae and Kuwaninidae. Setae on antennae mainly hair-like
but apical and subapical segments with strong, blunt, falcate, sensory setae. Sensory pore
always present on the second segment.
Eyes present.
Labium 1-3 segmented (Koteja 1974a, b) (generally 3), elongate (Fig. 3), very variable
in size in different genera, usually twice as long as wide but occasionally as wide as long.
Setae usually hair-like. Remains of basal segment partly fused with median segment, with
two pairs in Acanthococcidae and Cryptococcidae but only one pair in Eriococcidae s.s.;
basal segment in Kuwaninidae undeveloped.
Stylet loop very long on species living on woody plants, but somewhat shorter in
species living on herbaceous plants, when 2-4 times longer than labium and usually only
reaching the median coxae. On the other hand labium very short in genera and species
living on grasses except, R. greeni Newstead, which is currently known on Poaceae but has
a long stylet loop, which suggests a more recent switch in host plant:s.
Thoracic spiracles usually small, mostly without associated pores but with a loose
group of loculate pores around the spiracles in a few species and loculate pores even
occasionally present in atrium.
Legs well developed in Acanthococcidae and Eriococcidae, absent in some species
of Cryptococcidae and Kuwaninidae. When legs present, coxae normally developed
and posterior coxae often with pores and spinulae; spinulae also sometimes present on
median and anterior coxae. Trochanter-femur and tibia-tarsus segmentation distinct.
Tibia normally as long as tarsus (Fig. 4), but in some groups (e.g. Acalyptococcus, Eriococcus)
the ratio is different, with tibia shorter than tarsus. Claw long, elongated, with two setose
or hair-like digitules, usually longer than claw and knobbed, sometimes spine-like (see in
Fig. 4), claw usually with a denticle; legs with rows of spine-like or hair-like setae, those
on femur usually hair-like, but some on tarsus spine-like (e.g., Acalyptococcus, Acanthococcus,
Anophococcus, Asiacornicoccus, Greenisca, Neoacanthococcus, Rhizococcus). Trochanter with 2
sensory pores on each surface.
Pores. Pores on body surface of various types: small simple pores, cruciform pores,
three-, four- or five-locular pores and multilocular (6–11 locular) pores (Fig. 5); pore
microstructure variable between taxa.
Tubular ducts, if present, very variable. The microtubular ducts may be long and
narrow, with a bifurcate orifice, or short. Macrotubular ducts sometimes of three sizes.
In Eriococcidae s.s. (e.g., E. buxi), extra long (enlarged), wide macrotubular ducts present
on dorsal margin (Figs 6, 7).
Dorsum often covered with enlarged spine-like setae of various sizes and shapes.
Submargin of venter also often with enlarged spine-like setae. Only hair-like setae present
on midventer, rarely some of them may be capitate (Kotejacaoccus turcicus, R. matasovae, R.
saikwanensis).
Most genera have a pair of well-developed anal lobes but in the Cryptococcidae and
Kuwaninidae, and in the genera Borchseniococcus, Hispaniococcus and Ovaticoccus, the anal
lobes are degenerate or absent. Anal lobes often heavily sclerotised, with strong spines, or
setae on dorsal surface, normally three in number, similar in shape to marginal setae (Fig.
9). Each anal lobe often terminated by a long apical anal lobe seta, usually longer than
lobe. Ventral surface of each lobe with one or more shorter subapical setae. However,
shape and size of spines and setae highly variable between species and genera. Anal lobes
in some genera and species with normally a more or less sclerotised cauda (median plate
of some authors).
Anal ring usually well developed, with one or two rows of pores and 3 or 4 (rarely up
to 8) pairs of anal ring setae. Setae usually much longer than diameter of anal ring (Fig.
10). In some genera, anal ring with only one row of pores. A pair of suranal setae present
on either side of anal ring, each normally hair-like in species found in the Palaearctic
Region. Anal ring reduced in some genera, without pores but sometimes anal ring
modified, strongly sclerotised and with spine-like setae. Pores in outer row sometimes
with hardly visible spiculae, which are often not mentioned by authors. Anal ring mostly
on dorsum, only in a few groups on venter or on body margin.
Introduction 13
Figure 1 a-d. a – Gossyparia spurious, b – Rhizococcus coronillae male test,

c – Rhizococcus thymi, d – Rhizococcus thymi adult female.(Photo: G. Pellizzari and T.
Kondo).
Figure 1 e-h. e – Eriococcus sp., f – Ovaticoccus agavium, g – Ovaticoccus agavium male

test, h – Rhizococcus araucariae first instar larvae (Photo: G. Pellizzari and T. Kondo).
Introduction 15
Figure 2. Antennae of Acanthococcidae family group: a – Acanthococcus aceris,

b – Acanthococcus onukii, c – Acanthococcus corniculatus, d – Aculeococcus
yongpingensis, e – Anophococcus formicicola, f – Asiacornicoccus kaki,
g – Hispaniococcus agenjoi, h – Rhizococcus acutus, i – Rhizococcus brevenniae, j –
Cryptococcus aceris, k – Eriococcus buxi, l – Kuwanina betulae, m – Kuwanina parva.
Figure 3. Labium of Acanthococcidae family group. a – Acanthococcus aceris, b –

Acanthococcus onukii, c – Acanthococcus spiraeae, d – Anophococcus insignis, e
– Anophococcus oxyacanthus, f – Hispaniococcus agenjoi, g – Kaweckia glyceryae,
h – Kaweckia vanensis, i – Neoacanthococcus gracilispinus, h – Neoacanthococcus
tamaricicola, j – Rhizococcus astragali, l – Cryptococcus aceris, m – Eriococcus buxi, n –
Kuwanina betulae.
Introduction 17
Figure 4. Legs of Acanthococcidae family group. a – Acalyptococcus deformans, b –

Acanthococcus aceris, c – Anophococcus kotejai, d – Anophococcus oxyacanthus, e
– Asiacornicoccus kaki, f – Greenisca erwini, g – Neoacanthococcus gracilispinus, h –
Neoacanthococcus tamaricicola, i – Rhizococcus ammophilus, j – Rhizococcus astragali,
k – Eriococcus buxi, l – Kuwanina betulae.
Figure 5. Varieties of pores of Acanthococcidae family group, a – Anophococcus formicicola,

b – Borchseniococcus duzgunesae, c – Gossypariella juniperi, d – Gregoporia istriensis,
e – Kaweckia orientalis, f – Neoacanthococcus gracilispinus, g – Rhizococcus thaleri, h –
Uhleria marianae, i – Acanthococcus altaicus.
Introduction 19
Figure 7. Macrotubular ducts of Acanthococcidae family group: a – Acanthococcus

aceris, b – Neoacanthococcus gracilispinus, c – Anophococcus oxyacanthus,
d – Rhizococcus variabilis, e – Cryptococcus aceris, f – Pseudochermes williamsi, g –
Eriococcus buxi, h – Kuwanina betulae.
<<
Figure 6. Different kinds of micro tubular ducts of Acanthococcidae family group: a –
Acanthococcus aceris, b – Anophococcus formicicola, c – Gossypariella siamensis, d –
Hispaniococcus agenjoi, e – Kaweckia orientalis, f – Neoacanthococcus tamaricicola, g –
Neotrichococcus filifer, h – Uhleria araucariae, i – Uhleria mariannae, j – Pseudochermes
williamsi, k – Eriococcus buxi, l – Kuwanina parva.
Figure 8. Different kinds of enlarged setae of Acanthococcidae family group: a –

Acalyptococcus deformans, b – Acanthococcus aceris, c – Acanthococcus corniculatus,
d – Acanthococcus salicis, e – Acanthococcus uvaeursi, f – Anaphococcus oxyacanthus,
g – Asiacornicoccus kaki, h – Gossypariella juniperi, i – Gossypariella siamensis, j –
Gossyparia spuria, k – Greenisca erwini, l – Hispaniococcus agenjoi, m – Hujinlinococcus
nematospherus n – Kaweckia glyceryae, o – Kotejacoccus turcicus, p – Neokaweckia
laeticornis, q – Neotrichococcus filiferus, r – Ovaticoccus agavium, s– Rhizococcus acutus,
t – Rhizococcus coccineus, u – Rhizococcus palustris, v – Rhizococcus thaleri, w – Uhleria
araucaria, x – Uhleria mariannae, y – Eriococcus buxi.
Introduction 21
Figure 9. Different kinds of anal lobes of Acanthococcidae family group: a – Acanthococcus

aceris, b – Acanthococcus corniculatus, c – Acanthococcus spireae, d – Anophococcus
kotejai, e – Gossypariella siamensis, f – Kaweckia glyceryae, g – Kaweckia hellenica,
h – Neokaweckia laeticornis, i – Neokaweckia rubra, j – Noteococcus hoheriae, k –
Orentesicoccus lauri, l – Rhizococcus astragali, m – Rhizococcus canthium, n – Uhleria
mariannae, o – Eriococcus buxi.
Figure 10. Different kinds of anal rings of Acanthococcidae family group: a – Acanthococcus
aceris, b – Acanthococcus onukii, c – Anophococcus kotejai, d – Eriococcus buxi, e –
Greenisca erwini, f – Kaweckia vanensis, g – Kotejacoccus turcicus, h – Neoacanthococcus
gracilispinus, i – Neokaweckia rubra, j – Neotrichococcus filiferus, k – Rhizococcus
astragali, l – Rhizococcus thaleri, m – Cryptococcus aceris, n – Pseudochermes williamsi,
o – Ovaticoccus agavium, p – Kuwanina betulae.
Introduction 23
Morphology of the slide-mounted male

The male is known only in a few species of the family group Acanthococcidae (Hodgson
& Miller, 2010). Some other important literature on male morphology includes:
Acanthococcus (Fig. 53), Eriococcus, Gossyparia, Ovaticoccus and Pseudochermes (Afifi, 1968),
Calycicoccus (Gullan et al., 2006); Eriochiton (Henderson & Hodgson, 1995); Eriococcus
(Miller et al. 1992, Hodgson 2005); Pseudotectococcus (Hodgson et al., 2004); Kuenowicoccus
(Koteja, 1988a); Pseudomontanococcus (Kozár et al., 2008); Stibococcus (Miller & González,
1975), Apiomorpha and Opisthoscelis (Theron, 1968), and Cryptococcus (Wu, 2000).
The adult males of the Acanthococcidae FG in the Palaearctic differ from the other
non-acanthococcid clades of Cook & Gullan (Hodgson, in prep.). They can be diagnosed
by the following combination of character-states: mostly winged (macropterous)
although a few are wingless (apterous) or larvoid; antennae mostly with 10 segments;
postoccipital ridge strong and laterally forked; two pairs of simple eyes present; scutum
without a median membranous area; scutellum without a foramen; fleshy setae, when
present, almost entirely restricted to antennae and legs, and always much longer than
broad; legs well developed; claw broad at base, abruptly tapering apically; tarsus generally
two segmented; both claw and tarsal digitules with small capitate apices; tibia with 2
tibial spurs; each trochanter with 3 campaniform pores on each side, not in a straight
line; hamulohalteres present or absent; alar setae generally present; glandular pouches
generally present; penial sheath short, usually about as long as basal width; anus large, in
middle of dorsal surface of penial sheath; aedeagus and associated structures sometimes
complex (Fig. 54). A detailed description and drawing of the male of Acanthococcus
melnikensis (Hodgson & Trencheva, 2008) is presented under species description.
The studied winged (macropterous) males in the Region are: Acanthococcus melnikensis,
Anophococcus pseudinsignis, Eriococcus buxi, Gossyparia spuria, Ovaticoccus agavium, Rhizococcus
munroi, Uhleria araucaria, all with 9 or 10-segmented antennae some with brachypterous
males (Gossyparia spuria, G. salicicola). Wingless (apterous) forms have not been found in
the studied region, but the wingless Pseudomontanus martini, with 9-segmented antennae,
was found in New Guinea. The known larvoid males, such as Cryptococcus ulmi and
Pseudochermes fraxini, have 7- or 8-segmented antennae. Although the male is known
only for a few apterous species, a clear trend for a reduction in the number of antennal
segments can be observed.
Immature stages
The morphology of the immature stages has not been well studied in the
Acanthococcidae FG. In those that have, the morphology of the first- and second-
instar nymphs are quite distinctive (Hoy, 1962) but these stages need further study. In
the Palaearctic region, drawings of first- and second-instar nymph are sometimes added
to the female figures in some publications (Tereznikova, 1981). Some features of the
adult female and first-instar nymph from different genera and species are shown in
Fig. 11. They show considerable variation in the shape and distribution of the setae.
In Acanthococcus and some other genera living on woody plants (Gossyparia, Gossypariella,
Uhleria, Cryptococcus, Pseudochermes, Kuwanina, etc.), the enlarged spinose setae on the
margin and mid-dorsum of the adult female and nymph are quite similar. In the grass-
feeding group, the enlarged setae on the margin are often absent or, if present, are quite
different to the dorsal setae. Thus the structure of the nymphs provides good support
for generic and species diagnosis and sometimes supports differences between species
groups. Species living on herbaceous plants have similarly-shaped marginal and dorsal
enlarged setae on the female and on the nymph but the dorsal setae are usually much
shorter than the marginal setae in both stages. The great variation in the shape and size
of the dorsal setae clearly illustrates the diversity of the taxonomic structure within the
Acanthococcidae FG. On the other hand first-instar nymphs have few enlarged setae and
few microtubular ducts on the dorsum, few cruciform pores on the venter and have six-
segmented antenna.
Second-instar female nymphs have about twice as many enlarged setae and
microtubular ducts on the dorsum and many more cruciform pores on the venter. The
second-instar male nymphs have numerous macrotubular ducts on the dorsum and venter
(absent in second-instar females) and 7 segmented antennae (generally 6 segmented on
second-instar female nymphs). Second-instar females and second-instar males has been
described for Acanthococcus melnikensis, Anophococcus pannonicus, Eriococcus buxi, Kaweckia
glyceriae, Rhizococcus reynei, etc. (Hodgson & Trencheva, 2008; Williams, 1985a; and in the
present work).
In general, the morphological features of the nymph support the generic diagnoses of
the families in the Acanthococcidae FG within the Palaearctic Region. Further, detailed
studies of the morphology of all stages of each taxon could give more support for family
and generic separation and, in some cases, it could help even in species identification.
PHYLOGENY
Although, In “A Systematic Catalogue of the Eriococcidae (Felt Scales) (Hemiptera:

Coccoidea) of the World”, Miller & Gimpel (2000) reported 554 species in Eriococcidae
s.l., a total 668 species are listed in the family Eriococcidae s.l. in ScaleNet (Miller et al.,
2013), of which 346 were treated under the generic name “Eriococcus”. This is unnaturally
large for a genus. This pattern is also repeated in the Palaearctic region which has been
relatively well studied and which currently has 175 species in the Eriococcidae sensu lato,
of which 155 are regarded as “Eriococcus” species. In order to divide this family into more
natural groups, further studies will be needed, particularly on the morphology of males
and immature stages, and on their biology and ecological parameters, plus more analyses
using molecular data. We here consider that it is more useful to use the generic and family
names in a narrower sense under the general heading of Acanthococcidae FG.
Introduction 25
Figure 11. Variation of dorsal and marginal enlarged setae of females and first instar larvae
in different families, genera and species of Acanthococcidae family group.
The Acanthococcidae FG is considered sister to all other neococcids and the

pseudococcids (Foldi, 1997; Koteja, 1974a; Miller & Gimpel, 2000). The roots of this
family group may lie under the ice of the Antarctic (Hoy, 1962; Kozár, 2009). The study
of the phylogenetic relations of the family group is complicated, and quite problematic.
Cox & Williams (1987) indicated that the “Eriococcidae” s.l. was paraphyletic. They
pointed out that the “Eriococcidae” is separated from other families of Coccoidea
by: the absence of circuli, ostioles and trilocular pores from the Pseudococcidae; by the
absence of anal plates from the Coccidae; by the absence of a single triangular anal plate
from the Aclerdidae, and by the absence of cribriform plates from the Lecanodiaspididae
and Cerococcidae. In other words, the Eriococcidae s.l. appears to be diagnosed by
the absence of structures rather than by synapomorphies. On the other hand, the most
cited characteristic of the family is the shape of the macrotubular ducts which normally
have the inner end reflexed in the form of a cup (Figure 7), suggesting that this kind
of duct can be found in Eriococcidae, Pseudococcidae and Coccidae. More recently,
Cook & Gullan (2001) stated that the “Eriococcidae” were paraphyletic because species
currently placed in this family share morphological character states with the Aclerdidae,
Cerococcidae, Lecanodiaspidiae, Coccidae, Kermesidae and/or Putoidae, i.e. the anal
ring structure, cup-shaped macrotubular ducts and microtubular ducts. Although one
of the most important character-states of the felt scales is the presence of a pair of
elongate and often sclerotised anal lobes, many genera regarded as “Eriococcidae” s.l. lack
these, i.e. Kuwanina Cockerell, Cryptococcus Douglas, Borchseniococcus Kaydan & Kozár, and
Kotejacoccus Kaydan & Kozár. On the other hand, species from New Zealand, Australia
and Chile display forms of anal lobes that are intermediate between “Eriococcidae” s.l.
and Coccidae (Cox & Williams, 1987).
Koteja (1974a, b) studied the labium of the Coccoidea and included several families
in his Acanthococcidae FG. He found that most acanthococcids have a three segmented
labium (except Kuwanina) but with one pair of subapical setae only, and are, therefore,
easily distinguished from the Pseudococcidae, which have two pairs of subapical setae.
On this basis, Koteja (1974a) proposed that his Acanthococcidae family group contained
the following distinct groups: Acanthococcidae, Apiomorphidae, Dactylopiidae,
Kermesidae, Cerococcidae, Cryptococcidae, the Kuwania group (as family status) and
the unplaced genus Xerococcus).
Phylogenetic relationships of scale insects based on males, including some Eriococcidae
s.l. species, were studied by Afifi (1968), who could separate the Eriococcidae from the
Pseudococcidae, within which they had previously been considered a by some authors as
subfamily. In addition, Hodgson (2002) undertook a cladistic analysis based on adult male
morphology, including 9 species of Eriococcidae, and concluded, like Cox & Williams
(1987), that the family is paraphyletic. These conclusions have been clearly confirmed by
the work of Gullan & Cook (2007). To understand this complexity, more work needs to
be done using different methodologies, including larger samples of genera and species.
Introduction 27
Morphology
Ouvrard & Kozár (2009) conducted a phylogenetic analysis based on 132 morphological
characters of the adult females of 52 Palaearctic species in 9 genera, with one
pseudococcid, one coccid and one cryptococcid out-group species (Fig. 12). They
argued that Acanthococcidae FG “under the name of Eriococcidae sensu lato” was
“paraphyletic”. In this study, they determined that the family-group is supported by five
synapomorphic characters: distribution of inter-antennal setae (in elongate group between
antennae), the presence of a tarsal campaniform pore, “base of anal lobes not fused, presence of anal
lobes and presence of spines on the anal lobes”. Gullan & Cook (2007) found that Eriococcus buxi
(Boyer de Fonscolombe) fell into a different clade from Acanthococcus clade, along with
the Stictococcidae and Beesoniidae. In this analysis (Ouvrard & Kozár, 2009), E. buxi was
sister to all other species in the Acanthococcidae FG from the palaearctic and this sister-
clade was supported by two synapomorphic characters: presence of coxae with reticulations
and anal lobe setae longer than lobe. Thus, it could be argued that their results support the
findings of Gullan & Cook (2007).
The above study included just characters from the adult females. Unfortunately the
the morphology of the nymphal stages has only been studied in a very few species. As
indicated above, the addition of morphological characters from the immature stages can
also provide very useful characters, and this was recognized by Balachowsky (1953) in the
Palaearctic region and by Hendricks & Kosztarab (1999) in the Nearctic region in their
work on the Kermesidae. It is very likely that a better understanding of the morphology
of the adult males will also greatly improve our understanding of the relationships of the
group being studied here.
Molecular phylogeny
Using molecular methods, Cook et al. (2002, 2004) and Gullan & Cook (2001, 2007)
studied the phylogeny and higher classification of the scale insects, including the
“Eriococcidae s.l.” and, as in all recent phylogenetic studies with sufficient taxon sampling,
they considered that the Eriococcidae sensu lato was paraphyletic, as first suggested by Cox
& Williams (1987) based on adult female morphology. Phylogenetic analysis using 18S
data (Cook & Gullan, 2004) recovered four groups within the “Eriococcidae” s.l.: (i) the
BSE clade, which included the Beesoniidae, Stictococcidae and eriococcids related to E.
buxi; (ii) a Gondwanan group, which contained mainly the Australian and New Zealand
taxa which mainly feed on Myrtaceae); (iii) an Acanthococcid group (which included
species of Acanthococcus Signoret, mainly from the Holarctic) and (iv) Calycicoccus merwei
Brain as an isolated lineage (Cook & Gullan, 2004; Gullan et al., 2006). This division was
supported by the molecular results of Kondo et al. (2006) for the Nearctic region (North
America) and by Gwiazdowski et al. (2006).
Figure 12. Phylogenetic tree of Acanthococcidae family group based on female morphology
(Ouvrard & Kozár, 2009).
Introduction 29
Molecular methods
Specimens used in this study are listed in Table 1. Phenacoccus aceris Signoret was used as
outgroup. All specimens were kept in 95–100% ethanol prior to DNA extraction. Before
DNA extraction all specimens were examined under the microscope for the presence
of parasitoids in order to avoid contamination. DNA was extracted from single adult
females with the DNA-easy tissue kit (Qiagen, Inc, Valencia, CA) and the procedure
given by the maker was followed, except that the cuticle of the specimens were kept for
morphological identification.
PCR products were generated from a mitochondrial gene, cytochrome oxidase I
(COI), and one nuclear gene: a fragment of the D2 and D3 regions of the large subunit
ribosomal DNA gene (28S). Primers for both amplification and sequencing were 5' -
CAA CAT TTA TTT TGA TTT TTT GG - 3' (C1-J-2183 aka Jerry) (Simon et al., 1994)
and 5' - GCW ACW ACR TAA TAK GTA TCA TG - 3' (C1-N-2568 aka BEN3R,
designed by T. R. Schultz, Smithsonian Institution) (Brady et al., 2000) for COI; and 5' –
TCG GAR GGA ACC AGC TAC TA - 3' (A335 REVERSE) (Whiting et al., 1997) and
5' - GAG AGT TMA ASA GTA CGT GAA AC - 3' (S3660 FORWARD) (Dowton &
Austin, 1998) for 28S. PCR reaction components and final concentrations were 1.5 to
2.5 mM MgCl2, 0.2 mM dNTPs, and 1 unit Taq polymerase in a proprietary buffer (PCR
Master Mix, Promega Biotechnology), 0.2 µM each primer, and 5 µl DNA template in a
final volume of 25 µl. The PCR cycling protocol for COI was 95 ºC for 7 m, followed by
40 cycles of 95 ºC for 1 m, 45 ºC for 1 m, and 72 ºC for 1 m 30 s with a final extension
at 72 ºC for 5 m. Protocol for 28S was 94 ºC for 4 m followed by 35-45 cycles of 94 ºC
for 1 m, 49-52 ºC for 1 m, and 72 ºC for 1 m, 30 s with a final extension at 72 ºC for 4 m
(Gullan et al., 2010).
PCR products were purified by exonuclease I and shrimp alkaline phosphatase
digestion of single stranded DNA (primers) and dNTP’s (ExoSAP-IT, USB Corp.,
Cleveland, OH). Products were sequenced on both strands using the ABI Big Dye
terminator sequencing reaction kit (Perkin-Elmer/ABI, Weiterstadt, Germany) on an
ABI Prism 377 automatic sequencer with XL upgrade (Perkin-Elmer) on 5 % Acryl/
Bisacryl Long Ranger gels.
Contigs were assembled using CodonCode Aligner v. 3.7.1 (CodonCode Corp.) and
multiple alignment was performed with BioEdit.
Maximum likelihood (ML) and Bayesian analyses were used to estimate trees. The
data were analysed for each locus separately and with data for the two loci combined. ML
trees were reconstructed using Mega5 pocked program, with four data partitions: 3 codon
positions for COI, and one partition for 28S. A separate GTR, nucleotide substitution
model was applied to each partition. 10.000 non-parametric bootstrap replicates were
performed using GTR, with every 100 bootstrap tree used as the starting tree for ML
optimisation with GTR. Phylogenies were also reconstructed with Bayesian inference
methods using MrBayes v. 3.1.2 (Ronquist & Huelsenbeck, 2003) under the packed
program Geneious 5.6. We applied a separate GTR model with gamma-distributed
Table 1. Collection details of specimens and outgroup taxa from which DNA was extracted.
30
Coll.
Species name Code Location Date E N Host plant: Collector
Code
Van-Hakkari Road,
Acanthacoccus sp. MBK078 4222 23.v.2008 37°41’644’’ 043°56’266’’ Quercus sp. M.B.Kaydan
Turkey
Acanthococcus roboris Trabzon-Gümüşhane M.B.Kaydan,
MBK289 4724 07.vii.2010 40°37’406’’ 039°20’204’’ Quercus sp.
Goux road, Turkey F. Kozár
Anophococcus sp. MBK220 4694 Töreki, Hungary 26.ix.2008 Agropyron sp. F. Kozár
Anophococcus herbaceus Van - Gevaş - Tatvan ,

MBK175 4542 09.vi.2009 38°23’265’’ 042°47’444’’ Poaceae M.B.Kaydan
Danzig Turkey
Anophococcus agropyri
MBK001 4636 Ecséd, Hungary 09.ix.2009 Bromus sp. F. Kozár
(Borchsenius)
Anophococcus evelinae Trabzon Sümela,
MBK274 4725 07.vii.2010 40°4’167’’ 039°39’511’’ Melica sp. M.B.Kaydan
Kozár Turkey
Anophococcus insignis
MBK219 4695 Ecséd, Hungary 09.ix.2008 Bromus sp. F. Kozár
Newstead
Anophococcus formicicola
MBK026 4661 Úri, Hungary 31.v.2005 Cynodon dactylon F. Kozár
Newstead
Borchseniococcus duzgunesae Iğdır-Tuzluca-Gaziler
MBK066 4388 12.vi.2008 40°06’717’’ 043°34’183’’ Panderia pilosa M.B.Kaydan
Kaydan & Kozár road, Turkey
Gossyparia spuria Iğdır-Tuzluca road,
MBK032 1391 03.v.2005 40°02’611’’ 043°39’707’’ Ulmus sp. M.B.Kaydan
(Modeer) Turkey
Acanthococcidae and related families
Coll.
Code
Gossyparia spuria
MBK016 4651 Gárdony, Hungary 15.v.2005 Ulmus sp. M.B.Kaydan
Introduction
(Modeer)
Gossyparia spuria
MBK042 1619 Ağrı-Tutak, Turkey 01.vi.2005 39°32’698’’ 042°45’943’’ Ulmus sp. M.B.Kaydan
(Modeer)
Gossyparia spuria
MBK035 1425 Iğdır-Aralık, Turkey 02.v.2005 39°52’410’’ 044°30’997’’ Ulmus sp. M.B.Kaydan
(Modeer)
Greenisca sp. MBK002 4637 Ecséd, Hungary 18.vii.2009 Brachypodium sp. F. Kozár
Greenisca placida Bükkszentlélek,

MBK013 4648 23.x.2005 Brachypodium sp. F. Kozár
Green Hungary
Pseudochermes fraxini
MBK023 4658 Budapest, Hungary 11.v.2005 Fraxinus excelsior F. Kozár
(Kaltenbach)
Kotejacoccus turcicus Diyarbakır-Malabadi, M.B.Kaydan,
MBK064 4288 26.v.2008 38°09’335’’ 042°12’785’’ Quercus sp.
Kaydan & Kozár Turkey F. Kozár
Kotejacoccus turcicus Diyarbakır-Malabadi, M.B.Kaydan,
MBK054 4288 26.v.2008 38°09’335’’ 042°12’785’’ Quercus sp.
Kaydan & Kozár Turkey F. Kozár
Proteriococcus lauri M.B.Kaydan,
MBK069 4287 Hatay-Harbiye, Turkey 27.v.2008 36°07’859’’ 036°08’653’’ Laurus nobilis
Erkılıç F. Kozár
Van - Hakkari road,
Rhizococcus sp. MBK177 4504 06.vi.2009 38°22’248’’ 043°35’176’’ Euphorbia sp. M.B.Kaydan
Turkey
31
32
Coll.
Code
Rhizococcus acutus Van-Hakkari Road,
MBK119 4567 02.ix.2009 37°39’067’’ 043°52’992’’ Cynodon dactylon M.B.Kaydan
Goux Turkey
Rhizococcus greeni Hakkari-Başkale road,
MBK059 4271 23.v.2008 38°08’200’’ 044°04’546’’ Poaceae M.B.Kaydan
Newstead Turkey
Rhizococcus lactucae M.B.Kaydan,
MBK092 4340 Urfa-Birecik, Turkey 28.v.2008 37°01’781’’ 038°00’424’’ Echium sp.
(Borchsenius) F. Kozár
Rhizococcus micracanthus
MBK006 4641 Scabiosa sp. M.B.Kaydan
Danzig
Rhizococcus micracanthus Erzican-Erzurum Undetermined M.B.Kaydan,
MBK275 4800 08.vii.2010 39°46’131’’ 040°25’927’’
Danzig road, Turkey plant F. Kozár
Rhizococcus micracanthus Kars - Kağızman road,
MBK276 4913 17.vii.2010 40°16’283’’ 042°57’598’’ Dianthus sp. M.B.Kaydan
Danzig Turkey
Rhizococcus munroi Undetermined
MBK285 4928 Artvin-Borçka, Turkey 16.vii.2010 41°16’058’’ 041°46’108’’ M.B.Kaydan
(Boratynski) plant
Rhizococcus munroi
MBK222 4696 Óbarok, Hungary 26.ix.2008 Matricaria sp. F. Kozár
(Boratynski)
Rhizococcus munroi Nagykováicsi,
MBK011 4646 15.ix.2005 Scabiosa sp. F. Kozár
(Boratynski) Hungary
Rhizococcus munroi M.B.Kaydan,
MBK279 4744 Erzincan- Ağıl, Turkey 07.vii.2010 39°58’342’’ 039°28’163’’ Scabiosa sp.
(Boratynski) F. Kozár
Coll.
Code
Rhizococcus munroi Ardahan - Şavşat road,
MBK280 4866 15.vii.2010 41°09’977’’ 042°37’243’’ Artemisia sp. M.B.Kaydan
Introduction
(Boratynski) Turkey
Rhizococcus reynei Hakkari - Çukurca
MBK112 4555 01.ix.2009 37°29’902’’ 043°34’230’’ Cichorium sp. M.B.Kaydan
Schmutterer road, Turkey
Rhizococcus zernae Van - Çatak road,
MBK196 4486 06.vi.2009 38°14’778’’ 043°14’688’’ Umbellifera M.B.Kaydan
(Tereznikova) Turkey
Rhizococcus zernae Van-Hakkari Road, Undetermined
MBK096 4310 23.v.2008 37°41’644’’ 043°56’266’’ M.B.Kaydan
(Tereznikova) Turkey plant
Rhizococcus zernae Hakkari-Yüksekova- Undetermined
MBK095 4307 23.v.2008 37°40’881’’ 044°03’376’’ M.B.Kaydan
(Tereznikova) Ortaç, Turkey plant
Rhizococcus zygophylli
MBK009 4644 Alibotosh, Bulgaria 19.vi.2009 Achillea sp. I.Gavrilov
Archangelskaya
Outgroup
Phenacoccus aceris
MBK009 4662 Budapest, Hungary 11.v.2005 Fraxinus excelsior F. Kozár
Signoret
33
rates and a proportion of invariant sites (GTR) to each partition, using default priors.
Four Markov chains, three hot and one cold (program default), were run simultaneously
for five million generations, with trees sampled every 1000 generations. A plot of log-
likelihoods over time was examined, and the first 1000 trees, generated before the analysis
had achieved stationarity, were discarded as ‘burn-in’ (Gullan et al., 2010). The support
below 50% are not indicated on the tree.
Voucher specimens were slide-mounted by using the methodology Kosztarab &
Kozár (1988) from the cuticle of the actual specimens from which DNA was extracted,
and the slide-mounts deposited in the Coccoidea Collection of the PPI and Kaydan’s
collection (KPCT). Only adult female eriococcids were used for DNA extraction.
Molecular phylogenetic analyses of Acanthococcidae FG for the Palaearctic
region using Bayesian interference and Maximum likelihood [(ML tree, with bootstrap
proportions (BS)] is presented in Fig. 13. Seven main clades were obtained from these
analyses but the support for most clades was poor, although the support within each
of the clades was generally fairly good. Clade G is including species feeding on woody
plants included only Gossypria spuria, Acanthococcus sp. 1 and A. roboris (both Acanthococcus
species are closely related with A. aceris). Although it has been hypothesized in several
studies that Gossyparia species should be included in Acanthococcus, the adult females of
these two genera differ morphologically and this difference is supported with molecular
data here. In the phylogenetic study by Ouvrard & Kozár (2009) based on adult female
morphology, Gossyparia spuria was embedded within Acanthococcus (in a clade which also
included A. aceris and A. roboris), sharing characters such as presence of the frontal lobe;
shape of claw digitules; presence of multilocular pores on abdominal segments, and the
distribution of macrotubular ducts. Despite these similarities, the (G. spuria + Ac aceris)
clade had BS support of 80, suggesting that these two species are closely related but
we would like to keep the present generic name as Gossyparia and Acanthococcus. Clade F
included the other woody-feeding acanthococcid species: Kotejacoccus turcicus, Proteriococcus
lauri plus Pseudochermes fraxini.
The remaining species occurred in 5 further, often overlapping clades (A-E), but these
had poor BS values even though the groups can be easily separated morphologically (see
vignettes in Fig. 13). Ouvrard & Kozár (2009) considered that the most parsimonious
scenario for the evolution of host-plant choice in the “Acanthococcidae FG” was that
these insects originally evolved on woody plants but later moved to herbaceous plants
and finally to grasses. This trend can also be seen in Fig. 13. The “grass-feeding species”
are mainly found in clade A whilst the “herbaceous species” are mainly found in clades B,
C, D, E, and are mainly represented by Rhizococcus species. Several species could be found
in a group of species belonging to other clades which could be a result of incorrect
generic assignation, or may represent new genera, or they could be the result of multiple
evolutions or parallel evolution. The grass-living Rh. greeni may have moved to grasses
from herbaceous plants because of the length of the stylet loop. Unfortunately we were
unable to include E. buxi in our analyses because no PCR product was available for this
species. Nonetheless, our results appear to support the monophyly of the Palaearctic
Introduction 35
Figure 13. Phylogenetic tree for different genera and species of the Acanthococcidae family
group in the Palaearctic Region based on molecular data.
Acanthococcidae FG, although we included few species and the clades were not well
supported. Further studies need to be done to confirm these conclusions.
Based on the results of our molecular studies and on the morphology of the
Palaearctic Acanthococcidae FG, the group can be divided into two assemblages based
on feeding habits: a “woody-feeding group” or “Acanthococcinae” (Clades F and G)
and an “herbaceous-grass-feeding group” or “Rhizococcinae” (Clades A-E) apart from
Borchseniococcus dusgunasae (Clade C) which should be in the Ovaticoccini, according to
Kaydan & Kozár (2008)).
Adult female Acanthococcinae can be recognized by the presence of the following
features: (i) strong spines on the dorsum as well as on the body margin, (ii) narrow,
elongate microtubular ducts, (iii) only four setae on each posterior tibia, and (iv) a long
stylet loop, almost as long as the body.
Adult female Rhizococcinae can be recognized by the presence of the following
features: (i) strong spines on the body margin and sometimes on the anterior part of
dorsum but otherwise absent on dorsum, (ii) short microtubular ducts, and (iii) five setae
on each posterior tibia. In this subfamily, it seems likely that two new evolutionary lines
appeared, namely the tribes Rhizococcini (which mainly live on herbaceous plants) and
the Anophococcini (which live mainly on grasses). The Rhizococcini can be diagnosed
by: (i) spinose marginal setae, along the entire body margin, (ii) dorsum with only a few
enlarged setae similar to those along the margin, and (iii) the stylet loop usually reaching
the median coxae. The Anophococcini, on the other hand, are diagnosed by: (i) the
marginal spines normally only present on the last abdominal segments or restricted to the
anal lobes, (ii) all dorsal setae short and narrow, (iii) the stylet loop often short, as long as
the labium, and (iv) discoidal pores with a special structure and arrangement (generally
doubled rimmed). The Anophococcini generally live on the leaves or in the leaf sheaths
of grasses.
Distribution
The family group Acanthococcidae is widely distributed in the World apart from a few
tropical countries, such as Africa, with greatest species abundance in the Mediterranean
and temperate zones. Also we should note that they seem pretty abundant in the
Neotropics, but that genera should be restudied according to a new concept of the family
group.
Here we used the records in the World Catalogue of Miller & Gimpel (2000), the
scale insect data base ScaleNet (Miller et al., 2012) and the Palaearctic Catalogue of
Köhler (1998), with the addition of some new data published more recently and from
a zoogeographical analysis (after Kozár & Foldi, 2009). The zoogeographic regions we
have used are those in Kozár & Ben-Dov (1997) (i.e. with the island areas in the southern
hemisphere and in the Pacific divided into a New Zealand & Pacific region, an Austro-
Oriental region and an Australian Region).
Introduction 37
According to the survey by Kozár (2009), the Acanthococcidae FG contains 628

species in 97 genera. There has been a steady increase in the number of new genera and
species descriptions since 1758 (Fig. 14 a, b). It seems that the Palaearctic Region has
been somewhat better studied, and consequently the increase there has been smaller.
These data support the suggestion that the descriptive phase of the eriococcid fauna is
still very intense and that many new species remain to be discovered and described in all
zoogeographic regions.
Figure 14. Description rate of genera (a) and species (b) of Acanthococcidae family group
in the World and in the Palaearctic Region (Kozár & Foldi, 2009; Kozár, et al., 2010).
The Acanthococcidae FG has the greatest number of genera and species in the
Palaearctic Region (31 genera and 198 species), followed by the Neotropical (27 genera
and 67 species), Australian (26 genera and 171 species), New Zealand & Pacific (22 genera
and 102 species), and the Nearctic (15 genera and 86 species) regions. Remarkably, the
Acanthococcidae FG fauna of the Ethiopian (5 genera and 9 species), Austro-Oriental
(6 genera and 10 species) and Oriental (9 genera and 28 species) regions are very poor.
The number of endemic species is especially high in the Palaearctic, Australian and New
Zealand & Pacific and Neotropic regions (Fig. 15-16).
Although the greatest generic richness is found in Euro-Siberian subregion, the
species richness is generally the same (more than 70 in the Euro-Siberian, Irano-Turanian
and Far-Eastearn subregions but only 40 Irano-Turanian) in the Palaearctic Region (Fig
17a). The Acanthococcidae FG fauna of each of the main zoogeographic regions appears
to be unique, with low similarity both at the generic and specific level (Fig. 17. a, b). In
the Palaearctic and Nearctic regions, the Acanthococcidae FG fauna on herbaceous and
grass-like plants is rich. Although the gall-inducing groups on Myrtaceae are extremely
numerous in Australia (especially on Eucalyptus) and in South America (Hodgson &
Miller, 2010), the Nothofagaceae harbour many species in New Zealand, Australia and
South America. The strongest connection we could find was that between the Oriental
and Palaearctic regions, with 5 common genera and 20 common species. The number of
genera and species shared between the Nearctic and Palaearctic and between the Nearctic
and Neotropical regions was few. The Oriental and Austro-Oriental regions are connected
by species of Gossypariella and Sanguicoccus and the Australian Region is connected to
the New Zealand & Pacific region by Madarococcus species. The morphological similarity
of Pseudomontanococcus, from the Austro-Oriental region, to Montanococcus, from New
Zealand, suggests a new possible connection between these regions. New Zealand also
shares a few genera and species with the Neotropical Region (i.e., Madarococcus). On the
other hand, unusual distributions are found in a few genera, such as Cryptococcus, which
has several species in the Palaearctic but is also found in the Nearctic and in New Zealand
region. A rather similar distribution is shown by Kuwanina, which has species in Australia,
New Zealand and the Palaearctic regions.
The Venn analyses for the subregions of the Palaearctic Region show that the ratio
of local (“endemic”) species is greatest in the Mediterranean and Far-East subregions.
These connections are strongest between the Euro-Siberian, Mediterranean and Far
East subregions which share between 8 genera and 11 species. The close connection
of the Euro-Siberian and Mediterranean subregions is supported also by the similarity
analyses (Fig. 18). All the Venn diagrams, both at the generic and species level, suggest
that most species evolved locally and that the roots of the connections were lost in
ancient times (Fig. 19, 20). Only four species have been recorded from all subregions of
the Palaearctic (i.e., R. greeni, R. munroi, G. spuria, P. fraxini, An. insignis), and only 3 species
are cosmopolitan pests (O. agavium, R. coccineus, U. araucariae). Four species have a wider,
Holarctic, distribution (A. azaleae, R. greeni, G. spuria, An. insignis).
Introduction 39
Figure 15. Distribution of genera and species number of Acanthococcidae family group
(some species are distributed in several regions, so the sum of numbers higher than the
known number of species) in regions of the World (Kozár & Foldi, 2009; Kozár et al., 2010).
Figure 16. Distribution of genus and species numbers of Acanthococcidae family group in
subregions of the Palaearctic region (original and partly from Kozár & Foldi, 2009; Kozár
et al., 2010).
a b
Figure 17. Species level similarity of Acanthococcidae family group of the World (a) and in
the Palaearctic Region (b) (Kozár & Foldi, 2009; Kozár et al., 2010).
The species richness of the Australian and New Zealand regions is in accordance of
Hoy’s (1962) view that the roots of this family group is hidden under the Antarctic ice.
Williams (1984) also considered that the generic and species richness of the southern
Hemisphere indicated a southern origin of the family. It therefore seems that the main
evolution of this family occurred in the southern hemisphere and only subsequently
spread to the more temperate regions and the new data support this Hoy’s (1962)
opinion. The distribution of the Acanthococcidae FG suggests that, apart from South
Africa, the parts of the World with a Mediterranean climate (and thus with an arid climate
and sclerophyllous plants), are the richest in genera and species. The present day generic
and species composition is formed basically by extinction and vicariance and, to a smaller
extent, by dispersion. The relationships between zoogeographic regions presented here
differ slightly from those suggested by Hoy (1962). The New Zealand & Pacific region is
not a blind separate region, but is connected by a few genera and species with both the
Australian and the Neotropical regions, and provides a dispersal route to the Australian,
Nearctic, Oriental and Austro-Oriental regions. There is also a connection between the
Nearctic and the western part of the Palaearctic region (Fig 21).
Hoy (1962) also mentioned a possible connection between the Australian and
Palaearctic faunas as they share genera such as Gossyparia and Cryptococcus, which live on
woody plants. The genus Gossypariella, from the Oriental Region, was studied recently
by Kozár et al. (2007), and this genus could also connect the Australian and Palaearctic
regions. Indeed, species of Gossypariella, but known under different generic names, could
be present in both Australia and the Palaearctic. Furthermore, members of Acalyptococcus,
Sisyrococcus and Scutare, a complex of similar genera, have been found in New Zealand and
in the Oriental and Palaearctic regions. These observations suggest that New Zealand
should not be excluded from the Oriental-Palaearctic route of dispersal by members of
the Acathococcidae FG.
Introduction 41
Figure 18. Venn diagram of the number of total and “endemic” genera of the Acanthococcidae
family group in different regions of the World (Kozár & Foldi, 2009; Kozár et al., 2010).
Figure 19. Venn diagram of the number of total and “endemic” species of the
Acanthococcidae family group in different regions of the World (Kozár & Foldi, 2009; Kozár
et al., 2010).
Figure 20. Venn diagram of the number of total and “endemic” species of the
Acanthococcidae family group in different subregions of the Palaearctic Region (Kozár &
Foldi, 2009; Kozár et al., 2010).
Introduction 43
Figure 21. Modifications (in gray) to Hoy’s (1962) dispersion pattern of the Acanthococcidae
family group in different subregions of the World (Kozár & Foldi, 2009; Kozár et al., 2010).
Host plant relationships

Hoy (1962) considered that an appreciation of the host-plant specialisations within the
acanthococcid and eriococcid species could be important in an understanding of the
evolutionary development of the family. He found that almost 39% of all species in
the Eriococcidae sensu lato were found on the Fagaceae and Myrtaceae, whilst only 7%
were found on Poaceae. He therefore considered that the Poaceae offered little help
in the study of the zoogeography of this family. However, the Poaceae are particularly
important in the Palaearctic region, in much the same way as the Myrtaceae are in Australia
and Nothofagaceae in the New Zealand and Nearctic regions. This is clear when one
compares the number of species of “Eriococcidae” sensu lato on Poaceae in New Zealand
with that in the Palaearctic: in New Zealand, only two out of 86 species (2.3%) were
found on grasses as compared with 61 out of 190 in the Palaearctic Region (34.9%). In
the Palaearctic, the Poaceae harbors a specialized Acanthococcidae fauna. As can be seen
in Fig. 22, most species collected on Poaceae belong to a grass-living clade of mainly
Rhizococcus (most of them are treated as Anophococcus here) species but also including most
other genera like Acanthococcus (part of them are considered as Rhizococcus here). Even
polyphagous species from the herbaceous-living clade (mainly Acanthococcus spp.) do not
infest grasses. As can be seen from the figure 22, the species-rich genus Acanthococcus
(these are here regarded as real Acanthococcus) prefers woody plants (the woody clade), and
Rhizococcus species live on herbaceous plants (the herbaceous clade). Apart from Poaceae,
the most abundant host-plant species in the Palaearctic are in the Asteraceae (15 species),
Ericaceae (7) and Amaranthaceae (5). The herbaceous plants provide a habitat for about
40 species, grasses 27, and woody plants 20 species. Most of the polyphagous species
(about 12) are on herbaceous plants.
Unlike most well-known polyphagous scale insect pests, Acanthococcidae FG species
living on woody plants are mainly oligophagous, suggesting that a stabilizing selection
may have played a role (Kozár, 1990). The appearance of Poaceae in the Palaearctic
Region opened a new niche for speciation of scale insects and for the Acanthococcidae
in particular. This availability of new niches resulted in the acanthococcids evolving in
new directions, giving rise to several new genera and species. Most of these grass-living
species live on the leaves and these species show a greater morphological similarity to
the species on woody or herbaceous plants than to those living in the leaf sheaths of
grasses. The species present in the leaf sheaths required different protection systems and
consequently they evolved new morphological structures (Foldi, 1997) such as heavily
sclerotized, sometimes fused, groups of disc pores and the appearance of disc pores on
the dorsum. It is possible to speculate that, in the Palaearctic Region, the Acanthococcidae
FG fauna of woody plants is older than that on grasses because the latter have their
ancestral roots somewhere in the southern hemisphere. A habitat preference for the
leaves of woody plants is almost absent in the Palaearctic eriococcid fauna, and are found
only in Eriococcus buxi (on Buxus) and Orontesicoccus lauri Erkılıç (on Laurus sp.) whereas this
habit is very common in the Australian and New Zealand eriococcid fauna.
On the other hand, as was mentioned by Cook & Gullan (2001), several
Acanthococcidae FG species in Australia have enlarged tubular ducts on the dorsum
similar to those in E. buxi. The presence of enlarged tubular ducts could be a valuable
character, not only at the generic but also at higher taxonomic levels. Thus, the non-
gall inducing E. buxi and the gall-inducing Cylindrococcus spiniferus Maskell and Gallococcus
heckrothi Takagi all have enlarged tubular ducts and form a clade in the phylogenetic study
of Gullan & Cook (2007) based on their molecular data. The presence of these enlarged
tubular ducts might reflect their common leaf-living habitat. It seems likely that living on
the leaf surface (as with E. buxi) preceded gall-induction.
Ouvrard & Kozár (2009), in their study of the links between the Palaearctic
Acanthococcidae FG and their host plants, found that the most parsimonious scenario
for the evolution of the plant choice required 12 steps (Fig. 22) and that the ancestor of
the Palaearctic Acanthococcidae FG fed originally on woody plants. The ancestor of the
same clade, but excluding E. buxi, switched to Poaceae. This was later followed by the
ancestor of the large terminal clade with (A. tavignani + A. istresianus [(now we consider
them as R. reynei (=R. tavignani) + R. istresianus in this book)] the basal clade switching to
herbaceous plants. In this clade, some other independent switches back to woody plants
and Poaceae occurred.
The most parsimonious scenario for the evolution of feeding site (i.e. whether on
leaves, twigs or roots) required also 12 steps (Fig. 23). This suggests that the ancestor
Introduction 45
Figure 22. Evolution of plant type preference as reconstructed on the topology obtained
from the parsimony analysis of the reweighted morphological characters (Ouvrard & Kozár,
2009).
Figure 23. Evolution of feeding site preference on the host – plant as reconstructed on the
topology obtained from the parsimony analysis of the reweighted morphological characters
(Ouvrard & Kozár, 2009).
Introduction 47
of the Acanthococcidae FG fed on leaves of woody herbaceous plants, and that the
ancestor of large terminal clade with A. tavignani + A. istresianus [(now we consider them
as R. reynei (=R. tavignani) + R. istresianus in this book)] as the basal clade switched to
branches and trunks as their preferred habitat. We can observe some switches back to
leaves in this clade, as well as two independent switches to leaf sheaths.
Parasitoid complexes
The study of parasitoid-scale insect relationships can also help in our understanding of
the generic composition of scale insects. Sugonyaev (1984) said that “Sometimes parasitoids
are better taxonomists than coccidologists”.
Kozár & Japoshvili (2010) undertook a phylogenetic analysis of the insect-parasitoid
relationships of the Palaearctic Acanthococcidae FG. For this analyses, the World
Database of Chalcidoidea (2009) was used, which contains data for six “eriococcid”
genera, including 21 species as hosts, and 62 species from 23 genera of parasitoids of
Chalcidoidea. A large number of parasitoid species have been recorded on a world-
wide basis from Acanthococcus aceris (6 species), R. desertus (10), R. greeni (6), Eriococcus buxi
(7 all over the World), Gossyparia spuria (14), Greenisca brachypodii (6) and Neoacanthococcus
tamaricicola (9). Most of the better studied eriococcid genera have substantially different
generic complexes of parasitoids. The phylograms (UPGMA, cluster analyses) (Fig. 24)
show substantial differences between the parasitoid complexes of A. aceris, R. desertus, N.
tamaricicola, G. brachypodii, G. spuria and E. buxi when all parasitoid data were included.
A similar picture was found when the data for secondary parasitoids were omitted.
Although many of the numerous parasitoid records in the scale insect database
ScaleNet (Miller et al., 2012) and the Miller & Gimpel (2000) catalogue are questionable,
the records can provide some useful information. In the catalogue of Miller & Gimpel
(2000), more than 50 parasitoid species are mentioned from all families. Concerning
parasitoid species found only in the Palaearctic Acanthococcidae FG, there appear to be
8 species exclusive to Gossyparia spp., 17 species exclusive to Acanthococcus and Rhizococcus
spp., and 13 species only on Anophococcus species. On the other hand, some scale insect
species are hosts to polyphagous parasitoids, for example, R. greeni (Newstead) is host to
four parasitoid species, three of which are common parasitoids of the genus Anophococcus.
The relationships between parasitoid complexes and the Acanthococcidae FG
are more noticeable at the generic level of the parasitoids. For example, Acanthococcus
can be characterized by four obligatory parasitoid genera; the parasitoid complex of
Anophococcus is even more diagnostic as it has parasitoids belonging to ten genera, whilst
Gossyparia is parasitized by the specialized parasitoid genus Coccophagus (Encyrtidae). On
the other hand, Greenisca has very few parasitoids and this might be explained by the
young phylogenetic age of this group of species. However, although it could be argued
that the leaf sheath as a habitat might protect these species from parasitoids, numerous
examples are known of parasitoids of Pseudococcidae living in the same leaf sheaths,
suggesting that this is unlikely to be the answer. In the Palaearctic, E. buxi appears to be
Figure 24. Similarity of the parasitoid complex of the Acanthococcidae family group in the
Palaearctic region (Kozár & Japosvili, 2010).
totally without parasitoids, even where Buxus sp. bushes with infestations of E. buxi are
found intermingled amongst Ulmus sp. and Acer sp. trees with a high density of heavily
parasitized G. spuria and A. aceris. Even the oligophagous parasitoids infesting different
genera of Acanthococcidae do not appear to parasitise E. buxi in the Palaearctic. This
strongly suggests that the genus Eriococcus sensu stricto (and other species that may be
included later from other regions) did not evolve in the Palaearctic Region, and are older
than their Palaearctic relatives. One of the centers of diversity of Buxus sp. is East Asia
(Köhler & Brückner, 1989), and it is considered that the main route of dispersion of the
ancestral “eriococcid” groups from Australia toward the Palaearctic Region was through
the Oriental Region, perhaps E. buxi dispersed through Asia, leaving its parasitoids
behind. There is also no obvious explanation for the total absence of parasitoids on
Cryptococcus fagisuga and Pseudochermes fraxini either, but perhaps this situation is similar to
that of E. buxi. However species in these two genera are heavily attacked by predators in
this region.
Summary
The results of phylogenetic analyses of the Acanthococcidae FG such as Gullan &
Cook (2007), substantially support a traditional point of view of relationship “Complex
phylogenetic tree of Acanthococcidae FG based on female morphology” (Fig. 25) with
regard to some morphological and biological characters of the adult females. In this
figure we only summarize and combined data we have up to now. Several main lineages
are characteristic of some morphological characters such as setal shape, microducts,
anal ring structure etc., separated now in four families, of which four are treated here
(Acanthococcidae, Eriococcidae, Cryptococcidae and Kuwaninidae), following the
Introduction 49
views of Koteja (1974a) and Miller & Gimpel (2000). From these four families, three
(Cryptococcidae, Eriococcidae and Kuwaninidae) show some kind of degeneration in
certain organs at some stage in their development (e.g., antennal segmentation, reduction
of antennal segments, and lack of setae on the labium). These three families can be
characterized by the presence of one pair of setae on the basal labial segment, whereas the
Acanthococcidae have two pairs of setae on the basal labial segment in all developmental
stages and show little or no degeneration.
The family Acanthococcidae in the Palaearctic region is here divided into three
subfamilies (Acanthococcinae, Phloecoccinae and Rhizococcinae) plus the tribe
Ovaticoccini. The Phloecoccinae is a subfamily, whose sole representative in the
Palaearctic is Noteococcus hoheriae (Maskell), a species which has been introduced from
New Zealand, and which differs from all other “eriococcids” in the Palaearctic in having
numerous setae on the anal lobes. Species of Ovaticoccini differ by having a reduced
and simplified anal ring and in having unusually-shaped (truncated) marginal enlarged
setae. The Acanthococcinae are characterized by the presence of: (i) strong enlarged setae
on the dorsum as well as on the body margin, (ii) narrow, long microtubular ducts, (iii)
four setae on each posterior tibia, and (iv) a stylet loop almost as long as the body. The
Rhizococcinae are diagnosed by the presence of: (i) strong enlarged setae on the body
margin and sometimes also on the anterior part of the dorsum, (ii) short microtubular
ducts, and (iii) five setae on each posterior tibia (iv) stylet loop never reaching mid coxae.
It is also considered here that, within the Rhizococcinae, two new lineages have evolved,
tribes Rhizococcini and Anophococcini. The former lives on herbaceous plants and has
the whole body margin surrounded by spinose setae, the dorsum is at least partly covered
in enlarged setae similar to those on the margin, and it has a stylet loop that usually
reaches the median coxae. The Anophococcini live mostly on the leaves or in the leaf
sheaths of different grasses, and have the marginal enlarged setae usually restricted to
either the posterior abdominal segments or to the anal lobes; in addition, all dorsal setae
are short and narrow and the stylet loop is often short, only as long as the labium. In
addition the discoidal pores have a special structure and arrangement (e.g., doubled rim,
more sclerotized, in this tribe (Fig. 25 and Fig. 5)).
It is remarkable that character diversification (such as reduced anal ring like in
Borchseniococcus, having blunted enlarged setae like in Kotejacoccus) can be observed in the
Acanthococcidae FG and that this phenomenon is more common in the Palaearctic,
Nearctic and partly Neotropical regions. However the anal ring here shows also a trend
of degeneration i.e., Kuwaninidae or/and Cryptococcidae where the adult females have
lost their legs, the antennae are reduced, sometimes to one segment, and the anal ring is
is especially pronounced in the gall-inducing groups.
Males can be very useful in indicating relationships, but it is important to note that,
of the about 200 species found in the Palaearctic Region, the males are known only
for a few species. A study of the adult males of different genera currently included
in the Acanthococcidae FG (Chris Hodgson, unpublished) suggests that they are very
different from the males of the Pseudococcidae. It could be concluded on the basis of
50
Figure 25. Complex phylogenetic tree of Acanthococcidae family group based on female morphology.
Introduction 51
present knowledge that the Acanthococcidae FG originated in the Upper Cretaceous.

There are some molecular data connecting the Phenacoccinae and the basal families of
the Acanthococcidae family group, especially members of the family Acanthococcidae
FG. There is so far no data, except for that by Gullan & Cook (2007) who presented a
phylogenetic relationship of Coccoidea) regarding which ancestral groups are connected
to the each of the families and subfamilies within the Acanthococcidae FG.
Clearly more detailed analyses of the morphology of the females, males and nymphs,
as well as more molecular data, are necessary in order to split the family into more natural
groups, as mentioned by Cook et al. (2002). Because males are known for only a small
number of species, they might provide some additional data for the reconstruction of
the phylogenetic relationships of higher taxonomic categories when more taxa have been
studied.
All of the studies discussed above support to some degree the earlier generic systems
elaborated in the Palaearctic Region by Targioni Tozzetti (1868) and Signoret (1975),
and partly generalized by Borchsenius (1948, 1949) but with brilliant additions by Koteja
(1974a, b). These studies were variously followed by Danzig (1980), Miller & Williams
(1976), Kosztarab & Kozár (1988), quite differently by Tang & Hao (1995), Miller &
Gimpel (1996), and currently, in some way, by Kozár (2009), Hodgson & Miller (2010).
Several authors, use the work by Hoy (1963), the catalogue of Miller & Gimpel (2000)
and the papers of Miller & Miller (1992, 1993), which are based mostly on the Ferris
(1957b) system. This differs from the phylogeny presented here. In consequence, it
would need a lot of further systematic and nomenclatural work to find the right place for
all species known in the Acanthococcidae FG of the World. A more or less standardized
system would accelerate the exploration in different regions of the World?
It has been known for some time that the “Eriococcidae” (Acanthococcidae FG as
referred to here) are paraphyletic. These studies, but particularly the recent molecular
studies of Cook & Gullan (2001, 2007), suggest that there will have to be major
nomenclatorial changes in this group. With this present study we are aiming to clarify
generic and family structure of the Acanthococcidae FG in the Plaearctic region. On the
other hand we believe that further studies should be conducted to understand this group
more worldwide.
COLLECTING AND MOUNTING METHODS
The various methods useful for collecting Coccoidea were discussed in detail by
Kozár & Miller (1998). Of these, probably the most effective method for species of
Acanthococcidae FG, which have a white or cream wax cover, and live mostly on the
plant surface, on leaves, in galls, in the leaf sheaths of the grasses, twigs, bark, on the
root crown, are the visual methods. However, sometimes they live on the roots and in
litter mixed with moss and lichens and for these sifting or the use of a Berlese funnel
can help one to collect them. We have found that the soil, litter and moss samples in
museum collections of the Class Arachnida and of Collembola and Coleoptera are
important sources for some groups. In addition, a hand-held suction sampler, such as
a D-Vac (Samu & Sárospataki, 1995) is also very efficient in vegetation, as is beating.
Barber traps and pheromone traps can also provide important additional information
on Acanthococcidae FG fauna. For a quick exploration of a given locality, a variety of
collecting and trapping methods is recommended.
For mounting, Wilkey’s method is suggested (Kosztarab & Kozár, 1988), as

follows:
Because molecular studies have special requirements, specimens should be stored in both
70 % (for morphological studies) and 96 % (for molecular studies) ethanol.
For mounting, alcohol preserved specimens should be placed in a 10% solution of KOH.
The KOH can be heated gently (never boiling) or left overnight, until the specimens
are soft and have begun to clear. At this point, the specimen should be gently pulsated
to remove the inner contents. When the specimens are transparent, they are transferred
to 70% ethyl alcohol for 10-15 minutes; gentle cleaning of the body surface can be
continued, if necessary.
The specimens are then placed in 10-15 drops of staining solution (15 ml Essig’s Aphid
Solution with a 2% aqueous solution of 20 drops acid fuchsin, 20 drops lignin pink, and
20 drops of erythrosin) (Essig’s Aphid Fluid; 20 parts of lactic acid, 4 parts glacial acetic acid,
2 parts phenol, and one part of distilled water), and left without heating for 15-20 minutes.
They are then returned to 95% ethyl alcohol for 10 minutes, or until excess stain is
removed. The timing needs to be judged carefully.
Finally, the specimens are transferred to clove oil to remove excess stain and wax and are
left until they become glass-like (this sometimes takes up to 30 minutes). They are then
mounted in Canada balsam or other mountant on microscope slides and covered with a
cover slip.
The slides are placed on a tray to dry.
Introduction 53
DEPOSITORIES OF TYPES AND STUDIED INSECTS
In the course of preparation for this work, many specimens in the collections in the
BMNH, MNHN, USNM, ZMAS and other museums and institutes were studied.
Abbreviations of depositories according to ScaleNet:
AAKA: Institute of Zoology, Kazakhstan Academy of Science, Almaty, Kazakhstan

ASAR: Academie des Sciences Agricoles et Forestieres, Bucarest, Romania (Savescu
collection now with his wife)
BMNH: The Natural History Museum, London, England, UK
BPRL: Institute of Plant Protection, Riga, Latvia
DAFNAE: The Scientific Museums of the University of Padova, Department of
Agronomy, Food, Natural Resources, Animals and Enviroment, Entomology.
EISC: Entomological Institute, Shanxi Agricultural University, Shanxi, China
EMLS: Entomological Museum of the Zoological Institute, Lund, Sweden
GPPT: Scientific Research Institute of Plant Protection, Tbilisi, Georgia
HNHM: Hungarian Natural History Museum, Zoological Department, Budapest,
Hungary
HUSJ: Entomological Institute, Faculty of Agriculture, Hokkaido University, Sapporo,
Japan
ICVI: Department of Entomology, The Volcani Centre, Bet Dagan, Israel
IEBC: Institute of Entomology, Academy of Sciences, Beijing, China
ITLJ: Insect Taxonomy Laboratory, National Institute of Agricultural Environmental
Sciences, Konnon-dai, Yatabe, Tsukuba-shi, (I. Kuwana), Ibarakiken, Japan
KPCT: Kaydan’s Personal Collection, Turkey
KAYJ: S. Kanda Collection, Asano Senior High School, Kanagawa-ku, Yokohama, Japan
LSUK: The Linnean Society of London, England, UK
MNCN: Museo Nacional de Ciencias Naturales, Madrid, Spain
MNHN: Muséum national d'Histoire naturelle, Paris, France
NZAC: New Zealand Arthropod Collection, Landcare Research, Auckland, New Zealand
PASK: Institute of Systematic and Experimental Zoology, Polish Academy of Sciences,
Krakow, Poland
PPI: Plant Protection Institute, Centre for Agricultural Research, Hungarian Academy of
Sciences, Budapest, Hungary
SCHM: The Schmutterer Collection, Wetlenberg, Germany. Now in DEIC: Deutsches
Entomologisches Institut Leihschein, ZALF, Müncheberg, Germany
SIEC: Shanghai Institute of Entomology, Academia Sinica, Shanghai, China
TARI: Taiwan Agricultural Research Institute, Entomology Collection, Taichung,
Taiwan, China
UCD: The Bohart Museum of Entomology, University of California, Davis, California,

USA
USNM: United States National Entomological Collection, Coccoidea collection housed
at the U.S. Department of Agriculture, Beltsville, Maryland, 20705, USA
UZMH: University of Helsinki, Finnish Museum of Natural History, Helsinki, Finland
VASK: Zoological Institute, Academy of Sciences of Ukraine, Kiev, Ukraine
YPQS: Yokohama Plant Quarantine Service Station, Yokohama, Honshu, Japan
ZMAS: Zoological Museum, Academy of Science, St. Petersburg, Russia
ZMHB: Museum für Naturkunde der Humboldt Universität zu Berlin, Berlin, Germany
Introduction 55
ACKNOWLEDGEMENT
The first author wishes to thank the European Synthesys Programmes (GB-TAF-1318,
FR-TAF-1319, ES-TAF-2590) and the Hungarian Scientific Research Found – OTKA
(Grants No. T 034236, T 048801, T 075889) for financial support for this project. The
Authors thank Dr. Jon H. Martin for help in the Natural History Museum, London,
and Isabel Izquierdo and Paris Mercedes in the Museo Nacional de Ciencias Naturales,
Madrid (Spain), for the loan of specimens; and especially Dr. Douglas J. Williams (Natural
History Museum, London, UK) and Rosa Henderson (Landcare Research, Auckland,
New Zealand) for study of some species. Also to Mary Gimpel and Barbara Denno for
the help with rare reprints.
Thanks are due to Imre Foldi, Muséum national d’Histoire naturelle, Paris, France,
for his great financial support of the work of the senior author, and to Christopher J.
Hodgson for his permission to include his drawing and description of the male and
scientific discussions on some parts especially introduction part of this work.
The authors are especially grateful to Douglass R. Miller (USDA, Beltsville, USA),
D. J. Williams (BMNH, UK) and C. Hodgson (Cardiff, UK), and D. Matile Ferrero,
(Muséum National d’Histoire Naturelle, Laboratoire d’Entomologie, Paris, France), for
help and for making it possible to study the collections.
Studies and visits to the British Museum (UK) and Systematic Entomology Laboratory
USDA (USA) were partly financed by exchange agreements between the Hungarian
Academy of Sciences, the Royal Society (UK), and the National Science Foundation
(USA).
Thanks are also due to Prof. Jan H. Giliomee (Department of Botany and Zoology,
University of Stellenbosch, Private bag X1, Matieland, South Africa) for reading parts of
the manuscript and making valuable suggestions.
The publication of the book was supported by Hungarian Scientific Research Found–
OTKA (T 075889).
We thank Dr. Takumasa Kondo (CORPOICA, Palmira, Colombia) for kindly
reviewing the text. We are also very grateful to Dr. Wu San-an for the help with translation
of some Chinese descriptions, and to the more than 250 colleagues who have been kind
enough to put their published work at the author’s disposal, and who collected and sent
insects to us. Because of space limitations, unfortunately we cannot list their names.
Our librarians, Mrs. Dr. Tamásné Sárai, Lászlóné Papp and Veronika Hertelendyné
Bogdány (Plant Protection Institute, Hungarian Academy of Sciences, Budapest,
Hungary) were very helpful in searching for some of the literature.
Without all this help, this revision could never have been finished.
SYSTEMATIC PART
The systematics of the Acanthococcidae FG is still very controversial. Here we are using
the taxonomic concepts of Koteja (1974a), although it is clear from molecular studies
(e.g. Gullan & Cook, 2007) that the family group as currently understood is paraphyletic,
and that there is a close relationship with other families such as Apiomorphidae,
Beesoniidae, Calycicoccidae, Cerococcidae, Dactylopiidae, Kermesidae, Stictococcidae
plus the Xerococcus group.
The Acanthococcidae FG in the Palaearctic Region is here considered to include four
families: Acanthococcidae, Cryptococcidae, Eriococcidae and Kuwaninidae, mainly
following Koteja (1974a) but also partly Gullan and Cook (2007). However the present
classification needs still further study, such as with more molecular data, and on nymphal
and male morphology which could provide more information than that based purely on
female morphology.
Key to the superfamilies
1 – Abdominal spiracles present............................ Orthezioidea (= Archaeococcoids)

(Carayonemidae, Margarodidae Marchalinidae, Matsucoccidae, Monophlebidae,
Ortheziidae, Steingeliidae, Xylococcidae, etc.)
− Abdominal spiracles absent...........................................Coccoidea (= Neococcoids)
Key to the family groups in Coccoidea
1. – Often with well-developed, sclerotized anal lobes; body of adult female usually
covered by felt-like ovisac; pygidium, ostioles, circuli, cerarii, 8-shaped pores, scale
cover (test) always absent; trilocular pores usually absent; without anal plate or
plates, sometimes with cauda, never plate-like. . ..Acanthococcidae family group
− Without sclerotized anal lobes; body of adult female never covered by felt-like
ovisac; pygidium, ostioles, circuli, cerarii, 8-shaped pores, scale cover (test) present
in one or more families; trilocular pores sometimes present; often with an anal
plate or plates. ......................The rest of the families of Coccoidea superfamily
Acanthococcidae 57
Family group:
ACANTHOCOCCIDAE
Signoret, 1875 sensu Koteja, 1974
Acanthococcites Signoret. 1875: 314.

Acanthococcidae Signoret, 1875, family group, Koteja, 1974b: 294.
Family group diagnosis

Body of adult female in life generally elongate, but oval or rotund in Cryptococcidae and
Kuwaninidae; anal lobes well-developed, long and generally slightly to heavily sclerotized;
these occasionally undeveloped (e.g., in Cryptococcidae and Kuwaninidae). Body covered
with a felt-like ovisac (Fig. 1).
For morphological details please see the introduction part (pp. 11-23)
Host plants
The members of the Acanthococcidae FG are widely distributed on woody and
herbaceous plants and on grasses. The families Cryptococcidae, Eriococcidae and
Kuwaninaidae mostly live on woody plants. More information is given under Host
relationships in the Introduction.
Distribution
In the Acanthococcidae FG, members of the family Acanthococcidae are mostly found in
the Palaearctic, Nearctic and Neotropical regions, plus a few in the Oriental region. They
are less common in the Pacific & New Zealand Regions and in Australia. The Ethiopian
region is very poor. Members of the Cryptococcidae and Kuwaninaidae have a disjunct
distribution, being found in New Zealand and in the Palaearctic and Nearctic regions.
Perhaps the Cryptococcidae originated from New Zealand and “somehow” spread to
the North Temperate Zone (Gwiazdowski et al., 2006). The origin and distribution of
the Eriococcidae, in this narrow sense, is not clear but is unlikely to have been in the
Palaearctic because Eriococcus buxi (the type secies of Eriococcus) belongs to the BSE
(Beesoniidae-Stictococcidae-Eriococcidae) clade, according to molecular data (Gullan &
Cook, 2007), which is mainly found in Australasian, Neotropical and Oriental regions. In
addition, E. buxi is restricted to Buxus spp., which main distribution area is in the Oriental
and Pacific regions. On the other hand, according to Gullan & Cook, the BSE group is
not phylogenetically close to the rest of “Eriococcidae” sensu lato (in their sense) and is
sister to the Godwanian clade which includes species from south hemisphere. Clearly, the
relationships within this family group needs much more study in the future.
Biology
Many members of this family group feed on the bark of the trunk, branches or twigs,
often in crevices, of woody plants, with only a few on the leaves in the Plaearctic Region.
However, grass-living genera mostly prefer the leaves or the leaf sheaths and many
species living on herbaceous plants infest the roots. As far as it is known, most species
have one generation a year and overwinter as nymphs or eggs.
Below we describe and illustrate 28 genera and 188 species: Acanthococcidae: 22
genera and 174 species (fossils: 4 genera and 6 species); Cryptococcidae: 2 genera and 6
species, Eriococcidae: 3 genera and 6 species, and Kuwaninidae 1 genus and 2 species.
Key to the families in the Acathococcidae FG present in the Palaearctic Region
1. – Basal segment of labium with two pairs of setae ..........................Acanthococcidae

− Basal segment of labium with 1 or without seta............................................................2
2. – Anal ring, antennae and legs well developed; macrotubular ducts particularly large
and elongate..................................................................................................Eriococcidae
− Anal ring, antennae and legs reduced; particularly large and elongate macrotubular
ducts absent.........................................................................................................................3
3. – Dorsum with macrotubular ducts; labium three segmented; basal segment with
one pair of setae ....................................................................................Cryptococcidae
− Dorsum without macrotubular ducts, these replaced by large invaginated
quinqelocular pores; labium small, reduced, indistinctly two segmented; basal
segment absent.............................................................................................Kuwaninidae
Acanthococcidae 59
Family:
ACANTHOCOCCIDAE
Signoret, 1875
Type genus: Acanthococcus Signoret, 1875: 16.

Common name: Felt scales, acanthococcid scales.
Lit.: Afifi, 1968: 203; Borchsenius, 1948: 501, 1949: 48; Cook & Gullan, 2001: 60;
2004: 444; Danzig, 1975: 44; 1980: 3; Fernald, 1903: 70; Ferris, 1955: 69; Goux, 1948:
5; Green, 1922: 351; Hoy, 1963: 66; Gullan & Cook, 2001: 94; 2007: 413; Hodgson
& Henderson, 1996: 192; Hodgson & Miller, 2010: 7; Hodgson & Trencheva, 2008:
12; Kawecki, 1985: 27; Kaydan & Kozár, 2008:16; Kosztarab & Kozár, 1978: 6;
1988: 276; Koteja, 1974a: 296; 1974b: 76; Koteja & Zak-Ogaza, 1981: 512; Kozár,
2009: 91; Kozár & Walter, 1985: 73; Köhler, 1998: 371; Leonardi, 1920: 425; Matile-
Ferrero, 1988: 67; Miller & Gimpel, 2000: 12; Miller & González, 1975: 131; Miller &
Miller, 1992: 2; 1993: 9; Miller & Williams, 1976: 118; Ouvrard & Kozár, 2009: 101;
Schmutterer, 1952: 378; 2008: 77; Tang & Hao, 1995: 449; Tereznikova, 1981: 15;
Williams, 1969b: 318; 1985a: 357; Wu, 2000: 251.
Acanthococcites Signoret. 1875: 314.

Acanthococcidae (in part) Maskell, 1887: 88.
Dactylopites Signoret, 1875: 339.
Dactylopiidae (in part) Fernald, 1903: 39, Ferris, 1955: 69.
Kermidae Signoret; (in part) Ferris, 1937: 5.
Kerminae Signoret; Balachowsky, 1948: 253, partim;
Dactylopiidae Signoret; Ferris, 1955, partim;
Eriococcini Cockerell, 1899a: 389.
Eriococcidae, Hoy, 1963: 1.
Acanthococcidae Signoret, 1875; Koteja, 1974a: 296.
Comments
The status of Acanthococcidae family has always been controversial but has become more
so recently. The confusion regarding the selection of the correct family name and type-
genus has been discussed in the following major works: Afifi (1968); Borchsenius (1949),
Ferris (1955), Hoy (1962, 1963); Koteja (1974b); Koteja & Zak-Ogaza (1981), Miller and
McKenzie (1967); Morrison & Morrison (1966); Miller (1969); Miller & Williams (1976);
Tang & Hao (1995), Williams (1969b, 1985a), etc. Recently, molecular studies by Cook
and Gullan (Cook & Gullan, 2004; Gullan & Cook, 2007) have confirmed the findings
of Cox & Williams (1987) which suggested that the family is paraphyletic and that taxa
that are currently included in it belong to at least three quite separate clades. However, in
the present work, we follow partly the system of Koteja (1974a,b; 1990), Koteja & Zak-
Ogaza (1981), Kozár & Konczné Benedicty (2008a, 2008b) and Kozár (2009).
Family diagnosis
The body of the adult female and her eggs are often entirely enclosed in a felt-like sac (or
exceptionally only partly enclosed e.g., Gossyparia), thus the common name "felt scales".
Mounted adult female

The morphology is still very diverse in this family, here we have used a combination of
characters to define most genera
Adult females usually elongate-oval, rarely circular. Anal lobes sometimes sclerotized.
Most species with 6 - or 7 segmented antennae; large, membranous frontal lobes, or
minute, sometimes barely visible, frontal tubercles present near to base of each antenna,
sometimes both present together; labium normally 3 segmented, with 16˗18 setae;
median setae of apical segment of labium either hair-like or spine like; basal segment
of labium with 2 pairs of hair-like setae; legs normally 5-segmented; leg setae reduced in
number when compared with pseudococcids, tibia with 4˗6 setae; claws rarely without
a denticle; translucent pores normally present on hind coxae and femora, rarely also on
tibia; derm seldom with bilocular, trilocular or simple pores; if trilocular pores present,
these structurally different to those in Pseudococcidae; quinquelocular and multilocular
(7˗10 locular) disc pores present; often with oval disc (cruciform) pores, particularly
submarginally on venter; derm normally with distinctive microtubular ducts of variable
structure and size; invaginated macrotubular ducts present, often of 2 or 3 sizes; conical
setae generally present, each enlarged and conspicuous; anal lobes large and protruding;
anal ring normally sclerotized, with pores and 6 or 8 setae, rarely reduced; a lightly
sclerotized, cauda (median dorsal plate) often present anterior of anal ring, sometimes
even a ventral membranous present (e.g., in Acanthococcus, Rhizococccus); dorsal ostioles,
ventral circuli, and anal plates absent.
Mounted adult male

See under family group diagnosis. Described by Newstead (1903), Jancke (1955), Afifi
(1968), Hodgson & Trencheva (2008), Hodgson & Miller (2010).
Immature stages
These have been described for a few species, e.g., Gossyparia spuria (Herbert, 1924),
and Acanthococcus aceris, A. robosis and A. melnikiensis (Hodgson & Trencheva, 2008).
First instars and second-instar females are similar to adult females, but are smaller and
have fewer structures, including few microtubular ducts, and usually lack macrotubular
ducts. Second-instar males usually have macrotubular ducts and 7 segmented antennae
(generally six-segmented in second-instar females).
Acanthococcidae 61
The frequency of enlarged setae in adult females and first-instar nymphs of some
Acanthococcidae genera are shown in Fig. 11. It can be seen that there is a great variation
in the number of spines, and in their size and shape. For example, Orontesicoccus lauri
has unique setae, each with a microtubular duct on its base; Ovaticoccus agavacearum has
dome-shaped spines in both the adult female and nymphal stages, Aculeococcus yongpingensis
and Eriococcus buxi nymphs have nipple-shaped setae on the dorsum, while adult
female Hujinlinococcus nematosphaerus have both enlarged setae and nipple-shaped setae.
Gossypariella juniperi differs in having hair-like setae in both the adult female and first-
instar nymph. This variation in the structure and distribution of the dorsal setae in the
first-instar nymphs and adult females might be very useful for diagnosing the genera in
this family.
Biology
Species in this family feed on the bark of the trunk, branches or twigs of woody plants,
often in the crevices. Few adult females live on the leaves of woody plants, but those
taxa living on grasses prefer the leaves and the leaf sheaths. Species living on herbaceous
plants often infest the roots. There is usually one yearly generation and most species
overwinter as nymphs or eggs.
Host plant
Acanthococcidae are widely distributed and are found on species of such woody plant
genera as Acer, Populus, Quercus, Salix and Tamarix. On herbaceous plants, they are
common on species of Achillea, Artemisia, Astragalus, Calluna, Dianthus, Erica, Minuartia,
Teucrium, Thymus, Scabiosa, etc., while the grass-living species generally prefer species of
Agropyron, Brachypodium, Cynodon, Festuca, Koleria, Melica, Phleum, etc.
Distribution
Acanthococcidae are most abundant in the Palaearctic, Nearctic and Neotropical regions,
with a few species in the Oriental region. They appear to be less common in Pacific &
New Zealand Regions and in Australia and are very rare in the Ethiopian region.
At the present time, 174 species (plus 6 fossils) in 22 genera (plus 4 fossils) are known
from the Palaearctic region, and species richness is highest in the southern parts of the
Region. Few species are of economic importance, e.g. Acanthococcus aceris and Gossyparia
spuria, plus several species on herbaceous plants and grasses. Eriococcus cactearum, E.
coccineus and U. araucariae are only known in greenhouses and indoors in the northern part
of this Region.
Key to genera
1. – Base of enlarged setae with fused microtubular ducts..................................................2

– Base of enlarged setae without fused microtubular ducts............................................3
2. – Marginal and anal lobe enlarged setae setose ............... Orontesicoccus Kaydan gen. n.
– Marginal and anal lobe enlarged setae blunted
.....................................................................Pseudoacanthococcus Kozár & Kaydan gen. n.
3. – Middle abdomen with a narrow part as a constriction
..................................................................................Acalyptococcus Lambdin & Kosztarab
– Middle abdomen without a narrow part as a constriction............................................4
4. – Dorsum and margin with a mixture of large normal enlarged setae
and numerous cupoloid spines............................................. Hujinlincoccus Kozár & Wu
– Dorsum and margin without a mixture of large normal spines
and cupoloid spines............................................................................................................5
5. – Dorsal enlarged setae with three basal rings............... Neotrichococcus Miller & Gimpel
– Dorsal enlarged setae without three basal rings.............................................................6
6. – Dorsal enlarged setae drum stick-shaped........................ Kotejacoccus Kaydan & Kozár
– Dorsal enlarged setae not drum stick-shaped.................................................................7
7. – Anal lobes very strongly sclerotized, with numerous large and small
spines and teeth.......................................................................................... Noteococcus Hoy
– Anal lobes if present not strongly sclerotized, without numerous
large and small spines and teeth.......................................................................................8
8. – Both tarsal and claw digitules not capitated.................... Neoacanthococcus Borchsenius
– Tarsal and claw digitules generally capitated, rarely claw digitules can be
spinelike................................................................................................................................9
9. – Dorsal and marginal setae hair-like or flagellate...........................................................10
– Dorsal and marginal setae either spine like enlarged setae or smaller
spine like setae...................................................................................................................11
10. – Hind tibia with five setae, setae on dorsum and margin
flahellate......................................................................................Proteriococcus Borchsenius
– Hind tibia with four setae, setae on dorsum and margin
hairlike.........................................................................................Gossypariella Borchsenius
11. – Anal ring not well developed, often without pores.....................................................11
– Anal ring well developed, with a row, or rows of pores.............................................13
11. – Enlarged setae on dorsum present with or without spinelike setae..........................12
– Enlarged setae on dorsum absent only spinelike setae present
.........................................................................................Borchseniococcus Kaydan & Kozár
12. – Multilocular disc pores present on dorsum, cruciform pores on venter
in large groups on abdominal segments................................................Ovaticoccus Kloet
– Multilocular disc pores absent on dorsum, cruciform pores on venter
not in large groups on abdominal segments.................................. Hispaniococcus Kozár
Acanthococcidae 63
13. – Dorsum with cruciform pores or discoidal pores or with both................................14

– Dorsum without cruciform pores or discoidal pores..................................................17
14. – Dorsum with large sclerotized groups of sclerotized
quinquelocular pores............................................................................ Gregoporia Danzig
– Dorsum without large sclerotized groups of sclerotized 5-locular pores................15
15. – Dorsum without cruciform pores................................................ Greenisca Borchsenius
– Dorsum with cruciform pores........................................................................................16
16. – Marginal enlarged setae long, truncated................................................ Kaweckia Koteja
– Marginal enlarged setae short, wide, blunted........................Neokaweckia Tang & Hao
17. – Enlarged setae situated in a row only on the margin or restricted to the anal lobe
(except An. salsolae), live mostly on grasses..........................Anophococcus Balachowsky
– Enlarged setae situated all over the dorsum, lives where herbaceous
or woody plants.................................................................................................................18
18. – Microtubular ducts short, tibia with five setae, live mostly on
herbaceous plants............................................Rhizococcus Signoret (sensu Koteja, 1974)
– Microtubular ducts long, tibia with five setae, lives mostly on woody plants ........19
19. – Dorsal enlarged setae short and truncated, button-like small
setae present .................................................................................................Uhleria Cooke
– Dorsal enlarged setae not short and truncated, button-like small setae absent......20
20. – Macrotubular ducts present only on marginal part of dorsum;
dorsal enlarged setae of one size; ovisac do not cover the
dorsum of the body............................................................................ Gossyparia Signoret
– Macrotubular ducts present on entire dorsum; dorsal enlarged setae
normally of more than one size, labium with eight pairs of setae; ovisac
covers the whole body................................................................... Acanthococcus Signoret
Genus:
Acalyptococcus Lambdin & Kosztarab, 1977
Type sp.: Acalyptococcus eugeniae Lambdin & Kosztarab, 1977: 245, by monotypy.
Lit.: Kozár, 2009: 91; Miller & Gimpel, 2000: 19; Stoetzel & Miller, 1979: 14.
Description
Adult female pyriform, reddish brown in color and rests on a cushion of fluffy, white wax,
not completely enclosing female body. Mounted female pyriform, narrowed posteriorly,
with anal lobes conical and normally heavily sclerotized, sometimes strongly nodulose
with sclerotized teeth on inner margin, antennae 6 or 7 segmented; frontal lobes not
observed, labium three segmented, with 14 setae (of these 8 on apical segment); with
well developed segments and a weakly developed basal segment with two pairs of hair-
like setae. Legs well developed, inner side of hind tibia with only four setae, claw with
a denticle; tarsal and claw digitules clavate, longer than claw. Spiracles often with a few
associated disc pores; disc pores on venter only, quinquelocular, cruciform pores present
on ventral margin. Dorsum with bisclerotic segmental rows. Macro- and microtubular
ducts present or absent; if microtubular ducts present, bifurcate, with oval orifice, scattered
or forming transverse rows or bands on dorsum and on ventral margin. Enlarged conical
setae normally present on dorsal margin, 2 or 3 on each segment, on dorsum the setae
are of two categories, strong spine-like setae in a mid-dorsal longitudinal row, the smaller
setose spines form transverse rows on abdominal segments. Hair-like setae on venter
only. Anal ring well developed, sclerotized partly in double rows of pores and with 6 anal
ring setae often as long as apical seta on anal lobes; each anal lobe with a long apical seta
and usually with 3 short dorsal conical setae, ventral hair-like setae a1so present, suranal
setae hair-like, cauda usually conspicuous (Lambdin & Kosztarab, 1977).
Distribution
Only three species known in this genus from the Oriental and South East part of the
Palaearctic region.
Biology
Species of this genus have been found protected by the shelter of ants (Camponotus sp.)
and on Bambusa sp., Perotis indica and Perotis sp.
Comments
Acalyptococcus seems to be most closely related to the genus Scutare (Lambdin & Kosztarab,
1977), and shows some similarity with the genus Sisyrococcus Hoy (1962) (Miller et al.,
2013).
Acanthococcidae 65
Key to species
1. – Middorsal setae much smaller than the marginal ones.............................. A. trispinatus

– Some middorsal setae as long as marginal setae.............................................................2
2. – Margin of abdomen with three enlarged setae...............................................A. deformis
– Margin of abdomen with two enlarged setae................................................ A. graminis
Acalyptococcus deformis (Wang, 1974) (Fig. 26)

Eriococcus deformis Wang, 1974: 329.
Holotype: Female. China (Hainan), on Perotis sp., 28.v.1973, by T.C. Wang. By original
designation. Deposited in Beijing.
Lit.: Hua, 2000: 138; Kozár, 2009: 91; Kozár & Walter, 1985: 74; Köhler, 1998: 375;
Tao, 1999: 519; Wang, 1974: 329; 1980: 115; 1981: 287; 1982b: 143; 2001: 225; Yang,
1982: 104 (Miller et al., 2013).
Acanthococcus deformis; Kozár & Walter, 1985: 74. Change of combination.

Rhizococcus deformis; Tang & Hao, 1995: 526. Change of combination.
Eriococcus deformis; Miller & Gimpel, 2000: 186. Revived combination.
Acalyptococcus deformis; Kozár, 2009: 91. Change of combination.
Description
Unmounted female
Pyriform.
Mounted female
Female pyriform, 1.65–1.92 mm long, narrowed posteriorly, dorsum quite strongly
sclerotized. Anal lobes conical and normally heavily sclerotized sometimes strongly
nodulose with sclerotized teeth on inner margin, antennae 7 segmented; frontal lobes,
segments of labium, not mentioned. Anal lobe with a long apical seta and usually with 3
short dorsal conical setae.
Venter: Legs well developed, inner side of hind tibia with only four setae, claw with a
denticle; tarsal and claw digitules clavate, longer than claw. Spiracles with a few associated
disc pores; disc pores on venter only, quinquelocular, cruciform pores not detected. Hair-
like setae on venter only.Ventral hair-like setae also present on anal lobes, suranal setae
unusually long, hair-like, as long as anal lobe setae, 108 µm.
Dorsum: Dorsum with bisclerotic segmental rows. Macrotubular ducts numerous on
dorsum. Microtubular duct not mentioned. Enlarged conical setae normally present on
dorsal margin, 2 or 3 on each segment, on dorsum the setae are of two categories, strong
spine like forming a mid-dorsal longitudinal row, and smaller setose spines forming
transverse rows. Anal ring well developed, sclerotized, with 6 anal ring setae often as long
as apical seta on anal lobes. Cauda not conspicuous (Wang, 1974).
Ecology
Host plant: Bambusa sp., Perotis indica and Perotis sp.
Distribution: China (Guangdong, Hainan, Tibet).
Acanthococcidae 67
Figure 26. Acalyptococcus deformis (Wang, 1981), female. After Wang (1980) with
modifications.
Acalyptococcus graminis (Maskell, 1897) (Fig. 27) Combination nova

Eriococcus graminis Maskell, 1897: 243.
Syntype: Female. China (Hong Kong). Deposited in UCDC, USNM, NZAC.

Common name: Grass scale.
Lit.: Fernald, 1903: 75; Ferris, 1936:12; Hoy, 1963: 92; Hua, 2000: 137; Köhler, 1998:
377; Kuwana, 1917: 33; Tang & Hao, 1995: 448; Tao, 1999: 32; Wang, 1982b: 143;
WangZQ, 1982: 47; Yang, 1982: 104 (Miller et al., 2013).
Nidularia graminis; Lindinger, 1933a: 116. Change of combination.

Acanthococcus graminis; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus graminis; Miller & Gimpel, 2000: 219. Revived combination.
Acanthococcus graminis; Kozár, 2009: 92. Revived combination
Description
Unmounted female
Adult females enclosed in sacs of white cotton, closely felted. Male sacs are similar,
though smaller. Adult female elliptical, shriveling at gestation, dull greenish-brown, first
instars are yellow.
Mounted female
Body elongate oval. 1.5 mm long. Antennae 7 segmented. Frontal lobes not shown.
Venter: With long hair-like setae, few quinquelocular pores arranged in rows on
abdominal segments and some scattered around spiracles and on the margin. Legs long,
tibia shorter than tarsus. Claw with denticle. Macrotubular ducts on margin only.
Dorsum: Enlarged setae cylindrical, elongate, sides straight except basally where concave,
apices rounded, marginal setae conspicuously larger than other dorsal setae, 2 lateral
setae on margin of each abdominal segment. Dorsal setae few, on thorax longer than on
abdomen, form a pair of median longitudinal row. Macrotubular ducts few. Anal lobes
sclerotized, each with 3 enlarged setae. Cauda strongly sclerotized with teeth (Maskell,
1897).
Ecology
Host plant: Bambusa sp., Chrysopogon sp.
Distribution: China, Japan.
Biology: Unknown.
Acanthococcidae 69
Figure 27. Acalyptococcus graminis (Maskell, 1897), female. After Ferris (1936) with
modifications.
Acalyptococcus trispinatus (Wang, 1974) (Fig. 28) Combination nova

Eriococcus trispinatus Wang, 1974: 329.
Holotype: Female. China (Beijing). By original designation. Deposited in IEBC.

Lit.: Hua, 2000: Tang & Hao, 1995: 541; Tao, 1999: 35; Wang, 1980: 116; 1982b: 144;
WangZQ, 1982: 43; Yang, 1982: 105 (Miller et al., 2013).
Rhizococcus trispinatus; Kozár & Walter, 1985: 75. Change of combination.

Acanthococcus trispinatus; Miller & Gimpel, 1996: 604. Change of combination.
Rhizococcus trispinatus; Köhler, 1998: 401. Revived combination.
Eriococcus trispinatus; Miller & Gimpel, 2000: 367. Revived combination.
Rhizococcus trispinatus; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Unknown.
Mounted female
Body elongate oval. 1.68–2.28 mm long. Antennae 7 segmented: I: 30, II: 24, III: 49, IV:
36, V: 26, VI: 21, VII: 36 µm long; third segment without hair-like setae, other segments
with a few hair-like setae. Frontal lobes not shown. Eyes visible, situated on venter.
Venter: With long hair-like setae, few quinquelocular pores arranged in rows on
abdominal segments and some scattered around spiracles and on margin. Tibia and tarsus
of equal length. Claw with denticle. Macrotubular ducts few.
Dorsum: Enlarged setae cylindrical, elongate, sides straight except basally where concave,
apices rounded, marginal setae conspicuously larger than other dorsal setae, 3 lateral
setae on margin of each abdominal segment; 57–67 µm long. Dorsal setae few, small,
conical, slightly curved, 10 µm long. Macrotubular ducts numerous. Anal lobes each with
3 enlarged setae, mesal setae conspicuously thin. Cauda not shown (Wang, 1974).
Ecology
Host plant: Phragmites communis.
Distribution: China (Beijing).
Biology: Found on leaves.
Acanthococcidae 71
Figure 28. Acalyptococcus trispinatus (Wang, 1974), female. After Wang (1974) with
modifications.
Genus:
Acanthococcus Signoret, 1875
Type sp.: Acanthococcus Signoret, 1875: 16.
Lit.: Boratynski, 1962: 55, Borchsenius, 1948: 501; 1949: 331; Danzig, 1975: 42; 1980:
204; Ferris, 1955: 94; Hodgson & Miller 2010: 6; Hoy, 1963: 8; Kawecki, 1985: 27;
Kosztarab & Kozár, 1988: 276; Koteja 1974a: 296; 1974b: 276; Koteja & Zak-Ogaza;
1981: 582; Kozár, 2009: 111; Kozár & Konczné Benedicty 2008a: 140, 2008b: 256;
Kozár & Walter, 1985: 73; Miller & Gimpel, 1996: 605; Morrison & Morrison, 1966:
1; Ouvrard & Kozár, 2009: 101; Schmutterer, 2008: 77; Tang & Hao, 1995: 454,
Tereznikova 1981: 15; Williams, 1985a: 358 (Miller et al., 2013).
Acanthococcus Kiritchenko 1936: 156 is a junior homonym of Acanthococcus Signoret 1875

and is currently placed in the Pseudococcidae with the replacement name Spinococcus
Borchsenius, 1948: 953.
Thekes Maskell, 1892: 28. Type species: Acanthococcus multispinus Maskell, 1892: 217.
Synonymy by Ferris, 1955: 94. According to our concept this genus and species not
congeneric with Acanthococcus and should be re-established.
Acanthococcus Signoret, 1875: 35. Borchsenius, 1948: 501. Revived combination.
Acanthococcus Signoret, 1875: 34. Type species: Acanthococcus aceris Signoret, 1875: 35.
Placed on official list by Opinion 1203, Melville, 1982: 95.
Acanthococcus Signoret, 1875. Synonymy with Eriococcus by Miller & Gimpel, 2000: 105.
Acanthococcus Signoret, 1875. Kozár & Konczné Benedicty, 2008a: 128. Revived status.
Description
Ovisac ovoid, felt-like, white or gray, completely enclosing female body. Adult female
elongate-oval, narrowed posteriorly, with anal lobes conical and normally heavily
sclerotized, sometimes strongly nodulose with sclerotized teeth on inner margin, antennae
6, 7, rarely 8 segmented; frontal lobes or tubercles present, labium 3 segmented, with
16 setae (of these 10 on apical segment); with well-developed segments and a weakly-
developed basal segment with two pairs of hair-like setae, the outer one about half length
of the inner ones. Stylet short or very long, forming a double loop; legs well developed,
midcoxae and hindcoxae often with spinulae on anterior surfaces, translucent pores
usually absent, inner side of hind tibia with only four setae (medial setae absent), claw
usually with a denticle; tarsal and claw digitules longer than claw, knobbed, spiracles often
with a few associated disc pores; disc pores on venter only, most usually quinquelocular,
but the number of loculi vary between 3 to 9; oval disc pores (or cruciform pores)
absent from dorsum, but often on ventral side of prosomal in a marginal band; tubular
ducts of two types: micro- and macrotubular ducts present; microtubular ducts slender,
7 µm long, 2 µm wide, with bifid orifice, scattered or form transverse rows or bands on
dorsum, often associated with dorsal conical setae; macrotubular ducts often of 2 or
Acanthococcidae 73
3 sizes, usually forming transverse rows or bands on body surfaces; larger ducts each
with inner ductule longer than main ductule and with a flower-shaped terminal gland;
and smaller ducts each with short filamentous ductule. Enlarged conical setae normally
present at least in dorsal margin, but often on entire dorsum where they form transverse
bands or rows; hair-like setae on venter only; anal ring well developed, sclerotized with
partly double row of pores and 8 rarely 6 anal ring setae as long as apical seta on anal
lobes; each anal lobe with a long apical seta and usually with three short dorsal enlarged
conical setae, seldom with more, but at least with two, ventral hair-like setae also present,
suranal setae hair-like, cauda (median dorsal plate) usually well seen (Borchsenius, 1949;
Hodgson & Miller, 2010; Kosztarab & Kozár, 1988; Kozár & Konczné Benedicty, 2008a,
b).
In a comparative table (Fig. 29) we studied differences of enlarged setae variations
in females and first-instar nymphs of some species belonging to the genus Acanthococcus.
The type of enlarged setae are quite similar, what indicates the grouping of these
species in one genus. However, in this simple comparison can be seen great variation
and differences of number of spines, size and shape of these species. For example A.
aceris has truncated spines in the female, but pointed in the nymphs, A. uvaeursi in both
stages has pointed spines. A. kilinceri has a wide range of sizes of setae on dorsum of
females, in other cases, as A. melnikensis and A. aceris only one size category is present.
The greatest variation in spine size variation on female margins could be found in A.
glanduliferus. However, the largest variation of number of size categories and numbers of
rows of mid-dorsal longitudinal spines in the first-instar nymphs can greatly help in the
identification of species.
Disrtibution
With world-wide distribution, Kozár (2009) in the World list cited about 120 species in
this genus, however, some of them will be transferred to other genera in this work, but
others from the questionable “Eriococcus” sensu lato may belong to the genus Acanthococcus.
The composition of species in several regions needs revision. In the Palearctic Region,
37 species are listed. Except A. azaleae which is known in the Nearctic Region, all other
species seem to be restricted to the Palaearctic Region.
Biology
They feed on woody plants, usually specific to one part of the plant, for example A. aceris
is found at the bases at the forking of the twigs, while other species are found on the
bark, or in bark crevices. Most species have one generation per year; usually overwinter
in egg or second nymphal stage; males present.
Figure 29. Acanthococcus setae range on adult female and first instar.
Acanthococcidae 75
Key to species
Notes: The species A. azumae, A. chabohiba, A. shiraiwai could not be included into the
key because of the absence of necessary information. Even sometimes the position of
these species at the genus level was not clear, however, under comments of the species
we listed some possible similarities.
1. – Enlarged dorsal setae on abdominal segment VIII present.........................................2

– Enlarged dorsal setae on abdominal segment VIII absent.........................................18
2. – Enlarged setae on dorsum sharply pointed.....................................................................3
– Enlarged setae on dorsum blunt or truncated................................................................4
3. – Body elongate, enlarged dorsal tubular ducts on abdominal segment VIII
numbering 10-12........................................................................................... A. betulaefoliae
– Body round, enlarged dorsal tubular ducts on abdominal segment VIII
numbering 2-4.....................................................................................................A. abeliceae
4. – Number of enlarged setae on margin of abdominal segment VII only two.............5
– Number of enlarged setae on margin of abdominal segment VII more than two..9
5. – Enlarged setae on dorsum truncated............................................................... A. spiraeae
– Enlarged setae on dorsum blunt or pointed...................................................................6
6. – Venter of thorax without quinquelocular pores and tubular ducts............ A. tokaedae
– Venter of thorax with high number of quinquelocular pores and tubular ducts.....7
7. – On abdominal segment VII dorsal enlarged setae numbering
about 30......................................................................................................... A. koelreuterius
– On abdominal segment VII dorsal enlarged setae numbering between 11-20.........8
8. – Enlarged setae on dorsum of various sizes, dorsal setae much smaller
than marginal setae and reducing in size on the last abdominal segments
IV-VIII.................................................................................................................. A. ribesiae
– Enlarged setae on dorsum of various sizes, dorsal setae about half
the size of marginal setae and not reducing in size on the last abdominal
segments IV-VIII............................................................................................ A. castanopus
9. – Cruciform pores in groups on abdominal segments, and thorax, feeding on
Quercus spp.........................................................................................................................10
– Cruciform pores not in groups on abdominal segments, and thorax, feeding on
other trees..........................................................................................................................14
10. – Marginal and dorsal enlarged setae truncated...............................................................11
– Marginal and dorsal enlarged setae pointed..................................................................12
11. – Marginal enlarged setae on abdominal segment VII numbering 3................ A. roboris
– Marginal enlarged setae on abdominal segment VII numbering 5.........A. melnikensis
12. – Tubular ducts on venter in 2 of sizes......................................................A. macodeniensis
– Tubular ducts on venter in 3 of sizes............................................................................13
13. – Cruciform pores on submarginal of venter in large group, present on submedial

area of thorax, absent on head .......................................................................A. kılınceri
– Cruciform pores on submarginal of venter not in large group, present on
submedial area of thorax, present on head ....................................................... A. aceris
14. – Some of the enlarged setae on middorsum capitated.......................................A. salicis
– Capitated enlarged setae on dorsum absent..................................................................15
15. – Number of enlarged setae on abdominal segment VIII more than two.................16
– Number of enlarged setae on abdominal segment VIII two.....................................17
16. – Middorsal enlarged setae in longitudinal bands, trilocular pores absent
on venter.............................................................................................................A. ulmarius
– Middorsal enlarged setae not in longitudinal rows, some trilocular pores present
on venter.................................................................................................................A. populi
17. – Larger macrotubular ducts on mid venter, cruciform pores in bands on
abdominal segments III-VIII, on Populus spp............................................A. armeniacus
– Larger macrotubular ducts not on mid venter, cruciform pores in bands on
abdominal segments VII-VIII, on Castanae sp.............................................. A. rosannae
18. – Enlarged setae on middorsum forming large groups.............................A. kaschagariae
– Enlarged setae on middorsum not forming large groups...........................................19
19. – Two enlarged setae on the margin of each abdominal segment................................20
– More than two enlarged setae on the margin of each abdominal segment.............27
20. – Marginal setae sharply pointed........................................................................................21
– Marginal setae blunt..........................................................................................................26
21. – Dorsal enlarged setae much shorter than marginal enlarged setae,.................A. thymi
– Dorsal enlarged setae various sizes, some of them as long as marginal enlarged
setae....................................................................................................................................22
22. – Dorsal enlarged setae on abdominal segments VI and VII absent................ A. bezzii
– Dorsal enlarged setae on abdominal segments VI and VII present..........................23
23. – Dorsal enlarged setae on abdominal segment VII more than ten ..........A. isacanthus
– Dorsal enlarged setae on abdominal segment VII less than ten................................24
24. – Spiracles heavily sclerotized, enlarged setae on venter in wide submarginal band
on thorax................................................................................................................. A. onuki
– Spiracles not heavily sclerotized, enlarged setae on venter not in wide
submarginal band on thorax...........................................................................................25
25. – Macrotubular ducts on venter of three sizes, quinquelocular pores scattered all
over venter...........................................................................................................A. uvaeursi
– Macrotubular ducts on venter, of one size, quinquelocular pores concentrated on
abdominal segments...........................................................................................A. acericola
26. – Hind coxae and femur with pores, cruciform pores on venter scattered in mid
thorax, some of ventral setae capitated.................................................... A. glanduliferus
– Hind coxae and femur with pores, cruciform pores on venter only on submarginal
area, all ventral setae flagellate...........................................................................A. altaicus
Acanthococcidae 77
27. – Anal lobe and cauda strongly sclerotized, fused.......................................A. corniculatus

– Anal lobe and cauda not strongly sclerotized and fused.............................................28
28. – Middorsal enlarged setae in longitudinal bands, venter of thorax without
tubular ducts.........................................................................................................A. costatus
– Middorsal enlarged setae not in longitudinal bands, venter of thorax with
tubular ducts......................................................................................................................29
29. – Marginal enlarged setae on abdominal segment VII numbering
more than four..................................................................................................................30
– Marginal enlarged setae on abdominal segment VII numbering
less than four.....................................................................................................................31
30. – Dorsal and marginal enlarged setae sharp pointed, trilocular pores on
venter absent....................................................................................................A. transversus
– Dorsal and marginal enlarged setae pointed, trilocular pores on venter
present...................................................................................................................... A. ericae
31. – Cauda narrow, elongated, with a great group of quinquelocular pores
anterior to spiracles.................................................................................................A. sasae
– Cauda normal, not elongated, without a great group of quinquelocular
pores anterior to spiracles...............................................................................................32
32. – On abdominal segment VII dorsal enlarged setae numbering 10–12, enlarged
setae on dorsum of different shapes (from pointed to truncated).............. A. azaleae
– On abdominal segment VII dorsal enlarged setae numbering 12, enlarged
setae on dorsum not different shapes...........................................................................33
33. – Tubular ducts on venter of one size, enlarged setae on abdominal
segments VI-VII more than 16.......................................................................... A. latialis
– Tubular ducts on venter of two sizes, enlarged setae on abdominal
segments VI-VII around 16.......................................................................A. lagerstromiae
Acanthococcus abeliceae (Kuwana, 1927) (Fig. 30) Combination nova

Eriococcus abeliceae Kuwana, 1927: 111.
Syntypes: Female. Japan (Honshu, Kyoto City, Imperial Palace Grounds), on Abelicea
hirta, ?.v.1924, by S. Iwai. Deposited in ITLJ.
Lit.: Danzig, 1975: 44; 1977b: 50; 1980: 207; Hoy, 1963: 66; Kawai, 1980: 127; Koteja
& Zak-Ogaza, 1981: 513; Kozár & Walter, 1985: 73; Köhler, 1998: 372; Takahashi,
1957: 7; Tang & Hao, 1995: 31, Tao, 1999: 31; Wang, 2001: 208 (Miller et al., 2013).
Acanthococcus abeliceae, Kozár & Walter 1985: 74. Change of combination.

Eriococcus abeliceae; Miller & Gimpel, 2000: 111. Revived combination.
Acanthococcus abeliceae; Kozár, 2009:91. Revived combination.
Description
Unmounted female
Adult female dark purple, oval or broadly oval. Rounded anteriorly and gradually
narrowed posteriorly. Ovisac white or greyish white, elliptical in outline, with transverse
lines of depressions corresponding to the segments of the body of the insect and a
longitudinal median depression.
Mounted female
Mounted female oval, 2.2–2.3 mm long, and 1.2–1.9 mm wide. Anal lobes conical,
generally sclerotized. Antennae 7 segmented, II: 45 µm, III: 48 µm, IV: 38 µm, V: 25
µm, VI: 25 µm and VII: 35 µm long; with many long hair-like setae. Frontal lobes not
mentioned. Anal lobe with a 3 dorsal enlarged setae.
Venter: Labium, rather small, well sclerotised. Legs well developed, posterior legs
slightly larger, coxa: 125 µm, trochanter: 70 µm, femur: 130 µm, tibia: 90 µm, tarsus:
117 µm, claw: 25 µm long. Inner side of hind tibia probably with only four setae, claw
with a denticle; tarsal and claw digitules knobbed, longer than claw. Spiracles with a few
associated disc pores; discodial pores quinquelocular; few cruciform pores on ventral
margin present. Macrotubular ducts of two sizes. Hair-like setae on venter only; ventral
hair-like setae a1so present on anal lobes, suranal setae hair-like.
Dorsum: Enlarged conical setae normally present on dorsal margin, 2 or 3 on each
segment, on dorsum the setae form loose rows or bands, strong spine-like form. Two
setae present also in the middle of abdominal segment VIII. Macrotubular ducts
numerous. Microtubular ducts narrow, long. Anal ring well developed, with 8 anal ring
setae, each 105 um long, apical seta on anal lobes 157 µm long. Cauda not mentioned
(Kuwana, 1927; Tang & Hao, 1995).
Acanthococcidae 79
Figure 30. Acanthococcus abeliceae (Kuwana, 1927), female. After Tang & Hao (1995)
with modifications.
Ecology
Host plant: Abelicea hirta, Buxus microphylla, B. sinica, Ulmus sp.
Distribution: China, Japan.
Biology: Unknown.
Acanthococcus acericola (Tang & Hao, 1995) (Fig. 31) Combination nova
Eriococcus acericola Tang & Hao, 1995: 454.
Holotype: Female. Ningxia, on Acer truncatum, 30.ix.1963. By original designation.

Deposited in EISC.
Lit.: Tang & Hao, 1995: 452; Tao, 1999: 31; Wang, 2001: 208 (Miller et al., 2013).
Acanthococcus acericola; Miller & Gimpel, 1996: 598. Change of combination.

Eriococcus acericola; Miller & Gimpel, 2000: 112. Revived combination.
Acanthococcus acericola; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Unknown.
Mounted female
Adult female elongate-ovoid. 1.8 mm long, 0.9 mm wide. Antennae 7 segmented, the
size of the segments: I: 22.5 µm, II: 32.5 µm, III: 45 µm, IV: 25 µm, V: 25 µm, VI:
25 µm, and VII: 37.5 µm long; each segment covered with a few, strong hair-like setae
(except third segment); apical segment long with apical seta and 3 sensory falcate setae,
falcate sensory setae on two preapical segments. Frontal tubercle present. Eyes situated
on venter near margin. Anal lobes conical and normally heavily sclerotized, sometimes
strongly nodulose with sclerotized teeth on inner margin, each anal lobe with a long
apical seta and usually with 3 short enlarged conical setae.
Venter: Mouthparts well developed. Legs well developed, inner side of hind tibia with
only four setae, tibia shorter than tarsus, claw with a denticle; tarsal and claw digitules
clavate, longer than claw. Hair-like setae on venter only and suranal setae long hair-like.
Some quinquelocular pores present on last abdominal segments and head. Cruciform
pore present on ventral margin. Macrotubular ducts scattered on the venter, narrower
than on dorsum.
Dorsum: Enlarged conical setae normally present on dorsal margin, 2 on each segment,
on dorsum setae few, strong spine-like in shape, as on margin, spine on abdominal
segment 8 segment absent. Macrotubular ducts scattered on dorsum. Microtubular ducts
narrow, elongated, present on dorsum. Anal ring well developed, sclerotized, with 8 anal
ring setae, each as long as apical seta on anal lobes. Cauda present (Tang & Hao, 1995).
Acanthococcidae 81
Figure 31. Acanthococcus acericola (Tang & Hao, 1995), female. After Tang & Hao (1995)
with modifications.
Ecology
Host plant: Acer truncatum.
Distribution: China.
Biology: Unknown.
Acanthococcus aceris Signoret, 1875 (Fig. 32)

Acanthococcus aceris Signoret, 1875: 35. Unknown type status. Described: female.
Type material: "No original material has been traced. The specimens from Austria,
one of the type-localities, are from the Naturhistorisches Museum, Vienna, identified
by F. Löw, and the specimens from Switzerland are from the collection of P. Marchal.
There seems to be no doubt about the identity of the species now recognized as such
by many modern workers (Williams, 1985a)." Matile-Ferrero & Danzig visited the
Naturhistorisches Museum, Vienna in June of 1997 and were unable to locate any
type material of this species.
Additional material examined: Hungary, Selcepuszta, female on Acer campestre, C. P.
Malumphy, F. Kozár, 06. vii. 1989 (PPI: 9291); Austria, Vienna, 3 females on Acer sp.,
F. Kozár, 30. vii. 2009 (PPI: 8844); Hungary, Budapest, female on Acer campestre, F.
Kozár, 11. v. 2005 (PPI: 4664); Italy, Padova, 2 females on Acer campestre, F. Kozár,
05.v.1998 (PPI: 4992); Hungary, Igmánd, 2 females on Acer campestre, F. Kozár, 09. vii.
2007 (PPI: 7985).
Common name: Maple felt scale.
Lit.: Borchsenius, 1949: 131; Danzig, 1980: 3; Dziedzicka, 1977: 59; Fernald, 1903:
70; Gavrilov, 2010: 38; Cook & Gullan, 2001: 95; Hodgson & Miller, 2010: 7;
Hodgson & Trencheva, 2008: 12; Kosztarab, 1959: 402; Kosztarab & Kozár, 1988:
278; Koteja, 1974a: 296; 1974b: 76; 1976: 272; 1988b: 534; 2000: 172; Koteja & Zak-
Ogaza, 1979: 674; Kozár, 2009: 91; Kozár et al., 2013: 55; Kozár & Kosztarab, 1982:
204; Köhler, 1998: 372, Lindinger, 1912: 54; Rogojanu, 1966: 324; Schmutterer, 1952:
378; Tranfaglia & Esposito, 1985: 116; Tereznikova; 1981: 15, Tsalev, 1968: 207;
Zahradník, 1972:121 (Miller et al., 2013).
Eriococcus aceris; Cockerell, 1896: 323. Change of combination.

Nidularia aceris; Lindinger, 1933a: 108. Change of combination.
Acanthococcus aceris; Borchsenius, 1938: 131. Revived combination.
Eriococcus aceris; Hoy, 1963: 66. Revived combination.
Acanthococcus aceris; Kosztarab & Kozár, 1978: 65. Revived combination.
Eriococcus aceris; Miller & Gimpel, 2000: 113. Revived combination.
Acanthococcus aceris aceris, Kozár & Konczné Benedicty, 2008a. Revived status.
Eriococcus aceris kurdica; Bodenheimer, 1943: 21-22. Elevated here to species rank and
transfered to Rhizococcus genus.
Eriococcus melnikensis; Gavrilov, 2010: 38. Synonymy by Acanthococcus aceris. Notes: We
Acanthococcidae 83
studied all the Acanthococcus species which are related to A. aceris living on Quercus spp.
and other plants in the Palaearctic Region; concerning to first instar nymphs, there are
big differences between first instar nymphs of A. aceris and A. melnikensis which clearly
belongs to the Acanthococcus roboris group. On this base we reestablished here A. melnikensis
species status.
Description
Unmounted female
Adult female oval, chestnut colored, covered with white wax powder, 2.2 mm long, 1.5–
1.7 mm wide. Ovisac, felt-like test cream-white compact, greyish, up to 4 mm long, 2.2
mm wide.
Mounted female
Adult female broadly oval, largest specimens 2.75–3.75 mm long, 1.64–2.2 mm wide.
Antennae 7, rarely 6 segmented, 249–330 µm long; length of segments: I: 60.0–72.5
µm, II: 40–55 µm, III: 57.5–60.0 µm, IV: 47.5–57.5 µm, V: 27.5–30.0 µm, VI: 25.0–27.5
µm, and VII: 42.5–47.5 µm long, each segment covered with a few, strong hair-like setae
(except third segment); apical segment with apical seta 55–60 µm long; apical segment
also with 3 sensory falcate setae, each 25–37.5 µm long; segment VI with 1 sensory falcate
seta 25–30 µm long, segment V with 1 sensory falcate seta 22.5–27.5 µm long. Frontal
lobes developed but smaller than basal antennal segment and frontal tubercle present.
Eyes situated on venter near margin. Anal lobes strongly developed, about twice as long
as wide, sclerotised, each with 3 enlarged setae, each 45–60 µm plus 3–5 microtubular
ducts on dorsal surface; apical seta 155–200 µm; ventrally each with two flagellate setae,
ventral hair-like subapical seta 52.5–75.0 µm long and a suranal seta, latter longest but
shorter than anal ring setae, each 100–124 µm long.
Venter: Labium 145–170 µm long, 100–120 µm wide, shorter than clypeolabral shield,
basal segment with 2 pairs of setae, stylet loop extending to base of mesothoracic legs.
With normal flagellate setae in median areas, each short and slender but stouter and stiff
in lateral areas. Apical labial segment with 5 pairs of hair-like setae, with the median
pair somewhat shorter, median setae on apex of labium long, 20.0–22.5 µm long. Legs
well developed; lengths of segments and digitules of prothoracic legs; coxa: 65–80 µm,
trochanter: 57.5–65 µm, femur: 130–135 µm, tibia: 110 µm, tarsus: 127.5–140.0 µm and
claw: 27.5–30.0 µm, trochanther + femur: 185–195 µm, tibia + tarsus: 230.0–247.5 µm,
tarsal digitules: 50–55 µm, claw digitules: 25–30 µm long; length of segments and digitules
of mesothoracic legs; coxa: 65–85 µm, trochanter: 55–75 µm, femur: 120–135 µm, tibia:
115–120 µm, tarsus: 135 µm and claw: 30–35 µm, trochanther + femur. 185–190 µm,
tibia + tarsus: 255–260 µm, tarsal digitules: 52.5–55.0 µm, claw digitules: 30 µm long;
lengths of segments and digitules of metathoracic legs; coxa: 80–90 µm, trochanter: 60
µm, femur: 137.5–140.0 µm, tibia: 120–130 µm, tarsus: 145–155 µm and claw: 32.5–35.0
µm, trochanther + femur: 190–195 µm, tibia + tarsus: 270–280 µm, tarsal digitules: 55–
60 µm, claw digitules: 30–32.5 µm long. Meso and metathoracic coxae each with spinulae
on ventral surface. Tibiae of meso and metathoracic legs each with 4 setae (median seta
absent), tibiae of prothoracic legs each with 5 setae (median seta present), tarsi each with
6 setae. Length of spiracles 55.0–62.5 µm; diameter of spiracular peritreme 27.5–35.0 µm,
posterior spiracles slightly larger than anterior. Setae in median areas flagellate, long, each
17.5–75.0 µm long, setae in lateral areas spine like, each 17.5–20.0 µm long. Multilocular
pores each 3–5 µm in diameter and with 5 loculi (very few number of pores with 3 loculi),
distributed in sparse bands on all abdominal segments and scattered on thorax and head.
Macrotubular ducts of three sizes, smallest macrotubular ducts each 2.5–3.0 µm wide
and 10.0–15.0 µm long, present on median areas of abdominal segments, medium
macrotubular ducts each 5–7 µm wide and 22.5–25.0 µm long, present on median areas
of abdominal segments, larger macrotubular ducts each 7.5–10.0 µm wide and 25–30 µm
long, in a submarginal band on abdomen, thorax and head. Cruciform pores generally on
submedian area in a band on thorax and abdominal segments and a few on head, each
2.5–3.0 µm in diameter.
Dorsum: Strongly nodulose; with spinose setae of two sizes; marginal enlarged setae long,
blunted, each 40.0–57.5 µm forming a marginal row and differentiated from dorsal setae,
other dorsal setae conical, blunted, 25–35 µm long; arranged in transverse rows across
each body segment, rows irregular on head, with 6-8 enlarged setae on the abdominal
segment VIII. Macrotubular ducts, each 7.5–10.0 µm wide and 25–30 µm long, scattered
throughout dorsum, generally in segmental bands. Microtubular ducts long, each 10–13
µm long with oval or bifurcated dermal orifice, scattered over dorsum. Anal ring strongly
sclerotized, with 19-21 pores on each side, pores with spine-like spiculae 55–70 µm in
diameter, with 8 setae, each 125–155 µm long; anal ring situated on margin of dorsum.
Cauda triangular, slightly sclerotised (after Hodgson & Trencheva, 2008; Kosztarab &
Kozár, 1988; Williams, 1985a; with modifications).
Other stages
Mounted first instar nymph (Fig. 32, top left).

Body of slide-mounted specimens, oval. Antennae six segmented, apical three segments
with strong sensory setae as in adult female. Frontal tubercle present. Dorsum with
strong spines with pointed apices of one size, forming middorsal longitudinal pair of
rows, the submedian row contains only four large spines, followed by five very small
setae. Microtubular ducts long, few in two middorsal longitudinal rows on both sides.
Venter with transverse rows of six small hair-like setae on each abdominal segment;
median setae longer than others. Cruciform pores present on thoracic segments, with
one pair of quinquelocular pores on each thoracic segment, one near each spiracle, plus
one pair on frons, and two longitudinal submedian rows on abdomen. Anal ring with 6
spine-like setae (after Hodgson & Trencheva, 2008).
Acanthococcidae 85
Figure 32. Acanthococcus aceris Signoret, 1875, female, original. First instar nymph on top
left, after Hodgson & Trencheva (2008).
Ecology
Host plant: Acer campestre, A.cincinatum, A. platanoides, A. pseudoplatanus, Aesculus
hippocastanum, Carpinus betulus, Elaeagnus angustifolia, Fagus sylvatica, Malus sp., Myrtus
communis, Platanus orientalis, Pyrus sp., Quercus pubescens, Q. robur, Salix caprea, Ulmus campestris.
Distribution: Austria, Bulgaria, Croatia, former Czechoslovakia, Cyprus Island, France,
Germany, Hungary, Iraq, Italy, Moldova, Netherlands, Poland, Romania, Russia, Slovenia,
Switzerland, Turkmenistan, Ukraine, former Yugoslavia.
Biology: Feed on trees; ovisacs usually found at base of twig forks, or between the
corky bark strips and crevices of trunk. Bisexual; overwintering as second-instar nymphs;
according to Schmutterer (1952), adults develop by the first half of April and complete
their egg laying by the end of May; lay about 82–378 red eggs per female; eggs hatch in
about 30–35 days; first-instar nymphs feed on leaves, from where they return to the bark
for overwintering in September or October. In Hungary, all eggs hatched by mid June
1981 in Budapest. Rarely a pest in urban environment. The species is found in mountains
at up to 1.500 m elevation (Zahradník, 1972).
Acanthococcidae 87
Acanthococcus altaicus Matesova, 1967 (Fig. 33) Redescription

Acanthococcus altaicus Matesova, 1967: 1193.
Holotype: Female. Kazakhstan (Eastern Kazakhstan Oblast, Ubinski Ridge, Kirov

District, Orlovka), by original designation. Deposited in the ZMAS. Notes: Holotype
adult female mounted alone on slide, also 4 additional female paratypes on 4 slides
(Danzig, personal communication, 1996).
Lit.: Koteja & Zak-Ogaza, 1981: 513; Kozár & Walter, 1985: 73; Tang & Hao, 1995:
451 (Miller et al., 2013).
Eriococcus altaicus; Tang & Hao, 1995: 455. Change of combination.

Acanthococcus altaicus; Köhler, 1998: 372. Revived combination.
Eriococcus altaicus; Miller & Gimpel, 2000:123. Revived combination.
Acanthococcus altaicus; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female oval, chestnut coloured, covered with white wax powder, 2.2 mm long,
1.5–1.7 mm wide. Ovisac compact, milk-white colour, felted with waxy needles.
Mounted female
Adult female oval, Antennae 7 segmented, length of segments: I: 40 µm, II: 54 µm,
III: 46.9 µm, IV: 43.2 µm, V: 27 µm, VI: 21.6 µm and VII: 37.8 µm long. Frontal lobes
developed but smaller than basal antennal segment. Eyes situated on venter near margin.
Anal lobes protruding, about twice as long as wide, apically rounded, moderately
sclerotised, serrate on inner margin, with 3 dorsal conical setae, ventrally each with two
flagellate setae, one subapical, one on outer base and a suranal seta, latter longest but
shorter than anal ring setae and with long apical setae.
Venter: Labium 3 segmented, basal segment with 2 pairs of setae; with normal flagellate
setae in median areas, each short and slender, but stouter and stiff in lateral areas. Legs
well developed; lengths of segments of legs; coxa: 157 µm, femur: 119 µm, tarsus: with
claw: 157 µm long, claw with denticle; hind coxa without translucent pores, with spinulaes
on anterior side. Quinquelocular pores in transverse bands across abdominal sternites,
around margins to head, and on mid venter. Macrotubular ducts of two sizes; larger
tubular ducts about half-the size of dorsal enlarged ducts, not numerous, in more or less
single rows on abdominal segments and around submargins on head; narrower ducts,
about half size of larger ducts, present in small numbers across abdominal segments.
Microtubular ducts absent. Cruciform pores present in submarginal area from head to
abdominal segment IV.
Dorsum: Enlarged setae of three sizes, all with more or less straight sides tapering to a
truncate tip, in profile curved and bluntly thorn-like. Largest setae each 41–51 µm long
on margins of posterior segments; shortest setae more numerous, each about 17–23 µm
long on abdominal segments on each side forming a group of 2 large and 2–3 medium-
sized setae, other dorsal setae forming sparse, transverse segmental bands. On seventh
segment number of spines 12, on abdominal segment 8 spines absent. Macrotubular
ducts of one size scattered among spines. Microtubular ducts narrow, long, elongate,
with oval orifice; evenly distributed; forming transverse bands on dorsum of prosoma,
rows on abdomen. Anal ring oval, sclerotized, with two rows of pores with 8 setae each
about 132 µm long. Cauda triangular, slightly sclerotised, strongly nodulose (Matesova,
1967, with modifications based on type material).
Ecology
Host plant: Salix sp.
Distribution: Kazakhstan.
Biology: Lives in crevices on the bark of its host, density was very high. Lays lilac colored
eggs in mid June, between 76-116 eggs. Nymphs appear at end of June. Phyloxerina salicis
Linnaeus lives in a symbiotic relation with A. altaicus, 80% of egg sacs of the scale was
infested with Phyloxerina (Matesova, 1967).
Acanthococcidae 89
Figure 33. Acanthococcus altaicus Matesova, 1967, female. Original.

Acanthococcus armeniacus (Tang & Hao, 1995) (Fig. 34)

Eriococcus armeniacus Tang & Hao, 1995: 456.
Holotype: Female. China (Ningxia), on Prunus armeniaca, 18.x.1983. By original

designation. Deposited in EISC.
Lit.: Miller & Gimpel, 1996: 598; Tang & Hao, 1995: 451; Tao, 1999: 31; Wang, 2001:
208 (Miller et al., 2013).
Acanthococcus armeniacus; Miller & Gimpel, 1996: 598. Change of combination.

Eriococcus armeniacus; Miller & Gimpel, 2000: 134. Revived combination.
Acanthococcus armeniacus; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female spindle shaped, about 1.8 mm long, 0.8 mm wide.
Mounted female
Antennae 6 segmented (sometimes 7 segmented), length of segments: I: 57 µm, II: 43
µm, III: 75 µm, IV: 40 µm, V: 25 µm, VI: 20 µm and VII: 40 µm long, if six segmented
the third and fourth segments fused. Frontal lobes present. Eyes present. Anal lobes large
and sclerotized, with inner margins serrated, with three spines on dorsal side.
Venter: Mouth parts well developed, stylet loop very long, much longer than the body.
Legs medium size, tibia shorter than tarus, tarsal and claw digitules longer than claw,
with expanded tip, claw with denticle. Quinquelocular pores numerous on abdominal
segments, head and submarginal area of thorax, and cruciform pores in band on
submargin. Macrutubular ducts of two sizes; larger one as those on dorsum on margin
and thorax and head; narrower ones on abdominal segments.
Dorsum: Dorsum with spines of three sizes, the large and median size ones distributed in
marginal and medial longitudinal bands, medial band with 4 and marginal bands with 3–6
spines on each body segment. Two spines on abdominal segment VIII and a transverse
band of spines on abdominal segments VII. Macrotubular ducts of one size, scattered
among enlarged dorsal spines. Microtubular ducts present. Anal ring with pores and 8
setae. Cauda not mentioned (Tang & Hao, 1995 with modification).
Ecology
Host plant: Prunus armeniaca.
Biology: Unknown.
Acanthococcidae 91
Figure 34. Acanthococcus armeniacus Tang & Hao 1995, female. After Tang & Hao (1995)
with modifications.
Acanthococcus azaleae (Comstock, 1881) (Fig. 35)

Eriococcus azaleae Comstock, 1881: 338.
Lectotype: Female. USA (District of Columbia, in "Agr. Greenhouse"), on Azalea

sp., 06.i.1881, by D.C. Pergande. By subsequent designation Miller & Miller, 1992:
14-18. Deposited in USNM.
Common name: Azalea bark scale, eriococcus scale, spirea scale.
Lit.: Davids, 1974: 3; Fernald, 1903: 72, Gill, 1993: 156; Cook & Gullan, 2001: 95;
Hoy, 1963: 73; Johnson, 1982: 114; Kosztarab, 1996: 228; Koteja, 1974b: 76; 1976:
272; Koteja & Zak-Ogaza, 1981: 513; Köhler, 1998: 273; McDaniel, 1964: 103; Miller,
2005: 491; Miller & Miller, 1992: 3; 1993: 8; Williams, 1985b: 217; Zahradník, 1990:
16 (Miller et al., 2013).
Eriococcus borealis; Cockerell, 1899c: 369.

Syntype: Female. USA (Alaska, Dawson City). Deposited in USNM, and BMNH.
Synonymy by Miller & Miller, 1992: 14.
Nidularia borealis; Lindinger, 1933a: 108. Change of combination.
Nidularia azaleae; Lindinger, 1933a: 108, 117. Change of combination.
Eriococcus bezzii; Lindinger, 1943: 223. Incorrect synonymy; discovered by Tranfaglia &
Esposito, 1985: 121.
Acanthococcus azaleae; Borchsenius, 1949: 350. Change of combination.
Eriococcus azaleae; Ferris, 1955: 95. Revived combination.
Acanthococcus azaleae; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus azaleae; Miller & Gimpel, 2000: 137. Revived combination.
Acanthococcus azaleae; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Oval, posterior apex pointed, body dark red or purple. Ovisac pure white in color and
tapered.
Mounted female
Adult female, oval, 1.09–3.15 mm long, 0.61–2.06 mm wide. Antennae normally 7
segmented rarely with 8 segments, third or rarely fourth segment longest. Apical segment
with 4 sensory setae; second and third antennal segment from apex with one single
sensory seta. Frontal lobe present, large. Anal lobes strongly protruding, apically acute,
heavily sclerotized, with 0 to numerous teeth on mesal margin, each lobe dorsally with 3
enlarged setae (normally lateral seta largest or equal to posterior seta, anteromedial seta
smallest), with 3-7, usually 4, microtubular ducts; each lobe ventrally with 3 slender body
setae and rarely with a few quinquelocular pores.
Venter: Legs small; hind coxa and femur with spinulae on anterior surface, pores absent;
posterior 2 pairs of tibia with 4 setae, anterior pair with 5; inner apical tibial setae elongate,
Acanthococcidae 93
Figure 35. Acanthococcus azaleae (Comstock, 1881), female. After Miller and Miller
(1993) with modifications.
not enlarged; tarsi always longer than tibiae (hind tibia/tarsus ratio 0.61–0.77); claws with
conspicuous denticle near tip. Hair-like setae elongate; longest seta on abdominal segment
VII: 56–72 µm long, on segment II: 47–67 µm, medial setae apically acute. Enlarged setae
of small size only, present along body margin from abdominal segments V or IV up to
head. Multilocular pores of three kinds: 7-locular pores absent; quinquelocular pores
most numerous, variable in form, present over entire venter; trilocular pores sparse, more
numerous than septalocular pores, scattered over entire surface. Cruciform pores present
along lateral margin from abdominal segment V up to head. Macrotubular ducts of 2
sizes: larger size in small numbers along body margin; smaller size in reduced numbers
in medial or sublateral areas of abdominal segment VII through III, sparse on thorax,
absent on head. Microtubular ducts absent.
Dorsum: Enlarged setae normally of 2 sizes: 2 or 3 larger setae along margin of each
abdominal segment and on margin of thorax and head, additional large setae often
present in medial and sublateral lines, although sometimes absent; remaining setae small.
Largest enlarged seta 47–62 µm long, smaller setae 31–41 µm on abdominal segments VII
through II; longest large seta 1.2–1.8 times longer than longest small seta; setae straight,
with acute or rounded apices; all with thin setal rings, these setae in moderate numbers
over dorsum, e.g., abdominal segment IV with 22-35 large setae arranged in 3 pairs of
longitudinal lines (medial, sublateral, lateral), setae on abdominal segment VIII absent.
Macrotubular ducts present over dorsum. Macrotubular ducts numerous over dorsum.
Microtubular ducts, 10–12 µm long, with area farthest from dermal orifice sclerotized
and weakly divided into 2 parts, apical portion rounded, about one fourth length of
remaining sclerotized portion; total sclerotized portion much longer than unsclerotized
area; dermal orifice heavily sclerotized, with 2 protruding tubes. Anal ring usually ventral,
with 8, rarely 6, setae. Cauda (median dorsal plate) triangular, slightly sclerotized, strongly
nodulose (after Miller & Miller, 1993; with some modifications).
Comment
Acanthococcus azaleae is morphologically variable and therefore had been considered as 2
species (A. azaleae and A. borealis). The A. azaleae type was collected from Rhododendron
spp. from central Texas and east as well as from southern and Northern North America
(excluding Canada and Alaska). Specimens of the A. borealis type were not collected on
Rhododendron spp. and were from Utah west to California. Specimens collected from the
overlapping areas were intermediate in form, and the 2 species were therefore considered
to be 2 forms of 1 species (A. azaleae) (Miller & Miller, 1992). Gill (1993) treated A.
borealis as a separate species. Lindinger (1943) incorrectly considered this species to be a
senior synonym of A. bezzi (Leonardi).
Acanthococcidae 95
Ecology
Host plant: Acer sp., Azalea sp., Azalea hybrid ‘hino de giri’, A. indica, A. nudiflora, Celtis sp.,
Crataegus coccinea, Crataegus sp., Fremontodendron sp., Gaylussacia sp., Liquidambar sp., Populus
sp., Rhododendron catawbiense, Rhododendron sp., Ribes sp., Salix sp., Thuja sp., Vaccinium sp.,
V. macrocarpon.
Distribution: Canada, USA, Belgium, Germany, Russia.
Biology: Data indicates the species has one generation per year in colder places, but two
generations per year have been observed in warmer areas such as Alabama. This species
overwinters as eggs or first-instar nymphs. Females lay 50–250 reddish purple eggs
in spring (Kosztarab, 1996). On cranberries in Massachusetts the females were found
in crotches of the stems. In early June to early July 100 to 150 pink eggs were found
associated with the females in white fluffy sacs. Crawlers were present in late June and
early July and were light yellow unlike the red females (Franklin, 1952). On sprouts and
twigs. This species is a widespread economic pest, particularly on azaleas (Miller & Miller,
1992). Mentioned as a trouble some pest of azaleas in greenhouses (Franklin, 1952).
Acanthococcus azumae (Kanda, 1933) (Fig. 36)

Eriococcus azumae Kanda, 1933: 151.
Syntypes: Female. Japan (Honshu, Mt. Azuma), on Bambusa, 25.vii.1932, by S. Kanda.,

Deposited in KAYJ.
Lit.: Hoy, 1963: 74, Kanda, 1933: 151; Kawai, 1972: 5; Kawai, 1980: 129; Köhler,
1998: 373; Kozár & Walter, 1985: 74; Takahasi, 1957: 6; Tang & Hao, 1995: 450
(Miller et al., 2013).
Acanthococcus azumae; Kozár & Walter, 1985: 74. Change of combination.

Eriococcus azumae; Miller & Gimpel, 2000: 141. Revived combination.
Acanthococcus azumae; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female completely enclosed in a white ovisac. Body olivaceous brown, 3.05 mm
long.
Mounted female
Slide-mounted adult female with enlarged setae, conical, apices acute, abundant over
dorsum, marginal setae apparently larger than others, two spines present of the 8th
segment. Antennae seven segmented, length of segments: I: 39–48 µm, II: 41–43 µm, III:
43–44 µm, IV: 54–56 µm, V: 21–25 µm, VI: 24–28 µm and VII: 27–38 µm long. Tarsus
markedly longer than tibia, tarsal and claw digitules longer than claw, capitated. Anal ring
oval, with eight setae, subequal in length. Anal lobes well developed, sclerotized, with
long apical setae, and two stout spines on inner side and one on outer side. Cauda present
(Kanda, 1933).
Ecology
Host plant: Bambusa sp.
Distribution: Japan (Honshu).
Biology: Lives transversely across the axils of the leaves of bamboo, collected at the
end of July.
Comment
The species is similar to A. onukii, in which the fourth antennal segment is markedly
shorter than the third antennal segment.
Acanthococcidae 97
Figure 36. Acanthococcus azumae (Kanda, 1933), female. After Kawai (1980).
Acanthococcus betulaefoliae (Tang & Hao, 1995) (Fig. 37)

Eriococcus betulaefoliae Tang & Hao, 1995: 460.
Holotype: Female. China (Ningxia), on Pyrus betulaefolia, 12.ix.1983. By original

designation. Deposited in EISC.
Lit.: Tao, 1999: 31, Wang, 2001: 208.
Acanthococcus betulaefoliae; Miller & Gimpel, 1996: 598. Change of combination.

Eriococcus betulaefoliae; Miller & Gimpel, 2000: 144. Revived combination.
Acanthococcus betulaefoliae; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female ovoid, about 2.1 mm long, 1.2 mm wide.
Mounted female
Antennae 7 segmented, length of segments: I: 35 µm, II: 45 µm, III: 60 µm, IV: 60 µm,
V: 30 µm, VI: 20 µm and VII 50 µm long. Frontal lobes not mentioned. Eyes situated on
venter near margin. Anal lobes large and sclerotized, with inner margins serrated, with
three cylindrical spines on dorsal side.
Venter: Mouth parts well developed. Legs medium size, tibia shorter than tarsus, tarsal
and claw digitules longer than claw, with expanded tip, claw with denticle. Quinquelocular
pores and cruciform pores numerous in bands on abdominal segments, scattered on
thorax and head. Cruciform pores on submarginal area. Macrotubular ducts of 2 sizes,
larger ones on submarginal band, and narrower ones on middle part of abdominal
segments.
Dorsum: Enlarged setae of 2 sizes on dorsum, in general 30–40 µm long, scarcely about
10, distributed in bands, marginal bands with 3-5 spines on each body segment. With
16 spines on abdominal segment VIII and 26 of spines on abdominal segment VII.
Macrotubular ducts in one size, scattered through. Microtubular ducts scattered on derm.
Anal ring with pores in rows and 10 setae. Cauda not mentioned (Tang & Hao, 1995).
Ecology
Host plant: Pyrus betulaefolia.
Distribution: China (Ningxia =Ningsia).
Biology: Unknown.
Acanthococcidae 99
Figure 37. Acanthococcus betulaefoliae (Tang & Hao 1995), female. After Tang & Hao
Acanthococcus bezzii (Leonardi, 1907) (Fig. 38)

Eriococcus bezzii Leonardi, 1907: 148.
Lectotype: Female. Italy (Val Nerina, Sondrio, vaso 75 tubo no 7), on Rhododrendron
ferruguineum, by M. Bezzi. By subsequent designation Tranfaglia & Esposito, 1985: 121.
Deposited in IFSP.
Lit.: Goux, 1936: 352; Hoy, 1963: 75; Miller & Gimpel, 1996: 598; Ouvrard & Kozár,
2009: 101; Pellizzari & Kozár, 2011: 66; Tang & Hao, 1995: 451 (Miller et al., 2013).
Eriococcus uvaeursi; Lindinger, 1912. Misidentification; discovered by Leonardi, 1920.

Acanthococcus bezzii; Miller & Gimpel, 1996: 598. Change of combination.
Eriococcus bezzii; Miller & Gimpel, 2000: 144. Revived combination.
Acanthococcus bezzii; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female oval, 1.75–2.61 mm long, 1.12–1.82 mm wide.
Mounted female
Antennae strong, 7 segmented, 259–306 µm long, fourth segment longer than second
and third. Frontal lobes well developed. Eyes situated on venter near margin. Anal lobes
protruding, about twice as long as wide, apically rounded, moderately sclerotized, serrate
on inner margin, with 3 narrow, dorsal conical setae, 36.3–59.4 µm long, apical setae
152.4–188.7 µm long and with some microtubular ducts.
Venter: Labium 3 segmented, basal segment with a pair of setae; apical segment with
5 pairs of equal long setae; with normal flagellate setae in median areas, each short and
slender but stouter and stiff in lateral areas. Legs well developed: coxa-tarsus 188.8–324.5
µm, tibia 129.8–206.5 µm long, ratio of tibia and tarsus 0.65, claw with a denticle. Coxa
without translucent pores, but with spinules on anterior side. Discoidal pores scattered on
prosoma, cruciform pores scattered on margin; quinquelocular pores in transverse bands
across abdominal sternites. Few small macrotubular ducts present around submargins to
head. Some microtubular ducts present on margin.
Dorsum: Spine-like setae of 2 sizes, 47.1–61.7 µm long; with a truncated tip, 2 or 3
on margins of posterior segments; other dorsal setae form sparse, transverse segmental
bands; enlarged spines absent on abdominal segment VI, VII and VIII. Macrotubular
ducts one size, frequent on dorsum. Microtubular ducts narrow, long, elongate, with oval
orifice; evenly distributed. Form transverse bands on dorsum of prosoma, in rows on
abdomen. Anal ring oval, sclerotized, with two rows of pores with 8 setae, each about 132
µm long. Cauda not seen (Tranfaglia & Esposito (1985) with additons from lectotype).
Acanthococcidae 101
Figure 38. Acanthococcus bezzii (Leonardi 1907), female. After Tranfaglia & Esposito
Other stages
First-instar nymph described by Leonardi (1920).
Ecology
Host plant: Rhododendron ferrugineum.
Distribution: Germany, Italy, Spain.
Biology: Unknown.
Comments
According to Tranfaglia & Esposito (1985), Lindinger (1943) erroneously considered
Acanthococcus bezzii to be a junior synonym of A. azaleae. Lindinger (1912) incorrectly
considered A. bezzii to be a junior synonym of A. uvaeursi. Goux (1936 and 1936)
erroneously considered A. bezzii to be a junior synonym of A. bahiae = texanus. Later, in
1948, Goux incorrectly considered both A. bezzii and A. texanus to be junior synonyms
of A. uvaeursi. Information used to distinguish among A. bezzii, A. bahiae = texanus and
A. uvaeursi is given in the remarks of A. uvaeursi. We are considering all of these species
as distinct until more detailed studies could be undertaken. The authors of the present
work agree with the view of Tranfaglia & Esposito (1985).
Acanthococcus castanopus (Tang & Hao, 1995) (Fig. 39)

Eriococcus castanopus Tang & Hao, 1995: 463.
Holotype: Female. China (Guangxi, Long Sheng), on Castanopsis sp., 07.vi.1981. By

original designation. Deposited in EISC.
Lit.: Wang, 2001: 453; Tao, 1999: 32 (Miller et al., 2013).
Acanthococcus castanopus; Miller & Gimpel, 1996: 599. Change of combination.

Eriococcus castanopus Miller & Gimpel, 2000: 156. Revived combination.
Acanthococcus castanopus; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female ovoid, about 2.25–2.50 mm long, 1.10–1.45 mm wide.
Mounted female
Adult female ovoid. Antennae 7 segmented, length of segments: I: 35 µm, II: 32 µm,
III: 48 µm, IV: 30 µm, V: 20 µm, VI: 20 µm and VII: 30 µm long. Frontal lobe present.
Eyes situated on venter near margin. Anal lobes large and sclerotized, with inner margins
serrated, with three cylindrical spines on dorsal side.
Acanthococcidae 103
Figure 39. Acanthococcus castanopus (Tang & Hao, 1995), female. After Tang & Hao
Venter: Labium 3 segmented. Legs medium sized, tibia shorter than tarsus, tarsal and
claw digitules longer than claw, with expanded tip, claw with denticle. Quinquelocular
pores in row on abdominal segment, scattered on thorax and head. Cruciform pores
present on submarginal area. Macrotubular duct of 2 sizes; larger one in submarginal area
on abdominal segment, scattered on thorax and head.
Dorsum: Enlarged setae of three sizes on dorsum, distributed in bands, marginal bands
with 2 or 3 spines on each body segment; two spines on abdominal segment VIII and
around 14 of spines on abdominal segment VII. Tang & Hao (1995) mentioned only 10.
On anterior segments enlarged setae form two irregular transverse rows. Macrotubular
ducts one size, scattered through. Microtubular ducts long, scattered on body surface.
Anal ring with pores in rows and 8 hair-like setae. Cauda not mentioned (after Tang &
Hao, 1995, with modifications).
Ecology
Host plant: Castanopsis sp.
Distribution: China (Guangxi (=Kwangsi)
Biology: Unknown.
Acanthococcus chabohiba (Kuwana & Nitobe, 1918)

Eriococcus chabohiba Kuwana & Nitobe, 1918: 400.
Syntypes: Female. Japan, on Chamaecyparis obutosa (sic) var. breviramia. Deposited in

ITLJ. Notes: Five slides in UCDC.
Lit.: Hoy, 1963: 79; Kawai, 1972: 4; 1977: 151; 1980: 128; Köhler, 1998: 374;
Takahashi, 1957: 6; Tang & Hao, 1995: 453 (Miller et al., 2013).
Acanthococcus chabohiba; Kozár & Walter, 1985: 74. Change of combination.

Eriococcus chabohiba; Miller & Gimpel, 2000: 160. Revived combination.
Acanthococcus chabohiba; Kozár, 2009: 128. Revived combination.
Description
Unmounted female
With great, large, white eggsacs on braches of host plant.
Mounted female
Antennae 7 segmented. Legs normal, tarsus longer than tibia, tarsal and claw digitules
capitated, longer than claw. On the penultimate segment with 9 enlarged sharp pointed
spines. On margin 2 or 3 setae situated. Anal lobes elongated, serrated on inner side, with
three blunted spines. Anal lobe setae as long as anal lobe length. Anal ring with 8 pairs of
setae, as long as the lobe setae (after drawing of Kuwana & Nitobe, (1918) and Kawai, (1980).
Acanthococcidae 105
Other stages
Male is winged, typical for the genus.
Ecology
Host plant: Chamaecyparis obtusa var. breviramea.
Distribution: Japan.
Biology: On the young branches of its host, found in high density.
Comments
Hitherto, acanthococcids have not been found on Cupressaceae in the western Palaearctic
subregion.
Acanthococcus corniculatus (Ferris, 1950) (Fig. 40) Redescription

Eriococcus corniculatus Ferris, 1950: 7.
Holotype: Female. China (Yunnan Province, near Kunming, An-lin-wen-chian),

on Ternstroemia japonica var. wightii, 27.iv.1949, by G.F. Ferris. By original designation.
Deposited in UCDC.
Lit.: Borchsenius, 1960c: 916; Hoy, 1963: 82; Hua, 2000: 138; Tang & Hao, 1995:
450; Tao: 1999: 32; Wang, 1974: 329; 1982b: 41; 2001: 208; Yang, 1982: 101 (Miller
et al., 2013).
Proteriococcus corniculatus; Yang, 1982: 101. Change of combination.

Eriococcus corniculatus; Tang & Hao, 1995: 450. Revived combination.
Acanthococcus corniculatus; Miller & Gimpel, 1996: 600. Change of combination.
Eriococcus corniculatus; Miller & Gimpel, 2000: 175. Revived combination.
Acanthococcus corniculatus; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Ovisac usual type of the genus, white.
Mounted female
Adult female oval, 1.75–2.0 mm long, 1.2 mm wide. Antennae 7 segmented, length of
segments: I: 30 µm, II: 33 µm, III: 46 µm, IV: 35 µm, V: 22 µm, VI: 22 µm and VII: 30
µm long; third and fifth segments without hair-like setae, other segments with a few hair-
like setae; apical segment with apical seta 45 µm long; apical segment also with 3 sensory
falcate setae, longest 30 µm long; two preapical segments with one sensory falcate seta:
on segment V: 15, VI: 23 µm long. Frontal lobe present, frontal tubercle 3 µm wide.
Eyes situated on venter near margin. Width of anterior spiracles 21 µm. Anal lobes well
developed, sclerotized, with serrated distal margin, with 3 blunted enlarged spines, 36 µm
long, ventrally with two flagellate setae; anal lobes with 216 µm long apical setae.
Venter: Labium 3 segmented, 103 µm long; basal segment with 2 pairs of setae, apical
segment with 5 pairs of equal length hair-like setae. Legs well developed: prothoracic
legs: coxa 50 µm, trochanter 30 µm, femur 103 µm, tibia 67 µm long; mesothoracic legs:
coxa 50 µm, trochanter 30 µm, femur 100 µm, tibia 65 µm long; metathoracic legs: coxae
60 µm, trochanter 35 µm, femur 110 µm, tibia 78 µm, tarsus 117 µm; claw 28 µm long;
claw digitules 31 µm, tarsal and claw digitules slightly knobbed, longer than claw; meso
and metathoracic coxae with spinulae on anterior surface; claw with a denticle; legs with
a few hair-like setae; tibia with 4 setae (except prothoracic legs). Multilocular pores 5 µm
in diameter and with 3-5 loculi, mostly quinqueloculars; distributed in transverse bands
across abdominal sternites, in sparse rows on thoracic segments and head region; some
trilocular pores on abdomen. Macrotubular ducts of 2 sizes: larger macrotubular ducts
5–6 µm wide and 22 µm long, situated on margin; smaller macrotubular ducts 2 µm wide
and 17 µm long, in a submarginal band on the thorax, scattered in the head region and
on abdominal segments. Microtubular ducts sparse marginally and submarginally, some
submedially on thorax and head. Cruciform pores a few in submarginal area. A few
hair-like setae present on submedian venter, a few shorter hair-like setae submarginally.
Blunted enlarged spines on margin; half-sized as on the dorsum. Suranal setae hair-like.
Dorsum: The cuticular surface with nodulation. Enlarged setae blunt, conical, 24–43 µm
long, in a row on margins, 2 or 3 on segments, but absent from abdominal segment VIII.
Twelve dorsal setae on abdominal segment VII. Macrotubular ducts 5–6 µm wide and
22 µm long, scattered among enlarged setae. Microtubular ducts scattered, 13 µm long,
elongate with bifid orifice. Anal ring situated on margin of body; oval, sclerotized 42 µm
wide, 57 µm long, with one row of pores and with 8 setae 92 µm long. Cauda irregularly
toothed, 46 µm wide (after Ferris, 1950; present study: here we have redescribed and
redrawn the species from a paratype).
Ecology
Host plant: Camellia oleosa, Ternstroemia gymnanthera, T. japonica, T. japonica var. wightii.
Distribution: China (Yunnan).
Biology: On stem of the host plant.
Comments
Although the description and illustration of Ferris (1950) is quite good (Miller & Gimpel,
2000), for decision of the generic combination, some important characters are absent on
his drawing as frontal lobe, antennal tubercle, labial setae, cruciform pores, etc., thus the
authors felt better to present a new drawing.
Acanthococcidae 107
Figure 40. Acanthococcus corniculatus (Ferris, 1950), female. Original.

Acanthococcus costatus Danzig, 1975 (Fig. 41) Redescription

Acanthococcus costatus Danzig, 1975: 43.
Holotype: Female. Russia (Vladivostok, Okeanskaya), on Ulmus davidiana var. japonica,

01.vi.1963, by E. Danzig. By original designation. Deposited in ZMAS.
Lit.: Danzig, 1980: 205; 1988: 708; Hua, 2000: 137; Koteja & Zak-Ogaza, 1981: 513;
Hua, 2000; Kozár & Walter, 1985: 74; Tang & Hao, 1995: 450; Tao, 1999: 32; Wang,
1982b: 143; 2001: 208; WangZQ, 1982: 41; Xie, 1998: 95 (Miller et al., 2013).
Eriococcus ulmi; Tang, 1977: 45.

Syntypes: Female. Deposited in EISC. Synonymy by Danzig, 1983: 521.
Eriococcus costatus; Tang & Hao, 1995: 466. Change of combination.
Acanthococcus costatus; Köhler, 1998: 375. Revived combination.
Eriococcus costatus; Miller & Gimpel, 2000: 177. Revived combination.
Acanthococcus costatus; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Adult female oval, brown, 3 mm long. Ovisac white with obvious transverse ribs and a
depressed zone along the median body line.
Mounted female
Antennae 6 segmented, third segment longest. Large frontal lobe present. Anal lobes
strongly protruding, apically acute, heavily sclerotized, with numerous teeth on mesal
margin, dorsally with 3 enlarged setae, ventrally with 3 slender hair-like setae and with a
suranal setae.
Venter: Labium 3 segmented. Apical labial segment with 5 pairs of long setae. Legs small;
hind coxa and femur with spinulae on anterior surface, pores absent on coxa, each tibia
with 4 setae elongate, not enlarged; tarsus always longer than tibia, claws with denticle
near tip. Tarsal and claw digitules longer than claw, slightly knobbed. Hair-like setae
elongate; enlarged setae of small size only, present along body margin. Quinquelocular
pores numerous present over entire venter. Macrotubular ducts of two size; larger size in
high numbers along body margin; smaller size in reduced numbers in medial or sublateral
areas. Microtubular ducts absent. Cruciform pores present along lateral margin.
Dorsum: Enlarged setae normally of three sizes: 2 or 3 larger setae along margin of
abdominal segments and on margin of thorax and head, additional large setae often
present in two longitudinal bands median lines, enlarged setae absent on abdominal
segment VIII. Macrotubular ducts present over dorsum. Microtubular ducts numerous,
narrow, long. Anal ring dorsal, with 8 setae. Cauda present (Redescription after Danzig
(1975) and Tang & Hao (1995), with modifications based on the paratype).
Acanthococcidae 109
Figure 41. Acanthococcus costatus Danzig, 1975, female. After Danzig (1975) with
modifications.
Ecology
Host plant: Ulmus davidiana var. japonica, U. pumila.
Distribution: China, Russia.
Biology: Xie (1998) describes and illustrates a winged and a wingless male form of this
species. Eggs yellow, layed in June (Danzig, 1975).
Acanthococcus ericae (Signoret, 1875) (Fig. 42)

Eriococcus ericae Signoret, 1875: 31.
Type material: Unknown type status, type designation unknown. Notes: Matile-
Ferrero & Danzig visited the Naturhistorisches Museum, Vienna in June of 1997 and
were unable to locate any type material of this species.
Additional metarial examined: France, Post cros, female on Erica sp., A. S. Balachowsky,
19. vii. 1965 (MNHN: 3003-4).
Lit.: Balachowsky, 1933: 6; Brown, 1967: 131; Cockerell, 1896: 323; Fernald, 1903: 74;
Foldi, 2001: 305; 1937: 13; Gómez-Menor Ortega, 1957: 79; Goux, 1948: 67; Green,
1930: 11; Hulden, 1985: 59; Kosztarab & Kozár, 1978: 67; Kozár, 2009: 92; Leonardi,
1920: 434; Lindinger, 1957: 548; Martin-Mateo, 1985: 92; Miller & Gimpel, 1996: 600;
2000: 199; Ouvrard & Kozár, 2009: 102; Pellizzari & Kozár, 2011: 66; Tang & Hao,
1995: 450; Tranfaglia & Esposito, 1985: 126; Wünn, 1926: 49 (Miller et al., 2013).
Nidularia ericae; Lindinger, 1935: 134. Change of combination.

Eriococcus ericae; Hoy, 1963: 87. Revived combination.
Acanthococcus ericae; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus devoniensis; Lindinger, 1911: 357. Incorrect synonymy (Hoy, 1963).
Eriococcus thymi; Lindinger (1911 & 1912). Incorrect synonymy (Hoy, 1963).
Eriococcus ericae; Miller & Gimpel, 2000: 199. Revived combination.
Acanthococcus ericae; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Adult female oval, with evident segmentation
Mounted female
Body elongate oval, 2.17–2.8 mm long, 1.52–1.82 mm wide. Antennae 7 segmented,
206.5–266.0 µm long; segments V-VI with 1 sensory falcate seta. Frontal frontal lobe
well developed. Eyes situated on venter near margin. Anal lobes well developed, heavily
sclerotized but without medial teeth each with three enlarged setae on dorsal surface,
outer setae stronger (39.9–47.1 µm long) than those two on inner side (32.6–39.9µm
Acanthococcidae 111
Figure 42. Acanthococcus ericae (Signoret, 1875), female. After Tranfaglia & Esposito
long); enlarged setae more narrow than other dorsal setae; anal lobe seta 206–210 µm,
subapical setae 36.3–43.5 µm long.
Venter: Labium 3 segmented, basal segment with two setae present on each side. Legs
well developed; posterior tarsus 153.4–196 µm and tibia 137.9–159.7 µm long. Tarsal
and claw digitules knobbed, slightly longer than claw; claw with denticle at apex. Median
and posterior coxae with spinulae on anterior surface, posterior coxae with some pores.
Tibiae each with 4 setae. Derm with a few hair-like setae; macrotubular ducts of two
sizes, scattered. Multilocular pores each 4–6 µm in diameter and with 3-5 loculi, mostly
5, distributed in sparse rows on all abdominal segments, scattered on thorax and head.
Microtubular ducts not seen. Cruciform pores sparse on submargin.
Dorsum: Marginal dorsal enlarged setae narrowly conical spine-like, with slightly rounded
apices, setae of various sizes, scattered over dorsal surface; in rows across all segments;
setae on the middle part of thorax larger than others dorsal setae. Macrotubular ducts
sitiuated among enlarged setae in bands and rows. Microtubular ducts scattered. Anal
ring strongly sclerotized, with partly double rows of pores; slightly oval, with 4 pairs of
long setae, 98–115 µm long; Cauda not seen (after Tranfaglia & Esposito (1985) with
modifications).
Other stages
First instar described by Leonardi (1920), which shows some similarity with A. aceris, but
differs by the absence of shorter spines on dorsum.
Ecology
Host plant: Calluna vulgaris, Erica arborea, E. carnea, E. multiflora, E. tetralix.
Distribution: Algeria, Austria, France, Germany, Italy, Malta, Netherlands, Spain. The
distribution record for this species given by Tranfaglia & Esposito (1985) as "Gran
Bretagna" and the source is given as Hoy (1963). However, Great Britain is not listed in
Hoy (1963) or Williams (1985a) which treats the eriococcid fauna of Britain.
Biology: Unknown.
Comments
The drawing of Tranfaglia & Esposito (1985) was compared with slides deposited in
MNHN No 4943 (Boci ds Bordes, 18. x. 1908, Calluna vulgaris, St. Ent. De Paris, coll.
P. Marchal) and No 3003-4 (Port Cros, A.S.Balachowsky, 19.vii.65, Erica arborea). The
morphology of these specimens quite well agree with the drawing of Tranfaglia &
Esposito (1985). Here we provide a new drawing based on material from MNHN (3003-
4).
Acanthococcidae 113
Acanthococcus glanduliferus (Balachowsky, 1933) (Fig. 43) Redescription

Eriococcus glanduliferus Balachowsky, 1933:36.
Syntype: Female. France (Salins de Badon), on Salicornia fruticosa, 06.viii.1930, by A.

S. Balachowsky. Deposited in MNHN (type no. 4941). Notes: There are six slides
containing nine specimens deposited in the MNHN.
Lit.: Foldi, 2001: 305; 2002: 245; Goux, 1948: 67; 1989: 21; Kozár & Walter, 1985: 74;
Köhler, 1998: 376; Ouvrard & Kozár, 2009: 102 (Miller et al., 2013).
Nidularia glanduliferus; Lindinger, 1936:156. Change of combination.

Eriococcus glanduliferus; Hoy, 1963: 91. Revived combination.
Acanthococcus glanduliferus; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus glanduliferus; Miller & Gimpel, 2000: 214. Revived combination.
Acanthococcus glanduliferus; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Adult female is globular and elongate. Ovisac is globular, white or gray 2–3 mm long,
1.5–2.0 mm wide.
Mounted female
Adult female oval, 2–2.2 mm long, 1.2–1.3 mm wide. Antennae 7 segmented, length of
segments: I: 37 µm, II: 21 µm, III: 36.3 µm, IV: 24.3 µm, V: 24.5 µm, VI: 14.3 µm and 27.3
µm long; third and fifth segments without hair-like setae, other segments with a few hair-
like setae; apical segment with apical seta 32 µm long; apical segment also with 3 sensory
falcate setae, 21.3 µm long; two preapical segments with 1 sensory falcate seta: on fifth
segment 16.5, sixth 21.5 µm long. Frontal lobe absent, frontal tubercle 4 µm diameter.
Eyes situated on venter near margin. Anal lobes well developed, sclerotized, with 3 blunt
conical spines, ventrally with two flagellate setae; anal lobe with long apical seta.
Venter: Labium 3 segmented, basal segment with 2 pairs of setae, apical segment with
5 pairs of hair-like setae. Legs well developed: lengths of segments and digitules of
prothoracic legs; coxa: 48.3 µm, trochanter: 41.3 µm, femur: 112 µm, tibia: 86 µm; tarsus:
98.6 µm and claw: 33.5 µm long, tarsal digitules: 41.3 µm, claw digitules: 33 µm long;
lengths of segments and digitules of mesothoracic legs legs; coxa: 53.3 µm, trochanter:
47 µm, femur: 100.3 µm, tibia: 82.6 µm, tarsus: 96.3 µm, claw: 34 µm long, tarsal digitules:
44.3 µm, claw digitules: 32 µm long; lengths of segments and digitules of metathoracic
legs; coxa: 65.3 µm, trochanter: 48 µm, femur: 109 µm, tibia: 87.3 µm, tarsus: 104 µm;
claw: 32.6 µm long, tarsal digitules: 40.6 µm, claw digitules: 32.5 µm long, tarsal and
claw digitules slightly knobbed, longer than claw, meso- and metathoracic coxae with
spinulae on anterior surface; metathoracic coxae and femur also with small pores on both
anterior and posterior surfaces; claw with denticle; legs with a few hair-like setae; tibia
with 4 setae. Width of anterior spiracles 23 m. Multilocular pores 6–7 µm in diameter

and with 5 or 7 loculi, mostly 5; distributed in transverse bands across abdominal
sternites, in sparse rows on thoracic segments and head region. Macrotubular ducts
of two sizes: larger macrotubular ducts 7–8 µm wide and 21–24 µm long, situated on
margin; smaller macrotubular ducts 4 µm wide and 26–30 µm long scattered throughout
the venter. Microtubular ducts sparse marginally and submarginally. Cruciform pores a
few in submarginal area on each thoracic segments, submedially only on mesothoracic
segment. A few hair-like setae present on submedian venter, a few knobbed hair-like setae
submarginally on abdominal segments. Blunt conical enlarged setae in two sizes; larger 14
µm long on margin; smaller ones half-sized, submarginally. Suranal setae hair-like.
Dorsum: Enlarged setae cylindrical, apices rounded or truncate, marginal setae noticeably
larger than other dorsal setae, forming marginal line around body, lateral area of abdominal
segments with 1 or 2 setae, longer setae, 26–30 µm long and medium sized setae, 19–21
µm long, shorter setae more numerous, about 13–15 µm long, forming sparse bands
on thoracic and abdominal segments, and medial and submedial groups on the head
region, but absent on abdominal segment VIII. Ten dorsal setae on abdominal segment
VII. Macrotubular ducts 7–8 µm wide and 21–24 µm long, scattered among enlarged
setae. Microtubular ducts elongate, scattered, 8 µm long. Anal ring situated on margin
of dorsum; oval, sclerotized, 38–42 µm wide, 50–55 µm long, with one row of pores
and with 8 setae, each 89–107 µm long. Cauda hardly visible, divided (after Balachowsky,
1933 with a redescription and redrawing, measurements are based on holotype and 2
paratypes).
Ecology
Host plant: Salicornia fruticosa, Arthrocnemum macrostachyum.
Distribution: France.
Biology: Unknown.
Acanthococcidae 115
Figure 43. Acanthococcus glanduliferus (Balachowsky, 1933), female. Original.

Acanthococcus isacanthus Danzig, 1975 (Fig. 44) Redescription

Acanthococcus isacanthus Danzig, 1975: 45.
Holotype: Female. Russia (Vladivostok), on Spiraea salicifolia, 16.vi.1963, by E.

Danzig. By original designation. Deposited in ZMAS.
Lit.: Danzig, 1980: 205; 1986: 240; 1988: 708; Hua, 2000: 137; Koteja & Zak-Ogaza,
1981: 513; Kozár & Walter, 1985: 74; Kwon & Han, 2003: 151; Ouvrard & Kozár,
2009: 102; Tang & Hao, 1995: 451; Tao, 1999: 32; Wang, 2001: 208 (Miller et al., 2013).
Eriococcus isacanthus; Tang & Hao, 1995: 474. Change of combination.

Acanthococcus isacanthus; Köhler, 1998: 378. Revived combination.
Eriococcus isacanthus; Miller & Gimpel, 2000: 242. Revived combination.
Acanthococcus isacanthus; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Female is oval, brown. 2 mm long. Ovisac is white, with slightly apparent transverse ribs
and waxy spines on the margin.
Mounted female
Adult female oval. Antennae 7 segmented. Frontal lobe present. Anal lobes strongly
protruding, apically acute, heavily sclerotized, with numerous teeth on mesal margin,
dorsally with 3 enlarged setae, ventrally with 3 slender hair-like setae and with a suranal
setae.
Venter: Apical labial segment with 6 (?) pairs of long setae. Legs small; hind coxae and
femur with spinulae on anterior surface, coxae without pores, tibia with 4 setae elongate,
not enlarged; tarsi longer than tarsus, claws with denticle near tip. Tarsal and claw digitules
longer than claw, slightly knobbed. Quinquelocular pores numerous present over entire
venter. Macrotubular ducts of 2 sizes: larger size in high numbers along body margin;
smaller size in reduced numbers in medial or sublateral areas. Microtubular ducts absent.
Hair-like setae elongate; enlarged setae of small size only, present along body margin.
Cruciform pores present along lateral margin.
Dorsum: Enlarged setae normally of three sizes: 2 or 3 larger setae along margin of
abdominal segments and on margin of thorax and head, on abdominal dorsum spines
form 1 or 2 irregular rows of enlarged setae, enlarged setae absent on abdominal segment
VIII. Macrotubular ducts present over dorsum. Microtubular ducts numerous, narrow,
long. Anal ring dorsal, with 6 setae. Cauda present (Redescription after Danzig (1975) and
Tang & Hao (1995), with modifications based on paratypes).
Acanthococcidae 117
Figure 44. Acanthococcus isacanthus Danzig, 1975, female. After Danzig (1975) with
modifications.
Ecology
Host plant: Albizia kalkora, Spiraea salicifolia.
Distribution: China, North Korea, Russia.
Biology: This species appears to be monophagus on the mesophylic Spiraea salicifolia.
Female produce eggsacs in mid June, on the twig ramifications, at the base of the bud.
Acanthococcus kaschgariae Danzig, 1972 (Fig. 45)

Acanthococcus kaschgariae Danzig, 1972b: 339.
Holotype: Female. Mongolia (Bayan-Hongor Aymag, Edrengiyn-Nuru Ridge, 100

km SSW Bayan-Under), on Kaschgaria komorovi roots, 05.ix.1970, by I. Kerzhner. By
original designation (type no. 200-75). Deposited in ZMAS. Notes: 1 female on same
slide as holotype. 4 paratypes on 2 slides with same data in ZMAS.
Lit.: Köhler, 1998: 379; Kozár & Walter, 1985: 74; Matesova, 1976: 26; Miller &
Gimpel, 2000: 244; Tang & Hao, 1995: 451; Ter-Grigorian, 1983: 881.
Eriococcus kaschgariae; Tang & Hao, 1995: 474. Change of combination.

Acanthococcus kaschgariae; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Unknown.
Mounted female
Antennae 7 segmented. Frontal lobe not seen. Anal lobes protruding, apically acute,
dorsally with 3 enlarged setae.
Venter: Apical labial segment with 6 pairs of long setae, the median setae much shorter
than others. Legs medium sized; hind coxae and femora with spinulae on anterior surface
and pores on coxae, tibia with 4 strong setae; tarsi longer than tibiae, claws with denticle
near tip; tarsal digitules longer than claw, slightly blunted, claw digitules spinelike, much
shorter than claw, not blunted. Quinquelocular and some trilocular pores numerous,
present over entire venter. Macrotubular ducts scattered all over the venter. Microtubular
ducts absent. Hair-like setae short; enlarged setae of small size only, present along body
margin. Cruciform pores present along lateral margin.
Dorsum: Enlarged setae, conical, almost about the same sizes, without marginal row
of enlarged setae; on dorsal segments enlarged setae form in wide bands including
big groups in middorsum of abdomen and thorax. Number of setae smaller on head.
Macrotubular ducts present in small number all over dorsum. Microtubular ducts narrow,
long, numerous. Anal ring dorsal, with 8 setae with one row of pores. Cauda present
(after Danzig (1972b), with some modifications based on paratypes).
Acanthococcidae 119
Figure 45. Acanthococcus kaschgariae Danzig, 1972, female. After Danzig (1972b) with
modifications.
Ecology
Host plant: Kaschgaria komarovi.
Distribution: Mongolia.
Biology: Lives on the roots of host plant.
Acanthococcus kilinceri Kaydan Species nova (Fig. 46)
Holotype: Female. Turkey (Hakkari road), on Quercus sp., N: 37°41’644’’; E:

043°56’266’’; F. Kozár and M.B. Kaydan, 23. v. 2008 (KPCT: 4222). By original
designation. 7 paratypes of the same data as holotype, and 6 paratypes deposited in
Kaydan’s personal collection (KPCT) and one paratype in the PPI.
Description
Unmounted female
Adult females reddish; felt-like test cream-white.
Mounted female
Body elongate oval, 1.68–2.96 mm long, 1.44–1.58 mm wide. Antennae 7 (sometimes
8) segmented, 310–400 µm, length of segments: I: 40–70 µm, II: 57.5–75.0 µm, III:
57.5–70.0 µm, IV: 65–95 µm, V: 25–40 µm, VI: 27.5–30.0 µm, and VII: 40–45 µm long,
segments covered with a few, strong hair-like setae (third segment with 1–3 setae); apical
segment with apical seta 67.5–70 µm long; apical segment also with 3 sensory falcate
setae, 27.5–40.0 µm long; segment VI with 1 sensory falcate seta 20.0–27.5 (20) µm long,
segment V with 1 sensory falcate seta 27.5 µm long. Frontal lobes and frontal tubercle
present. Eyes situated on venter near margin. Anal lobes strongly developed, sclerotised,
with 3 enlarged setae, 40–60 µm plus 3-5 microtubular ducts on dorsal surface; apical seta
285–325 µm; ventral hair-like subapical seta 135–170 µm long.
Venter: Labium 250–270 µm long, 135–165 µm wide, median setae on apex of labium
25–35 µm long. Stylet loop long, reach to last coxae. Legs well developed; lengths of
segments and digitules of prothoracic legs; coxa: 85–115 µm, trochanter: 60.0–85.0 µm,
femur: 140–170 µm, tibia: 120–135 µm, tarsus: 137.5–160.0 µm and claw: 30–35 µm,
trochanther + femur: 200–240 µm, tibia + tarsus: 280–290 µm, tarsal digitules 52.5–55.0
µm, claw digitules 32.5–35 µm long; lengths of segments and digitules of mesothoracic
legs; coxa: 100.0–112.5 µm, trochanter: 60–85 µm, femur: 160–170 µm, tibia: 120.0–
137.5 µm, tarsus: 145–170 µm and claw: 32.5–37.5 µm, trochanther + femur: 200–245
µm, tibia + tarsus: 270–310 µm, tarsal digitules 55 µm, claw digitules 35–40 µm long;
lengths of segments and digitules of metathoracic legs; coxa: 90–110 µm, trochanter: 75–
80 µm, femur: 145–170 µm, tibia: 140–160, tarsus: 155–175 µm and claw: 32.5–37.5 µm,
trochanther + femur: 215–250 µm, tibia + tarsus: 310–330 µm, tarsal digitules 52.5 µm,
claw digitules 35.0–37.5 µm long; meso- and metathoracic coxae with spinulae on ventral
Acanthococcidae 121
Figure 46. Acanthococcus kilinceri Kaydan, sp. n., female, with first instar on top right.
surface; tibiae of meso and metathoracic legs with 4 setae (median seta absent), tibiae of
prothoracic legs with 5 setae (median seta present), tarsi with 6 setae. Length of spiracles
65–85 µm; diameter of spiracular peritreme 37.5–50.0 µm, posterior spiracles slightly
larger than anterior. Setae in median areas flagellate, 20–135 µm long, setae in lateral
areas flagellate but stronger than setae on median area, 50–70 µm long. Multilocular
pores 3–5 µm in diameter and with 5 loculi, distributed in sparse bands on all abdominal
segments and scattered on thorax and head. Macrotubular ducts of three sizes, smallest
macrotubular ducts 2.5–3.0 µm wide and 15–17 µm long, present on median areas of
abdominal segments, thorax and around labium, medium macrotubular ducts 7–10 µm
wide and 20–25 µm long, present on submargin of body, largest macrotubular ducts
7.5–10.0 µm wide and 22.5–27.5 µm long, on submarginal band on abdomen, thorax
and head. Cruciform pores on on submedian area in great groups and band on meso and
metathorax and abdominal segments I–IV, 2.5–3.0 µm in diameter.
Dorsum: Marginal dorsal setae long blunted 40–65 µm on the (abdominal segment VII
with two long two short setae) forming a marginal row and differentiated from dorsal
setae, other dorsal setae conical, truncated, 20.0–37.5 µm long; arranged in transverse
rows across body segments, rows irregular on head, on the abdominal segment VIII
6–8 enlarged setae situated. Macrotubular ducts, 7.5–10.0 µm wide and 20.0–27.5 µm
long, scattered throughout dorsum, generally in segmental bands. Microtubular ducts
long, 7–11 µm long with oval dermal orifice, scattered over dorsum. Anal ring strongly
sclerotized, with 18-22 pores on each side, 65–100 µm in diameter, with 8 setae, each
125–155 µm long; anal ring situated on margin of dorsum. Cauda present.
Other stages
Mounted first instar nymph (Fig. 46, top right).

Body of slide-mounted specimens, oval. Antennae 6 segmented, apical three segments
strong spines with pointed apices of one sizes, forming middorsal longitudinal pair of
rows on head and thorax, the submedian row contains only three large spines, what is
continued by 5–8 very small setae, and one pair of large spines in front of the anal ring.
Microtubular ducts long, few in two middorsal longitudinal rows on both side. Venter
with transverse rows of six small hair-like setae on each abdominal segment; median setae
longer than others. Cruciform pores present on thoracic and first abdominal segments,
with 1 pair of quinquelocular pores on each thoracic segment, 1 near each spiracle, plus 1
pair on frons, and two longitudinal submedian rows on abdomen. Anal ring with 6 spine-
like setae. First instar nymph similar to nymph of A. roboris.
Acanthococcidae 123
Etymology
The new species is named in honour of Prof. Dr. Neşet Kılınçer who made valuable
entomological studies in Turkey.
Ecology
Host plant: Quercus sp.
Distribution: Turkey.
Biology: This species was found on the root crown or stem near to the soil or on the
small branch which was lying on the ground.
Acanthococcus koelreuterius (Wei & Wu, 2004) (Fig. 47)

Eriococcus koelreuterius Wei & Wu, 2004: 118.
Holotype: Female. China (Beijing). By original designation. Deposited in the Insect

Collection of Beijing Forestry University.
Lit.: Kozár, 2009: 92 (Miller et al. 2013).
Acanthococcus koelreuterius; Kozár, 2009: 92. Change of combination.
Description
Unmounted female
Adult female: the body spindle-shaped, 1.88–2.2 mm long, 1.0–1.15 mm wide.
Mounted female
Antennae 7 or 8 (usually 7) segmented, length of segments: I: 50 µm, II: 45 µm, III: 55
µm, IV: 65 µm, V: 30 µm, VI: 25 µm, and VII: 45 µm long. Frontal tubercles developed.
Eyes present. Anal lobes large and sclerotized, with inner margins serrate and 3 equal
long spines on its dorsal surface. Anal lobe setae 280 µm long.
Venter: Mouth part well developed. Legs medium size, tibia shorter than tarsus, tarsal
and claw digitules longer than claw, with expanded tip, claw with denticle, hind coxae
without translucent pores. Quinquelocular pores with few septalocular or quadrilocular
pores present all ventral surface. Cruciform pores, present in margins of head, thorax and
abdominal segment. Enlarged setae of two sizes, the large one 37.5 µm long, the small
one 25 µm long, present along the margin; hair-like setae about in four sizes, distributed
in middle part of venter. Macrotubular ducts of two sizes, the largest one 17.5 µm long
and 7.5 µm wide, present along the margin; the small one 17.5 µm long and 5.0 µm wide,
distributed in venter of thorax and abdomen, more in middle area of abdomen.
Dorsum: Dorsum with blunted enlarged setae of three sizes, the largest one about 75 µm
long, 15 µm in diameter at its base, present along margin and midline, the median one
about 50 µm long and 9 µm in diameter, the small one about 40 µm long and 7.5 µm in
diameter, the large and medium size ones distributed in marginal and medial longitudinal
rows, medial row with 1 or 2 and marginal row with 2 or 3 spines on body segments.
There are two enlarged setae on segment VIII and a transverse band of enlarged setae
on abdominal segment VII. Macrotubular ducts of one size, scattered through body,
202-282 in total number. Microtubular ducts narrow, long, scattered, 194-228 in total.
Anal ring with two rows pores and 8 setae. Cauda not mentioned (after Wei & Wu (2004),
with modifications).
Other stages
First-instar nymph typical for the genus, morphologicaly near to A. aceris first-instar
nymph.
Acanthococcidae 125
Figure 47. Acanthococcus koelreuterius (Wei & Wu, 2004), female. After Wei & Wu (2004)
with modifications.
Ecology
Host plant: Koelreuteria paniculata.
Biology: Unknown.
Acanthococcus lagerstroemiae (Kuwana, 1907) (Fig. 48)

Eriococcus lagerstroemiae Kuwana, 1907: 182.
Syntypes: Female. Japan (Ichijiku and Sarusuberi), on Ficus carica and Lagerstroemia
indica. Deposited in Tachikawa.
Common name: Crapemyrtle scale.
Lit.: Ali, 1970: 76; Bielenin & Weglarska, 1992: 422; Foldi, 1983: 164; Green, 1915:
177; Kawai, 1972: 4; 1977: 152; 1980: 127; Kwon & Han, 2003: 151; Köhler, 1998:
379; Paik, 1978: 165; Takahashi, 1957: 7; Tang, 1977: 43, 1984: 125; Tang & Hao,
1995: 449; Tao, 1999: 32; Wang, 1974: 329; 2001: 208; Williams, 1985a: 374; Yang,
1982: 105 (Miller et al. 2013).
Nidularia lagerstroemiae; Lindinger, 1933a: 116. Change of combination.

Acanthococcus lagerstroemiae; Borchsenius, 1960b: 214, 217. Change of combination.
Eriococcus lagerstroemiae; Hoy, 1963: 99. Revived combination.
Acanthococcus lagerstroemiae; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus lagerstroemiae; Miller & Gimpel, 2000: 251. Revived combination.
Acanthococcus lagerstroemiae; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Egg sac is snow white in color, about 6 mm long, 3 mm wide. Body is oval or broad
elliptical, about 4 mm long, dark purple in color. Dorsum has very spiny hairs. Eggs are
purple-red and turn pinkish-red with age (Miller et al., 2013).
Mounted female
Antennae 7 segmented, second and third segments subequal, and the longest is the third,
segments bears a few rather long strong setae. Frontal lobes developed but smaller than
basal antennal segment. Anal lobes protruding, apically rounded, moderately sclerotized,
serrate on inner margin, with 3 dorsal equal long conical setae, ventrally with two flagellate
setae, anal lobe with long apical setae.
Venter: Labium 3 segmented, basal segment with 2 pairs of setae; stylet loop relatively
short, somewhat longer than labium; with normal flagellate setae in median areas, short
and slender but stouter and stiff in lateral areas. Legs subequal, comparatively small,
tarsus longer than tibia, claw with denticle, tarsal and claw digitules longer than claw, with
Acanthococcidae 127
Figure 48. Acanthococcus lagerstroemiae (Kuwana, 1907), female. After Tang & Hao
capitated apices; coxae without translucent pores but with spinulaes on anterior side.
Quinqelocular pores in transverse bands across abdominal sternites, around margins to
head, and on mid venter. Macrotubular ducts of two sizes: not numerous, in more or less
single rows on abdominal segments and around submargins to head. Microtubular ducts
absent. Cruciform pores present in submarginal area from head to about abdominal
segment IV.
Dorsum: Enlarged setae spine-like setae of 2–3 sizes; enlarged setae cylindrical, apices
rounded, abundant over dorsal surface; largest setae form sparse, transverse segmental
bands; on abdominal segment VII number of spines 14–18, on segment VIII spines
absent, on the abdominal margin 3 or 4 setae present. Macrotubular ducts of two sizes,
large duct numerous among enlarged setae. Microtubular ducts narrow, long, elongate,
with one sclerotized area, orifice bifurcate; evenly distributed. Anal ring oval, sclerotized,
with two rows of pores and with 8 setae. Cauda present (after Kawai, (1980), Kuwana,
(1907), Tang & Hao (1995) with modifications).
Ecology
Host plant: Anogeiussus sp., A. latifolia, Buxus microphylla koreana, Celtis sinensis, Dalbergia
sp., Diospyros kaki, Ficus carica, Glochidion puberum, Glycine max, Lagerstroemia speciosa, L.
indica, L. japonica, Ligustrum obtusifolium, Mallotus japonicus, Malus pumila, Myrtus sp., Punica
granatum, Rubus sp.
Distribution: China, India, Japan, Mongolia, South Korea, United Kingdom (England).
Biology: This species has two generations in a year. The first adults appear in late April
or early May, the second from late August to late October, 3 generations per year in China
and that the overwintering stage is nymphal. It can be a pest on Myrtus spp., Punica spp.
and Rubus spp. Tang & Hao (1995).
Acanthococcidae 129
Acanthococcus latialis (Leonardi, 1907) (Fig. 49)

Eriococcus latialis Leonardi, 1907: 144.
Lectotype: Female. Italy (Marino, near Rome), on undetermined host, by F. Silvestri.

By subsequent designation Tranfaglia & Esposito, 1985: 130. Deposited in IFSP.
Lit.: Goux, 1938: 333; Hoy, 1963: 99; Kozár & Walter, 1985: 74; Köhler, 1998: 380;
Leonardi, 1918: 215; 1920: 436; Lindinger, 1912: 349; Longo et al., 1999: 121; Pellizzari
& Kozár, 2011: 66; Tang & Hao, 1995: 450 (Miller et al., 2013).
Nidularia latialis; Lindinger, 1933a: 116. Change of combination.

Acanthococcus latialis; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus latialis; Miller & Gimpel, 2000: 255. Revived combination.
Acanthococcus latialis; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Unknown.
Mounted female
Adult female elongate elliptical, 3.22–3.36 mm long, 1.96–2.04 mm wide. Antennae 7
segmented, 252–280 µm long long second and third segments subequal, fourth is the
longest, each segment bears a few strong setae. Frontal lobes not mentioned. Anal lobes
protruding, heavily sclerotized, apically rounded, serrate on inner margin, with 3 dorsal
equal long enlarged setae, ventrally with two flagellate setae, and one suranal setae, anal
lobe, with long apical setae.
Venter: Labium 3 segmented, basal segment with 2 pairs of setae. Stylet loop relatively
short, somewhat longer than labium. With normal flagellate setae in median areas, short
and slender but stouter and stiff in lateral areas. Legs well developed, claw and tarsus
185.1–203.2 µm long, tibia 123.4–137.9 µm long, tarsus longer than tibia, ratio of tibia/
tarsus 0.6, claw with denticle, tarsal and claw digitules knobbed, longer than claw; coxa
without translucent pores but with spinulaes on anterior side. Quinqelocular and some
trilocular pores in transverse bands across abdominal sternites, around margins to head,
and on midventer. Macrotubular ducts of one size present only around submargin.
Microtubular ducts absent. Cruciform pores not mentioned.
Dorsum: Enlarged setae cylindrical, apices almost truncated, 39.9–54.4 µm long marginal
setae on abdomen slightly larger than others, abundant over dorsal surface; spine-like
setae of 2 or 3 sizes; largest setae form sparse, transverse segmental rows and bands;
on abdominal segment VII number of spines 22, on segment VIII enlarged setae not
mentioned, on the abdominal margin 3-4 setae present. Macrotubular ducts of one size,
large ducts scattered among enlarged setae. Microtubular ducts narrow, long, elongate,
with 1 sclerotized area, evenly distributed. Anal ring oval, sclerotized, with two rows of
pores and with 8 setae. Cauda not mentioned (Tranfaglia & Esposito, 1985).
Other stages
First instar described by Leonardi (1920), very similar to that of A. aceris.
Ecology
Host plant: Undetermined.
Distribution: Italy.
Biology: Unknown.
Acanthococcus macedoniensis Fetykó & Kaydan, Species nova (Fig. 50)
Holotype: Female. Macedonia (Dorjan), on branches of Quercus ilex, F. Kozár and K.

Fetykó, 10. iv. 2010 (PPI: 9291). Deposited in PPI. Paratypes: 3 adult females, same data
as holotype (marked) (PPI: 9291).
Description
Unmounted female
Adult female reddish; felt-like test cream-white.
Mounted female
220–260 µm, length of segments: I: 55–60 µm, II: 32.5–35.0 µm, III: 42.5–45.0 µm, IV:
35–45 µm, V: 20–25 µm, VI: 20–25 µm, and VII: 35–40 µm long, the segments covered
with a few, strong hair-like setae (except third segment); apical segment with apical seta
50–55 µm long; apical segment also with 3 sensory falcate setae, 22.5–32.5 µm long;
segment VI with 1 sensory falcate seta 25 µm long, segment V with 1 sensory falcate
seta 25 µm long. Frontal lobes and frontal tubercle present. Eyes situated on venter near
margin. Anal lobes strongly developed, with 3 enlarged setae, 40–70 µm long, plus 4-6
microtubular ducts on dorsal surface. Apical seta 225–245 µm; ventral hair-like subapical
seta 95–135 µm long.
long, 25.0–32.5 µm long, stylet loop long, longer than body length. Legs well developed;
lengths of segments and digitules of prothoracic legs; coxa: 60 µm, trochanter: 45–60 µm,
femur: 115 µm, tibia: 160–170 µm, tarsus: 85.0–92.5 µm and claw: 30 µm, trochanther +
femur: 160–170 µm, tibia + tarsus: 195 µm, tarsal digitules 45 µm, claw digitules 32.5 µm
long; lengths of segments and digitules of mesothoracic legs; coxa: 60–65 µm, trochanter:
Acanthococcidae 131
Figure 49. Acanthococcus latialis (Leonardi, 1907), female. After Tranfaglia & Esposito
47.5–60.0 µm, femur: 105–120 µm, tibia: 95–115 µm, tarsus: 100–115 µm and claw: 30
µm, trochanther + femur: 155–170 µm, tibia + tarsus: 210–220 µm, tarsal digitules 45 µm,
claw digitules 30 µm long; lengths of segments and digitules of metathoracic legs; coxa:
67.5–70.0, trochanter: 45–55 µm, femur: 100–120 µm, tibia: 95–105 µm, tarsus: 120–130
µm and claw: 31–32.5 µm, trochanther + femur: 167.5–170 µm, tibia + tarsus: 215–235
µm, tarsal digitules 50.0–52.5 µm, claw digitules 30 µm long; tarsal and claw digitules
knobbed; meso- and metathoracic coxae with spinulae on ventral surface; tibiae of meso
and metathoracic legs with 4 setae (median seta absent), tibiae of prothoracic legs with 5
setae (median seta present), tarsi with 6 setae. Length of spiracles 50–70 µm; diameter of
spiracular peritreme 27.5–35.0 µm, posterior spiracles slightly larger than anterior. Setae
in median areas flagellate, long, 25–75 µm long, setae in lateral areas spine like, 15.0–17.5
µm long. Enlarged setae, situated on submargin in a row, 15.0–17.5 µm long, on head like
dorsal setae 25.0–27.5 µm long. Multilocular pores 3–5 µm in diameter and with 5 loculi,
distributed in sparse bands on all abdominal segments and scattered on thorax and head.
Macrotubular ducts of two sizes, smaller macrotubular ducts 3–4 µm wide and 15.0–17.5
µm long, present on median areas of abdominal segments, larger macrotubular ducts
5.0–7.0 µm wide and 17.5.0–22.5 µm long, sparse in a band on last abdominal segments,
and in a submarginal band on abdomen, thorax and head. Cruciform pores on thorax
generally on submedian area in a band on thorax and first for abdominal segments and a
few on head, 3.0–4.0 µm in diameter.
Dorsum: Marginal dorsal enlarged setae long, blunted, 35–60 µm forming a marginal row
and differentiated from dorsal setae, other dorsal setae conical, blunted, seta 25.0–27.5
µm long; arranged in transverse rows across body segment, 8 enlarged setae in midle of
segment VIII, rows irregular on head. Macrotubular ducts 7.5–10.0 µm wide and 20.0–
27.5 µm long, scattered throughout dorsum, generally in segmental bands. Microtubular
ducts long, 10.0–12.5 µm long with oval dermal orifice, scattered over dorsum. Anal ring
strongly sclerotized, with 19-21 pores on each side, 62.5–70.0 µm in diameter, with 8
setae, 115–145 µm long; anal ring situated on margin of dorsum. Cauda present.
Etymology
The new species is named after the country of collection.
Ecology
Host plant: Acer sp., Carpinus betulus, Quercus ilex.
Distribution: Macedonia, Poland, Switzerland.
Biology: On branches.
Acanthococcidae 133
Figure 50. Acanthococcus macedoniensis Fetykó & Kaydan sp. n., female.
Comments
The adult female of A. macedoniensis is a remarkable species having high number of
enlarged setae on last abdominal segments of dorsum, and by two sizes of macrotubular
ducts on venter.
There are two females in Kozár’s collection from Poland (Warsaw, 06.03. 1981, from
Acer sp. Leg: Kozár and Nagy (PPI 1443), female, and from Switzerland (Glarus, 01.05.
1992, Carpinus betulus, Leg: Kozár (PPI 3890), female, male, which belongs to this species,
and present new distribution records for these countries.
Acanthococcus melnikensis (Hodgson & Trencheva, 2008) (Figs. 51, 52,

53, 54) Reestablishment
Eriococcus melnikensis Hodgson & Trencheva, 2008: 12.
Holotype: Female. Bulgaria (Pirin Mountains, Melnik), on Quercus pubescens, 04.x.2008,

by K. Trencheva. Deposited in BMNH.
Lit.: Gavrilov, 2010: 38; Kozár, 2009: 93; Kozár et al., 2013: 55 (Miller et al., 2013).
Acanthococcus melnikensis; Kozár, 2009: 93. Change of combination.

Acanthococcus melnikensis; Gavrilov, 2010: 38. Synonymy with A. aceris. Here A. melnikensis
is reestablished. See notes under A. aceris.
Description
Unmounted female
Young female 3.25–5.0 mm long; venter wider than dorsum; dorsum 2.1–3.5 mm widest,
total width 4.12 mm widest. More or less oval but slightly more pointed posteriorly.
Ovisac felted, cream-white, up to 9 mm long, 6 mm wide. Adult female oval, red raspberry
colored.
Mounted female
250–310 µm long the segments covered with a few, long hair-like setae (third segment
without setae); apical segment with apical seta 40–60 µm long; apical segment also with
3 sensory falcate setae, segment VI with 1 and segment V with 1 sensory falcate seta.
Frontal lobes and frontal tubercle present. Eyes situated on venter near margin. Anal
lobes strongly developed sclerotised, relatively large, with 3 enlarged setae, plus several
microtubular ducts on dorsal surface; apical seta 70–115 µm; ventral hair-like subapical
seta 30–40 µm, suranal setae 85–112 µm long.
Venter: Labium 145–170 µm long, apical segment with 5 pairs of setae; stylet loop
relatively short, reaching posteriorly to median coxae. Legs well developed; lengths of
segments and digitules of metathoracic legs; coxa: 100–135 µm, trochanter+femur:
Acanthococcidae 135
Figure 51. Acanthococcus melnikensis (Hodgson & Trencheva, 2008), female, with first
instar on top right. After Hodgson & Trencheva (2008).
153–195 µm, tibia: 100–125 µm, tarsus: 125–150 µm and claw: 30–35 µm long, tarsal
digitules and claw digitules longer than claw, capitated; meso- and metathoracic coxae
with spinulae on ventral surface, tibiae of meso and metathoracic legs with 4 setae
(median seta absent), tibiae of prothoracic legs with 5 setae (median seta present), tarsi
with 6 setae. Spiracles quite small; diameter of spiracular peritreme 26–33 µm, posterior
spiracles slightly larger than anterior. Setae in median areas flagellate, 16–115 µm long,
enlarged setae in lateral areas spine-like, 25–30 µm long. Multilocular pores about 5 µm
in diameter and with 5 loculi distributed in sparse bands on all abdominal segments and
submargin and scattered on thorax and head. Macrotubular ducts of two sizes, smallest
macrotubular ducts 9–13 µm long, 0-10 present on median areas of abdominal segments,
larger macrotubular ducts forming a submarginal band on abdomen, thorax and head.
Cruciform pores generally few on thorax and abdominal segments and a few on head,
3.5–4.0 µm long and 2 µm wide.
Dorsum: Marginal enlarged setae 30–50 µm long, truncated, forming a marginal row and,
other dorsal enlarged setae conical, truncated, seta 35–50 µm long; arranged in transverse
rows across body segments, rows irregular on head, with 4-5 enlarged setae on the last
abdominal segment. Macrotubular ducts, 7–8 µm wide and 18–28 µm long, scattered
throughout dorsum, generally in segmental bands. Microtubular ducts long, 7–10 µm
long with oval dermal orifice, scattered over dorsum. Anal ring strongly sclerotized, with
a row of pores on each side, with 8 setae, 85–155 µm long; anal ring situated on margin of
dorsum. Cauda present (after Hodgson & Trencheva (2008), with some modifications).
Other stages
Mounted first instar nymph (Fig. 51, top right)

with strong sensory setae as in adult female. Frontal tubercle present. Dorsum with strong
spines with pointed apices of one size, forming middorsal longitudinal pairs of double
rows on head and thorax, the submedian row contains five large spines, followed by 4-6
very small setae, and two pairs of large spines in front of the anal ring. Microtubular
ducts long, few in three middorsal longitudinal rows on both side. Venter with transverse
rows of six small hair-like setae on each abdominal segment; median setae longer than
others. Cruciform pores present on thoracic segments, with 1 pair of quinquelocular
pores on each thoracic segment, 1 near each spiracle, plus 1 pair on frons, and two
longitudinal submedian rows on abdomen. Anal ring with 6 spine-like setae. First-instar
nymph similar to that of A. roboris.
Excellent drawings of adult female, male, second instar female and male, and first instar
nymph are shown here after Hodgson & Trencheva (2008), for completeness with
kind permission of Prof. Chris Hodgson we show here all stages to see the possible
differences, among different developmental stages, what could be a good example for
study of other species in the family.
Acanthococcidae 137
Ecology
Host plant: Myrtus communis, Quercus pubescens, Quercus sp.
Distribution: Bulgaria, Cyprus Island, Hungary and Israel.
Biology: On twigs and stems.
Comments
This species differs from A. roboris by having only two size of macrotubular ducts on
venter, by having smaller number of enlarged spines on dorsal abdominal segments VII
and VIII, by having smaller number of cruciform pores. Also these two species are
different from each other by some morphological characters of the first-instar nymph,
and by different size categories of most of the body parts (Hodgson & Trencheva, 2008).
Figure 52. Acanthococcus melnikensis (Hodgson & Trencheva, 2008), second instar a -
female and b - male. After Hodgson & Trencheva (2008).
Figure 53. Acanthococcus melnikensis (Hodgson & Trencheva, 2008), male prepupa. After
Hodgson & Trencheva (2008).
Acanthococcidae 139
Figure 54. Acanthococcus melnikensis (Hodgson & Trencheva, 2008), adult male. After
Hodgson & Trencheva (2008).
Acanthococcus onukii (Kuwana, 1902) (Fig. 55) Redescription

Eriococcus onukii Kuwana, 1902: 51.
Syntypes: Female. Japan (Honshu, Tokyo), on Arundinaria hindsii var. graminae, by I.

Kuwana., female. Deposited in UCDC, and ITLJ.
Lit.: Borchsenius, 1950: 236; Danzig, 1975: 46; Fang et al., 2001: 104; Fernald, 1903:
76; Hoy, 1963: 105; Hua, 2000: 137; Kawai, 1972: 153; 1980: 129; Kozár & Walter,
1985: 74; Köhler, 1998: 380; Kwon & Han, 2003: 151; Paik, 1978: 166; Takahashi,
1957: 7; Tang, 2001: 3; Tang & Hao, 1995: 450; Tao, 1999: 32; Wang, 1974: 329; 2001:
208; Yang, 1982: 105 (Miller et al., 2013).
Nidularia onukii; Lindinger, 1933a: 116. Change of combination.

Acanthococcus onukii; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus onukii; Miller & Gimpel, 2000: 285. Revived combination.
Acanthococcus onukii; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Female is oval in shape. Sac strongly convex. Its color is white to gray. With four or five
transverse ridges on dorsum. Adult female reddish brown and turns red when boiled in
KOH. Ovisac 3.5 mm long, 2.0 mm wide, the female somewhat smaller, grayish white,
nearly oval. Male sac is similar to female, but smaller (Kuwana, 1902).
Mounted female
Slide-mounted adult female 3.0 mm long, 2.5 mm wide. Antennae 7 segmented, 230 µm
long, second and third segments subequal, the third segment is the longest, all segments
except third with few rather long spine-like setae, fifth and sixth segment with one
falcate sensory seta, and the seventh with 3 falcate setae. Frontal lobe present. Anal
lobes protruding, apically rounded, moderately sclerotized, serrate on inner margin, with
3 dorsal equally long conical setae and with three short hair-like setae and a long apical
setae.
Venter: Labium small, well developed, 3 segmented; according to Kuwana (1902) the
stylet loop is short, but on our material is medium long, much longer than labium. Legs
subequal, comparatively small, tarsus longer than tibia, tarsal digitules longer than claw,
claw with minute digitules. Posterior and median coxae with spinulae; tarsus longer than
tibia, tarsal and claw digitulae knobbed, longer than claw. Normal flagellate setae in
median areas, short and slender but stouter and stiff in lateral areas, with rather large
pores especially in abdominal region. Quinquelocular pores in rows on last five abdominal
segments, scattered on thorax and head. Macrotubular ducts of three sizes, the largest
macrotubular ducts on margin, medium size macrotubular ducts scattered on abdominal
Acanthococcidae 141
Figure 55. Acanthococcus onukii (Kuwana, 1902), female. Original.

segments, thorax and head, smallest size on medial area of the surface. Cruciform pores
present in few numbers on margin.
Dorsum: Enlarged setae, stout spine-like setae of 2 or 3 sizes, those on the margin
somewhat larger, sides straight, apices pointed, some medium sized spines very short and
wide, without dorsal setae on abdominal segment VIII, anterior of anal ring; largest setae
form sparse, transverse segmental rows; on abdominal segment VII, number of enlarged
setae about 6, with 2 or 3 setae present on each margin of all abdominal segments.
Macrotubular ducts of one size, scattered on surface. Microtubular ducts scattered
through surface among dorsal enlarged setae. Anal ring oval, sclerotized, with 8 setae.
Cauda present (Redescription based on material; Japan, Tokyo, 2 females on Bambusa sp.,
25. ix. 1913, collected and determined by Kuwana).
Ecology
Host plant: Arundinaria hindsii var. graminae, Bambusa nana var. normalis, Sasa sp., S.
peniculata.
Distribution: China, Japan, Russia.
Biology: On twigs and stem.
Acanthococcus populi Matesova, 1967 (Fig. 56) Redescription

Acanthococcus populi Matesova, 1967: 1195.
Holotype: Female. Kazakhstan (Alma-Ata Oblast, Charyn River, Sartagoy), on

Populus pruinosa, 22.vi.1964, G. Matesova. By original designation. Deposited in
ZMAS. Notes: 5 paratypes on 2 slides with same data as holotype in; a second
collection of paratypes from Alma-Ata Oblast, between Chilik and Ayak-Kalkan, on
Populus diversifolia, 05.v.1963, Makarov with 3 females on 3 slides deposited in ZMAS.
Lit.: Danzig, 1977b: 50; 1980: 207; Koteja & Zak-Ogaza, 1981: 513, 514; Kozár,
2009:94; Kozár & Walter, 1985: 74; Matesova, 1971: 27; Tang & Hao, 1995: 449; Tao,
1999: 33, Wang, 2001: 208 (Miller et al., 2013).
Eriococcus populi; Tang & Hao, 1995: 486. Change of combination.

Acanthococcus turanicus Matesova, 1967: 1197.
Holotype: Female. Kazahstan (Alma Ata Oblast, Chulac-Tau Ridge, Kzyl-Aus
Gorge), By original designation. 15 paratypes on 15 slides from Kazakhstan and
Tajikistan, deposited in ZMAS; paratypes also in AAKA. Synonym nova based on
type material.
Eriococcus turanicus; Tang & Hao, 1995: 450. Change of combination.
Acanthococcus populi; Köhler, 1998: 382, 383. Revived combination.
Eriococcus populi; Miller & Gimpel, 2000: 306. Revived combination.
Acanthococcus populi; Kozár, 2009: 93. Revived combination.
Acanthococcidae 143
Description
Unmounted female
Adult female wine red, almost black; felt-like test cream-white.
Mounted female
250–290 µm long, length of segments: I: 44–75 µm, II: 35–50 µm, III: 45–60 µm, IV:
29–45 µm, V: 17.5–26 µm, VI: 20–24.2 µm, and VII: 35–48.4 µm long; each segment
covered with a few, strong hair-like setae; apical segment with apical seta 52.5–65 µm
long; apical segment also with 3 sensory falcate setae, each 30.0–37.5 µm long; segment
VI with 1 sensory falcate seta 22.5–25.0 µm long, segment V with 1 sensory falcate seta
20–22 µm long. One or two frontal tubercles present at the base of antennae. Eyes
situated on venter near margin. Anal lobes strongly developed, each with 3 enlarged setae,
each 20–37 µm plus 2 or 3 microtubular ducts on dorsal surface; apical seta 193–303 µm;
ventral hair-like subapical seta 84–165 µm long.
Venter: Labium 160–200 µm long, 105–130 µm wide, with 5 pairs of setae, median
setae on apex of labium 22.5–25.0 µm long. Legs well developed; lengths of segments
and digitules of prothoracic legs; coxa: 60–75 µm, trochanter: 45–65 µm, femur: 115–160
µm, tibia: 90–123 µm, tarsus: 125–130 µm and claw: 27.5–40 µm, trochanther + femur:
160–180 µm, tibia + tarsus: 210–240 µm, tarsal digitules: 52–55 µm, claw digitules: 27.5–
30 µm long; lengths of segments and digitules of mesothoracic legs; coxa: 60–80 µm,
trochanter: 45–60 µm, femur: 120–125 µm, tibia: 105–120 µm, tarsus: 120–135 µm and
claw: 27.5–40 µm, trochanther + femur: 165–175 µm, tibia + tarsus: 230–240 µm, tarsal
digitules: 52–60 µm, claw digitules: 25–35 µm long; lengths of segments and digitules
of metathoracic legs; coxa: 62–85 µm, trochanter: 40–60 µm, femur: 125–160 µm, tibia:
123–135 µm, tarsus. 125–150 µm and claw: 35–42.5 µm, trochanther + femur: 165–190
µm, tibia + tarsus: 235–260 µm, tarsal digitules: 55.0–62.5 µm, claw digitules: 25–35 µm
long; tarsal and claw digitules slightly knobbed, longer than claw; metathoracic coxae
each with spinulae on ventral surface on coxa; tibiae each with 4 setae (median seta
absent, present on prothoracic legs), tarsi each with 5 setae. Length of spiracles 45–70
µm; diameter of spiracular peritreme 25–35 µm, posterior spiracles slightly larger than
anterior. Derm with a sparse covering of scattered flagellate hair-like setae, each 15–80
µm long. Enlarged setae situated on submargin in 1 or 2 rows, each 15–20 µm long.
Multilocular pores each 4–5 µm in diameter and with 3-5 loculi, distributed in sparse
bands on all abdominal and thoracic segments and head. Macrotubular ducts of one size,
each 5.0–7.5 µm wide and 25–32.5 µm long, present on scattered on abdomen, thorax
and head. Cruciform pores few, each, 3–4 µm in diameter, distributed on the submargin
of thorax, head and first abdominal segments.
Dorsum: Marginal enlarged setae slightly truncated, 33–70 µm, forming a marginal row
and not strongly differentiated from dorsal setae; 2-4 setae on margin of abdominal
segments, other dorsal setae slightly truncated almost parallel ssides, each seta 11–33 µm
long, those on thorax, head and anterior abdominal segments slightly larger than setae
on posterior 2 or 3 abdominal segments; arranged in transverse rows across each body
segment, rows irregular on head. There are 3-8 setae on abdominal segment VIII, and
about 20-28 on abdominal segment VII segment. Macrotubular ducts, 7–10 µm wide and
30–35 µm long, scattered throughout dorsum, generally in segmental bands. Microtubular
ducts, each 7–10 µm long scattered over dorsum. Anal ring strongly sclerotized, with a
row of pores on each side, 60–70 µm in diameter, with 8 setae, each 95–193 µm long; anal
ring situated on margin of dorsum. Cauda present with large spinules as on dorsal surface
(after Matesova (1967); Tang & Hao (1995); with modifications and new measurements
besed on type material).
Ecology
Host plant: Populus sp., P. euphratica, P. pruinosa, Salix sp. S. wilhelmsiana.
Distribution: China, Kazakhstan.
Biology: In bark crevices of trunk and thick branches. Often in large colonies. Female
appears in egg sacs in the begining of May. Egg laying starts in the last days of June.
Comments
According to our study of the type material of A. turanicus Matesova (1967), we found
that the distinguishing characters mentioned by Matesova (1967) for the species are highly
variable and that its characters overlap with those of A. populi. Thus we here synonymize
the two species, with A. turanicus becoming becoming a junior synonym of A. populi.
Acanthococcidae 145
Figure 56. Acanthococcus populi Matesova, 1967, female. Original.

Acanthococcus ribesiae Borchsenius, 1960 (Fig. 57)

Acanthococcus ribesiae Borchsenius, 1960a: 193.
Lectotype: Female. Kazakhstan (Dshungar Alatau, Marka-Kul Sea). One

paralectotype with the same data. Designated here: Deposited in ZMAS.
Lit.: Borchsenius, 1963: 97; 1973: 97; Danzig, 1972a: 198; Hoy, 1963: 112; Koteja &
Zak-Ogaza, 1981: 514; Matesova, 1968: 114 (Miller et al., 2013).
Eriococcus ribesiae; Hoy, 1963: 112. Change of combination.

Acanthococcus ribesiae; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus ribesiae; Tang & Hao, 1995: 448. Revived combination.
Acanthococcus ribesiae; Köhler, 1998: 382. Revived combination.
Eriococcus ribesiae; Miller & Gimpel, 2000: 316. Revived combination.
Acanthococcus ribesiae; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female elongated oval, 2.5 mm long. Egg sacs felt-like, cream-white, 2.5 mm long,
1.5 mm wide.
Mounted female
Antennae 7 segmented, length of segments: I: 40.0 µm, II: 40.0 µm, III: 42.0 µm, IV:
46.0 µm, V: 24.0 µm, VI: 24.0 µm, and VII: 38.0 µm long, each segment covered with
a few, hair-like setae (except third segment); apical segment with 3 sensory falcate setae,
segment V and VI with 1 sensory falcate seta. Frontal lobe and tubercle present. Eyes
situated on venter near margin. Anal lobes strongly developed, with 3 enlarged setae,
apical seta 180–190 µm; two ventral subapical hair-like setae 60–70 µm, suranal setae 70
µm long.
Venter: Labium well developed, apical segment with 5 pairs of setae, median setae on
apex of labium long. Legs well developed: prothoracic coxa: 100 µm long, tarsus+claw:
170.0 µm, tarsal digitules and claw digitules slightly knobbed. Meso- and metathoracic
coxae with spinulae on ventral surface. Tibiae of legs with 4 setae (median seta absent).
Setae in median areas flagellate, setae in lateral margin enlarged, in a row. Multilocular
pores with 5 loculi, numerous, distributed in bands on all abdominal segments and
scattered on thorax and head. Macrotubular ducts small, 4.0 µm wide and 16.0 µm long,
scattered on median areas of abdominal segments, larger ducts present a marginal band.
Cruciform pores in sparse bands on thorax and first abdominal segments.
Dorsum: Marginal enlarged setae straight, blunt, the larger 25–30 µm long, medium
sized 20–25 µm and smaller setae 14–18 µm long, 2 lateral setae present on margin of
abdominal segments, dorsal setae much shorter, arranged in sparse transverse rows and
bands. Enlarged setae on other surface much shorter than marginal ones, on the median
Acanthococcidae 147
Figure 57. Acanthococcus ribesiae Borchsenius, 1960, female. After Borchsenius (1960),
with modifications.
part of thorax a longitudinal band of large spines present. Two setae on abdominal
segment VIII, and about 16 on abdominal segment VII. Macrotubular ducts of one
size, 8.0 µm wide and 20.0 µm long, scattered throughout dorsum. Microtubular ducts
long, scattered over dorsum. Anal ring strongly sclerotized, with partly double rows of
pores, with 8, or ten setae, 110–120 µm long; anal ring situated on margin of dorsum.
Cauda present, triangular (after Borchsenius (1960a), with modifications based on type
material).
Ecology
Host plant: Ribes meyeri.
Distribution: Kazakhstan, Russia.
Biology: In bark crevices of trunk and wide branches. Often in large colonies. Female
appears in egg sacs in the end of May. Egglaying finished by the end of July. Heavily
infested branches of plants dryed out (Borchsenius, 1960a).
Acanthococcus roboris (Goux, 1931) (Fig. 58) Redescription

Eriococcus roboris Goux, 1931: 58.
Lectotype: Female. The slide contains 2 females, the lectotype designated here
(marked), following the drawing of Goux. France: Rhone, Courzieu, by L. Goux, on
Quercus robur, 30. vii. 1927. Paralectotypes: one female on slide of lectotype (MNHN
14508-01). Two slides MNHN 14508-02-03, with same data as the lectotype. Seven
slides MNHN 14507-01-07 from the same place as lectotype collected in 30.vii. 1928.
Notes: According to Hoy (1963), Lindinger (1936, 1957) erroneously considered
Eriococcus roboris to be a synonym of Nidularia aceris (Signoret, 1875) and E. roboris to
be a synonym of Nidularia quercus (Comstock, 1881).
Additional material examined: 8 ♀♀, Turkey, Hakkari road, on Quercus sp., N:
37°41’644’’; E: 043°56’266’’; F. Kozár and M.B. Kaydan, 23.v.2008 (KPCT: 3475); 8
♀♀, Turkey, Bitlis river, on Quercus sp., N: 38°17’051’’; E: 041°58’925’’; M.B. Kaydan,
31.v.2007 (KPCT: 3455); 8 ♀♀, Turkey, Bitlis Narlıdere, on Quercus sp., N: 38°13’768’’;
E: 041°54’007’’; M.B. Kaydan, 31.v.2007 (KPCT: 3475).
Common name: Oak felt scale
Lit.: Borchsenius, 1949: 346; Danzig, 1964: 632; 1971a: 821; 1980: 207; Erdős,
1957: 370; Foldi, 2000: 81; 2001: 305; Hadzibejli, 1956a: 50; 1983: 269; Hodgson
& Trencheva, 2008: 2; Kaydan et al., 2007: 90; Kiritchenko, 1940: 136; Kosztarab
& Kozár, 1988: 277; Koteja & Zak-Ogaza, 1981: 514; Kozár et al., 2013: 55; Kozár
& Kosztarab, 1982: 2040; Kozár & Walter, 1985: 74; Köhler, 1998: 382; Lindinger,
1957: 543; Longo et al., 1999: 121; Marotta, 1993: 160; Ouvrard & Kozár, 2009:
102; Pellizzari & Kozár, 2011: 66; Rogojanu, 1966: 323; Tang & Hao, 1995: 450;
Tereznikova, 1981: 36; 1982: 35; Ter-Grigorian, 1983: 879; Ülgentürk, et al., 2003: 52;
Zahradník, 1972: 402; 1977: 121.
Acanthococcidae 149
Nidularia roboris; Lindinger, 1933a: 116. Change of combination.

Acanthococcus roboris; Borchsenius, 1949: 346. Change of combination.
Eriococcus roboris; Hoy, 1963: 112. Revived combination.
Acanthococcus roboris; Kosztarab & Kozár, 1978: 74. Revived combination.
Eriococcus roboris; Miller & Gimpel, 2000: 316. Revived combination.
Acanthococcus roboris; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Rather large, 3.25–5 mm long; venter wider than dorsum-dorsum 2.1–3.5 mm widest,
total width 4.12 mm widest. More or less oval but slightly more pointed posteriorly.
Ovisac felted, cream-white, up to 9 mm long, 6 mm wide. Adult female oval, red raspberry
colored.
Mounted female
340–360 µm, length of segments: I: 60–85 µm, II: 50.0–67.5 µm, III: 50.0–67.5 µm, IV:
52.5–80.0 µm, V: 30–35 µm, VI: 25–30 µm, and VII: 40.0–47.5 µm long, the segments
covered with a few, strong hair-like setae (third segment without setae); apical segment
with apical seta 47.5–55 µm long; apical segment also with 3 sensory falcate setae, 25–40
µm long; segment VI with 1 sensory falcate seta 25.0–27.5 µm long, segment V with 1
sensory falcate seta 25 µm long. Frontal lobes and frontal tubercle present. Eyes situated
on venter near margin. Anal lobes strongly developed sclerotised, with 3 enlarged setae,
40–60 µm long plus 3–5 microtubular ducts on dorsal surface; apical seta 285–325 µm;
Venter: Labium 195–205 µm long, 122.5–180.0 µm wide, median setae on apex of labium
long, 35–40 µm long; stylet loop relatively short, reaching to the median coxae. Legs
well developed; lengths of segments and digitules of prothoracic legs; coxa: 85–110 µm,
trochanter: 65–70 µm, femur: 142.5–165 µm, tibia: 120–140 µm, tarsus: 145–165 µm and
claw: 27.5–35 µm, tarsal digitules: 45.0–52.5 µm, claw digitules: 27.5–35 µm long; lengths
of segments and digitules of mesothoracic legs; coxa: 85–105 µm, trochanter: 65–75 µm,
femur: 150–165 µm, tibia: 135–145 µm, tarsus: 155–160 µm and claw: 35.0–37.5 µm,
tarsal digitules: 52.5–55.0 µm, claw digitules: 32.5–35.0 µm long; lengths of segments and
digitules of metathoracic legs; coxa: 90–115 µm, trochanter: 55–85 µm, femur: 160–170
µm, tibia: 145–155 µm, tarsus: 160–177.5 µm and claw: 35 µm, tarsal digitules: 52.5–60.0
µm, claw: digitules 27.5–37.5 µm long; meso- and metathoracic coxae with spinulae on
ventral surface; tibiae of meso and metathoracic legs with 4 setae (median seta absent),
tibiae of prothoracic legs with 5 setae (median seta present), tarsi with 6 setae; tarsal and
claw digitules knobbed, slightly longer than claw. Length of spiracles 75 µm; diameter of
spiracular peritreme 40.0–47.5 µm, posterior spiracles slightly larger than anterior. Setae
in median areas flagellate, 25–110 µm long, setae in lateral areas spine-like, 25–30 µm
long. Multilocular pores 3–5 µm in diameter with 5 loculi, distributed in sparse bands on
all abdominal segments and scattered on thorax and head. Macrotubular ducts of three
sizes, smallest macrotubular ducts 2.5–3.0 µm wide and 12.5–17.5 µm long, present on
only median areas of abdominal segments, medium macrotubular ducts 5–7 µm wide and
25–32.5 µm long, present on submargin of body, larger macrotubular ducts 7.5–9.0 µm
wide and 20–25 µm long, on submarginal band on abdomen, thorax and head. Cruciform
pores generally on submedian area in band on thorax and abdominal segments and a few
on head, 2.5–3.0 µm in diameter.
Dorsum: Marginal enlarged setae 45–70 µm long, truncated, forming a marginal row and,
other dorsal setae conical, truncated, seta 30.0–67.5 µm long; arranged in transverse rows
across body segments, rows irregular on head, on the last abdominal segment 4-5 enlarged
setae situated. Macrotubular ducts, 6.0–7.5 µm wide and 25–30 µm long, scattered
throughout dorsum, generally in segmental bands. Microtubular ducts long, 8–10 µm
long with oval dermal orifice, scattered over dorsum. Anal ring strongly sclerotized, with
25–28 pores on each side, 60–70 µm in diameter, with 8 setae, 135–165 µm long; anal ring
situated on margin of dorsum. Cauda present (after Goux (1931); Hodgson & Trencheva
(2008); Kosztarab & Kozár (1988), with some modifications).
Other stages
Mounted first-instar nymph (Fig. 58, top right).
strong spines with pointed apices of one size, forming middorsal longitudinal pair of
rows, the submedian row contains five large spines, followed by 5 or 6 medium small
setae and two pairs of large spines in front of the anal ring. Microtubular ducts long,
few in two middorsal longitudinal rows on both side. Venter with transverse rows of
six small hair-like setae on each abdominal segment; median setae longer than others.
Cruciform pores present on thoracic segments, with 1 pair of quinquelocular pores on
each thoracic segment, 1 near each spiracle, plus 1 pair on frons, and two longitudinal
submedian rows on abdomen. Anal ring with 8 spine-like setae (after Goux (1931) and
Hodgson & Trencheva (2008)).
Ecology
Host plant: Aesculus sp., A. hippocastanum, Castanea sp., C. sativa, Pterocarya pterocarpa,
Quercus sp., Q. robur imeretina, Q. petraea, Q. pubescens, Q. robur.
Distribution: Armenia, Azerbaijan, former Czechoslovakia, France, Georgia, Hungary,
Italy, Kazakhstan, Netherlands, Portugal, Romania, Russia, Switzerland, Turkey, Ukraine.
Biology: Lives in bark crevices of trunk and branches, on the stem and base of young
twigs, or at base of buds or twig forkings, sometimes under the litter on the roots. Female
appears with egg sacs in May. Lays red eggs in June, these hatched at end of July in
Hungary; males not known (Kosztarab & Kozár, 1988).
Acanthococcidae 151
Figure 58. Acanthococcus roboris (Goux, 1931), female, original. First instar on top right,
after Hodgson & Trencheva (2008).
Comments
Comparing the size categories given by Hodgson & Trancheva (2008), and what is
presented here are very similar, but very different from data given for A. melnikensis.
Acanthococcus rosannae (Tranfaglia & Esposito, 1985) (Fig. 59)

Eriococcus rosannae Tranfaglia & Esposito, 1985: 130.
Holotype: Female. Italy (Monte Faito), on Castanea sativa, 07.v.1983-31.vii.1983.

Deposited in IFSP.
Lit.: Kozár & Walter, 1985; Köhler, 1998: 383; Longo et al., 1999: 113; Miller &
Gimpel, 1996: 113; Tang & Hao, 1995: 452. (Miller et al., 2013)
Acanthococcus rosannae; Longo et al., 1995: 113. Change of combination.

Eriococcus rosannae; Miller & Gimpel, 2000: 319. Revived combination.
Acanthococcus rosannae; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female oval-elongate, 2.46–5.68 mm long, 1.1–3.72 mm wide.
Mounted female
Antennae robust, 7 segmented, 350–420 µm long, fourth segment longer than second
and third. Antenna with hair-like setae, longer than segments; sensory setae on segments
V, VI and VII, long, narrow; hair-like setae present on third segment, too. Frontal lobes
present, weakly developed. Anal lobe with two hair-like setae; suranal seta also very long
as long as anal lobe.
Venter: Apical segment of labium with 5 pair of equal long setae; stylet loop very long,
almost as long as the body. Legs well developed: posterior tibia-tarsus 221.4–304.9 µm,
tibia 174.2–196 µm long, ratio of tibia and tarsus 0.60–0.78 µm, claw with denticle,
claw and tarsal digitules knobbed, longer than claw. Coxae without translucent pores,
with spinulaes on anterior side of all legs. Quinquelocular and some trilocular pores in
transverse bands across abdominal sternites, and scattered on prosoma. Macrotubular
ducts two of sizes; larger macrotubular ducts present around submargin, narrower
macrotubular ducts present on median part of segments. Some microtubular ducts
present on margin. Setae, normal flagellate setae in median areas, most of them long, and
slender, on last abdominal segments longer than width of the segments. Cruciform pores
in wide bands on margin.
Dorsum: Enlarged setae cylindrical, sides straight, apices truncate, marginal setae slightly
larger than other setae on dorsum, other dorsal setae form sparse, transverse segmental
bands, 4 or 5 lateral setae on margin of abdominal segments. Setae of two sizes, 54.4–83.4
Acanthococcidae 153
Figure 59. Acanthococcus rosannae (Tranfaglia & Esposito, 1985), female. After Tranfaglia
& Esposito (1985) with modifications.
µm long; Macrotubular ducts shown one size, frequent on dorsum. Microtubular ducts
narrow, long, elongate, with bifurcated orifice; evenly distributed, form transverse bands
on dorsum of prosoma, rows on abdomen. Anal ring oval, sclerotized, with two rows
of pores with 8 setae. Anal lobes protruding, about twice as long as wide, sclerotized,
serrate on inner margin, with 3 narrow, dorsal conical setae, posterior 76.2–98.0 µm,
basal median 61.7–79.8 µm, basal marginal 76.2–90.7 µm long, apical setae 254.1–392.1
µm long and with some microtubular ducts. Cauda present, weakly developed (after
Tranfaglia & Esposito (1985), with modifications based on lectotype and paralectotypes).
Ecology
Host plant: Castanea sativa.
Biology: In bark crevices of trunk and branches of its host at elevation about 500 m
above see level. Female appears in egg sacs in May.
Comments
The collection data in the manuscript differs from the original label data, in manuscript
written 14.vi. 83, however, on the slide label and cover envelop it is 14.v.1983. Tranfaglia
and Espinosa (1985) mentioned in the original description “tipo and paratipi” (11 slides),
and following them Miller & Gimpel (2000) as “Holotype”. However on the slides it was
not marked, and because of this here a lectotype and 10 paralectotypes are designated.
The authors thank Prof. Dr. Antonio Garonna, Prof. Dr. Francesco Pennacchio and
Bruno Espinosa for the help in search of the type material and clarification of the status
of the types.
Acanthococcidae 155
Acanthococcus salicis (Borchsenius, 1938) (Fig. 60)

Eriococcus salicis Borchsenius, 1938, 135.
Lectotype: Female. Russia (Siberia, Guberoro), on Salix sp., 25.vi.1934, by S.V.

Brown. By subsequent designation Danzig, 1980: 208 (type no. S-T.7). Deposited in
ZMAS.
Lit.: Borchsenius, 1950: 120; 1963: 23; 1973: 231; Danzig, 1975: 43; 1977b: 57; 1980:
205, 1986: 239; 1988: 708; Hua, 2000: 137; Koteja & Zak-Ogaza, 1981: 514; Köhler,
1998: 382; Matesova, 1967: 1194; Tang, 1977: 45; Tang & Hao, 1995: 449; Tao, 1999:
33; Wang, 2001: 208; WangZQ, 1982: 43; Xie, 1998, 97; Xu, 1983: 77; Zhang & Hu,
1980: 152–153 (Miller et al., 2013).
Acanthococcus salicis; Borchsenius, 1949: 345. Change of combination.

Nidularia salicis; Lindinger, 1957: 544. Change of combination.
Eriococcus salicicola;Tang, 1984: 125. Nomen nudum; discovered by Tang & Hao, 1995: 489.
Notes: Tang (1984) used the name Eriococcus salicicola n. sp. and indicated that it occurred
widely in Northwest China on willows. He indicated in the paper's introduction that
the new species would be published in the future. We believe this species was described
as Gossyparisa [sic] salicicola Borchsenius. This species should be attributed to Tang not
Borchsenius.
Gossyparia salicicola; Tang in Tang & Li, 1988: 72. Described: female. Synonymy by Tang
& Hao, 1995: 595. Notes: In addition to treating this species as a junior synonym of
Eriococcus salicis Borchsenius, we believe that this is the first time that Eriococcus salicicola
Tang was described. It was previously mentioned in Tang (1984), but because there was
no description it is now placed as a nomen nudum.
Eriococcus salicis; Hoy, 1963: 114. Revived combination.
Acanthococcus salicis; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus salicis; Miller & Gimpel, 2000: 325. Revived combination.
Acanthococcus salicis; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Ovisac is oval, creamy grey in color. Adult female elongate oval, dark violet. Eggs are
crimson in color. (Borchsenius, 1938, 1949; Danzig, 1980)
Mounted female
Adult female elongated oval, about 2.1–2.5 mm long, 1.3–1.4 mm wide. Antennae 7
seldom 8 segmented, I: 30.4 µm, II: 35.4 µm, III: 52.6 µm, IV: 31.9 µm, V: 24.0 µm, VI:
24.0 µm, and VII: 38.1 µm long, sometimes segment four divided, the segments covered
with a few, hair-like setae apical segment with 3 sensory falcate setae, segment V and VI
with 1 sensory falcate seta. Frontal lobe present. Eyes situated on venter near margin.
Anal lobes strongly developed, with 3 enlarged setae, about 55 µm long, apical seta 220
µm; two ventral subapical hair-like 80 µm, and 27 µm long.
Venter: Labium well developed, wide, apical segment with 5 pairs of setae, median setae
on apex of labium short; stylet loop long, near to base of posterior legs, on the drawing
of Tang & Hao (1995) it is short. Legs well developed: all coxae with spinulae on ventral
surface, according to Borchsenius (1949) there are pores too, however on type material
studied by us, these were not seen. Femur of posterior legs 125 µm, tibia 110 µm, tarsus
125 µm long, claw with denticle, claw and tarsal digitules knobbed, longer than claw.
Tibiae with 4 setae (median seta absent). Setae in median areas short, flagellate, setae in
lateral margin spine like, in a sparse band. Multilocular pores with 5 loculi, numerous,
distributed in bands on all abdominal segments and scattered on thorax and head.
Macrotubular ducts only one sized, 6.0 µm wide and 27.0 µm long, scattered on median
areas of abdominal segments, and in a marginal band. Cruciform pores in sparse band on
thorax and first for abdominal segments. Suranal setae 80 µm long.
Dorsum: Marginal enlarged setae elongate, slender, curved, sides concave basally, apices
rounded, 44–56, µm long, dorsal setae in midline capitated, 35–44 µm long, form two
longitudinal rows, others 18–30 µm long, arranged in transverse rows and bands. There
are 8 setae on abdominal segment VIII, and 20 on abdominal segment VII. Macrotubular
ducts 8.0 µm wide and 27.0 µm long, numerous throughout dorsum. Microtubular ducts
long scattered over dorsum. Anal ring strongly sclerotized, with partly double row of
pores, with 8 setae, each 110–140 µm long; anal ring situated on margin of dorsum.
Cauda triangular (after Borchsenius (1949); Danzig (1980); with modifications based on
type material).
Ecology
Host plant: Salix sp., S. alba.
Distribution: China, Mongolia, Russia, Turkey.
Biology: On the stem, branches, and twigs, partly pest on Salix. Danzig (1986) states that
this species is an oligophagous species living on willow and that egg laying was seen in
mid-June. Xu (1983) gives a detailed description of the life history.
Comment
According to drawings of Danzig (1986) and Tang & Hao (1995) there are several setae
on abdominal segment VIII, anterior to the anal ring and anal lobes apparently with
4 setae; microtubular ducts short, with 2 sclerotized areas. These characters were not
verified on type material.
Acanthococcidae 157
Figure 60. Acanthococcus salicis (Borchsenius, 1938), female. After Danzig (1980) with
modifications.
Acanthococcus sasae Danzig, 1975 (Fig. 61)

Acanthococcus sasae Danzig, 1975: 69.
Holotype: Female. Russia (Kunashir Island, Sernovodsk), on Sasa sp., 11.vi.1967, by

E. Danzig. By original designation. Deposited in ZMAS.
Lit.: Danzig, 1977b: 52; 1978a: 13; 1980: 206; 1988: 708; Fang et al., 2001: 104; Kozár
& Walter, 1985: 74; Tang & Hao, 1995: 451 (Miller et al., 2013).
Eriococcus sasae; Tang & Hao, 1995: 490. Change of combination.

Acanthococcus sasae; Köhler, 1998: 383. Revived combination.
Eriococcus sasae; Miller & Gimpel, 2000: 328. Revived combination.
Acanthococcus sasae; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Female is round and brown, 2.5 mm long. Ovisac is oval, cream colored, markedly convex,
with 2 longitudinal and several transverse ribs. Eggs raspberry-colored (Danzig, 1975).
Mounted female
Antennae 7, seldom 8 segmented, I: 30.4 µm, II: 35.4 µm, III: 52.6 µm, IV: 31.9 µm,
V: 24.0 µm, VI: 24.0 µm, and VII: 38.1 µm long, sometimes segment four divided, the
segments covered with a few, hair-like setae apical segment with 3 sensory falcate setae,
segment V and VI with 1 sensory falcate seta. Frontal lobe probably present. Eyes
situated on venter near margin. Anal lobes strongly developed, with 3 enlarged setae.
Venter: Labium well developed, wide, apical segment with 5 pairs of setae, median setae
on apex of labium long. Legs well developed: coxae with spinulae on ventral surface;
median seta present on tibiae of front legs, others with 4 setae, tarsal and claw digitules
knobbed, longer than claw. Setae in median areas short, flagellate, setae in lateral margin
enlarged, spine like, in a sparse row; suranal and two subapical setae short. Multilocular
pores with 5 loculi, numerous, distributed in bands on all abdominal segments and
scattered on thorax and head. Macrotubular ducts two of sizes, larger ones on margin;
smaller ones scattered through. Cruciform pores in sparse band on thorax and first for
abdominal segments.
Dorsum: Marginal enlarged setae conical, sides straight, apices acute, marginal setae
slightly longer than other dorsal enlarged setae, setae abundant over surface, without
enlarged setae anterior of anal ring, form bands around margin and two longitudinal
bands in midline, arranged in dense transverse rows and bands, on all segments. Setae
absent on abdominal segment VIII, about 20 on abdominal segment VII. Macrotubular
ducts numerous throughout dorsum. Microtubular ducts long scattered over dorsum.
Anal ring strongly sclerotized, with partly double row of pores, with 8 setae; anal ring
Acanthococcidae 159
Figure 61. Acanthococcus sasae Danzig, 1975, female. After Danzig (1975) with
modifications.
situated on margin of dorsum. Cauda triangular (Danzig (1975, 1980), with modifications
based on type material).
Ecology
Host plant: Sasa sp.
Distribution: Russia.
Biology: This species is always associated with bamboo growing in open sectors, not
extending beneath the forest canopy. An abundant species, females overwintering in leaf
axils. Oviposition was noted at the beginning of July, general hatching of the first-instar
nymphs by late July. First-instar nymphs migrate to young plants (Danzig, 1975).
Acanthococcus shiraiwai (Kuwana, 1931) (Fig. 62)

Eriococcus shiraiwai Kuwana in Kuwana & Muramatsu, 1931: 654.
Syntypes: Female. Japan (Honshu, Yokohama customs), on unidentified cactus, by H.

Shiraiwa. Deposited in ITLJ.
Lit.: Hoy, 1963: 115 (Miller et al., 2013).
Acanthococcus shiraiwai; Miller, 1996: 79. Change of combination.

Eriococcus shiraiwai; Miller & Gimpel, 2000: 332. Revived combination.
Acanthococcus shiraiwai; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female is oval, tappering behind. 3.5 mm long, 1.5 mm wide.
Mounted female
Slide-mounted adult female 3 mm long, 2 mm wide. Antennae 6 segmented, third
segment the longest. Anal lobes protruding, apically rounded, moderately sclerotized,
with 3 dorsal strong conical setae, one smaller than others and with three ventral hair-like
setae, suranal setae similar size and with a very long apical setae
Venter: Legs typical well developed, tibia with 4 setae, median seta absent, tarsus slightly
longer than tibia, tarsal digitules longer than claw, claw digitules as long as claw, digitules
knobbed.
Dorsum: Enlarged setae, elongate conical, sides straight, apices acute, 2 or 3 sizes,
abundant over dorsum setae all approximately same size. Anal ring oval, sclerotized, with
8 setae and two rows of pores (Kuwana & Muramatsu, 1931).
Acanthococcidae 161
Figure 62. Acanthococcus shiraiwai (Kuwana & Muramato, 1931), female. After Kuwana &
Muramato (1931) with modifications.
Ecology
Host plant: On unidentified cactus
Distribution: Japan, Mexico.
Biology: On plant surface.
Comments
Cactus originated from Mexico and intercepted at the Yokohama customs. The validity
of this species needs further study. Its establishment in Japan till now was not confirmed.
The Authors discuss similarity with “Eriococcus” stanfordianus (Ferris, 1920), what is now
“Eriococcus” dubius (Cockerell, 1896), which is a polyphagous species infesting many
species of plants.
Acanthococcidae 163
Acanthococcus spiraeae Borchsenius, 1949 (Fig. 63), Redescription

Acanthococcus spiraeae Borchsenius, 1949: 348.
Lectotype: Female. Georgia (Tiflis, Tbilisi), on Spiraea hypericifolia twigs, 17.vii.1934.

By subsequent designation Danzig, 1996: 522. Deposited in ZMAS. Notes: Two
paralectotype females on same slide lectotype (Danzig, 1996).
Lit.: Borchsenius, 1950: 121; 1963: 147; 1973: 148; Danzig, 1975: 45; 1977a: 200;
1978b: 76; 1988: 708; 1996: 522; Hadzibejli, 1983: 269; Hoy, 1963: 117; Hua, 2000:
137; Koteja & Zak-Ogaza, 1981: 514; Kozár & Walter, 1985: 74; Lindinger, 1957: 543;
Matesova, 1968: 114; Tang, 1984: 125; Tang & Hao, 1995: 450 (Miller et al., 2013).
Nidularia spiraeae; Lindinger, 1957: 543. Change of combination.

Eriococcus spiraeae; Hoy, 1963: 117. Change of combination.
Acanthococcus spiraeae; Matesova, 1968: 114. Revived combination.
Eriococcus spiraeae; Tang & Hao, 1995: 450. Revived combination.
Acanthococcus spiraeae; Köhler, 1998: 384. Revived combination.
Eriococcus spiraeae; Miller & Gimpel, 2000: 340. Revived combination.
Acanthococcus spiraeae; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female body is egg-shaped, raspberry in color, 1.7 mm long, 1.1 mm wide. Sac is
grayish, compact, entirely covering the insect, 2.4 mm long, 1.4 mm wide (Borchsenius,
1949).
Mounted female
Adult female oval, 2.1–2.2 mm long, 1.1–1.3 mm wide. Antennae 7 segmented, length of
segments: I: 24.7, II: 25.3, III: 32, IV: 36.3, V: 24, VI: 17.3, VII: 32.7 µm long; segments
III and V without hair-like setae, other segments with a few hair-like setae; apical segment
with apical seta 43.7 µm long; apical segment also with 3 sensory falcate setae, longest
28.7 µm long; two preapical segments also with one sensory falcate seta; falcate setae
on segment V, 17 µm long; on segment VI, 27.3 µm long. Frontal lobe present; frontal
tubercle 3 µm wide. Eyes situated on venter near margin. Anterior spiracles 28 µm in
diameter. Anal lobes well developed, sclerotized, with 3 truncate conical enlarged setae,
43 µm long, ventrally with two flagellate setae, 55 and 20 µm long; anal lobe with 169 µm
long apical seta.
Venter: Labium 3 segmented, joint length of two apical segments 85.3 µm; basal segment
with 2 pairs of setae, apical segment with 5 pairs of hair-like setae, apical seta shorter and
thicker; stylet loop twice longer than labium. Legs well developed, lengths of segments
of prothoracic legs, coxa: 56, trochanter: 33.7, femur: 105.7, tibia: 79; tarsus: 95 and claw:
28.3, tarsal digitules 46, claw digitules 29.7 µm long; lengths of segments of mesothoracic
legs; coxa: 56, trochanter: 32, femur: 109, tibia: 82, tarsus: 107, claw: 29, tarsal digitules:
31, claw digitules: 30.3 µm long; lengths of segments of metathoracic legs; coxa: 60.3,
trochanter: 30.3, femur: 111.3, tibia: 93, tarsus: 110.7; claw: 29.7, tarsal digitules: 40.5,
claw digitules: 32 µm long, tarsal and claw digitules slightly knobbed, longer than claw;
meso- and metathoracic coxae with spinulae on anterior surface; claw with a denticle;
legs with a few hair-like setae; tibia with 4 setae. Quinquelocular pores 4 µm in diameter;
distributed in sparse rows across abdominal sternites, a few on thoracic segments and
head. Macrotubular ducts of two sizes: larger macrotubular ducts 7 µm wide and 35
µm long, situated on margin; smaller macrotubular ducts 3 µm wide and 17 µm long,
in a submarginal band on the thorax, scattered in the head and on abdominal segments.
Microtubular ducts sparse marginally, and submarginally on abdomen and head. A few
cruciform pores in submarginal area. A few hair-like setae scattered on submedian and
submarginal venter. Enlarged setae blunted 22–31 µm long on submargin. Suranal setae
hair-like, 96.7µm long.
Dorsum: The cuticular surface with nodulation. Enlarged setae in two sizes, longer setae
truncate, conical, 45–48 µm long, in a row on margins, two on each segments, and a
few medially on the head; shorter setae more numerous, about 22–31 µm long, forming
bands on segments. 14 enlarged setae on abdominal segment VII and 2 on abdominal
segment VIII. Macrotubular ducts 7 µm wide and 35 µm long scattered among spines.
Microtubular ducts scattered, 7–8 µm long, elongated. Anal ring situated on margin of
dorsum; oval, sclerotized, 53.3 µm wide, 73 µm long, with partly two rows of pores and
with 8 setae 108 µm long. Cauda present (after Borchsenius (1949), with modifications
Ecology
Host plant: Spiraea sp., S. crenata, S. hypericifolia, S. media, S. salicifolia, Betula fruticosa, B.
platyphylla, Duschekia fruticosa, Salix sp.
Distribution: China, Georgia, Kazakhstan, Russia.
Biology: Lives on branches of host plants.
Comments
The sizes given by Borchsenius (1949) are somewhat differs from material which we
examined, what could be the result of using different equipment.
Acanthococcidae 165
Figure 63. Acanthococcus spiraeae (Borchsenius, 1949), female. Original.

Acanthococcus thymi (Schrank, 1801) (Fig. 64) Combination nova

Coccus thymi Schrank, 1801: 146.
Type material: Germany (Bavaria, Kehlheim) (probably lost). Unknown type status,
type designation unknown. Notes: Lindinger (1933a) considered Eriococcus thymi
Schrank (1801) as different from E. thymi Signoret (1875). Williams (1985a) disagrees
in the following statement: “Although there is no original material available, there
seems to be no doubt that the specimens examined and identified by Signoret and
others identified by Marchal and Goux represent the species described by Schrank on
Thymus as C[occus] thymi and on Daphne gnidium as Rhizococcus gnidii.” Matile-Ferrero and
Danzig were unable to locate any type material of this species in Naturhistorisches
Museum, Vienna in June of 1997 and. Syntypes. Germany: Kehlheim Eriococcus thymi
(Schrank, 1801) Signoret, 1875: 32; Fernald, 1903: 79.
Additional material examined: Turkey: 1 ♀, Van-Çaldıran-Serpmetaş, on root
of Thymus sp., N: 39°11’143’’, E: 043°54’748’’, 2094 m, M.B. Kaydan, 20.vii.2005
(KPCT: 2215); 3 ♀♀, Van-Doğubayazıt road, on root of Artemisia vulgaris, N:
39°27’383’’, E: 044°14’982’’, 1694 m, M.B. Kaydan, 07.vi.2006 (KPCT: 2809); 3
♀♀on Van-Doğubayazıt road, on root of A. vulgaris, N: 39°27’026’’, E: 044°14’526’’,
1807 m, M.B. Kaydan, 07.vi.2006, (KPCT: 2832); 2 ♀♀, Iğdır-Aşağıaktaş, on root
of Compositae, N: 40°06’089’’, E: 043°30’858’’, 1197 m, M.B. Kaydan, 13. vi.
2007, (KPCT: 3580); 2 ♀♀, Van-Gürpınar-Çatak road, on root of Compositae, N:
38°03’868’’, E: 043°25’108’’, 1926 m, 29.vi.2007, M.B. Kaydan, (KPCT: 3838); 1 ♀,
Van-Çatak road, on root of Compositae, N: 38° 03’142’’, E: 043° 02’954’’, 1750
m, 03.vii.2007 (KPCT: 3935); 1 ♀, Hakkari-Yüksekova-Şemdinli road, on root of
Compositae, N: 38°03’142’’, E: 043°02’954’’, 1750 m, M.B. Kaydan, 05.vii.2007,
(KPCT: 3970); 2 ♀♀, Hakkari- road, on root of Thymus sp., N: 37°41’873’’, E:
043°52’296’’, 1538 m, M.B. Kaydan, 04.vii.2007, (KPCT: 3973). France: Montpellier,
on Thymus vulgaris (Lamiacea) (det. as E. thymi by V. Signoret), Cannes, on Daphne
gnidium (Thymelaeaceae) (det. As R. gnidii by V. Signoret), Bouc-Bel-Air, on T. vulgaris,
v.1932 (L. Goux).
Lit.: Ali, 1970: 76; Borchsenius, 1948: 501; Cockerell, 1896: 324; Danzig, 1975: 42;
Dziedzicka & Koteja, 1971: 558; Fernald, 1903: 79; Ferris, 1955: 94; 1957b: 85; Foldi,
2000: 81; 2001:305; 2002: 246; Gómez-Menor Ortega, 1937: 346; 1948: 105; 1954:
144; 1957: 79; 1958: 9; 1968: 553; 1960: 201; Goux, 1931: 73; 1936: 353; 1936a: 294;
1938: 328; 1944: 137; 1946: 90; 1948: 67; Hoy, 1963: 92; Kaydan, et al., 2007: 90;
Kaydan, et al., 2002: 255; Kiritchenko, 1940: 127; Koteja, 1974a: 276, 1974b: 296;
1986: 28; Koteja & Zak-Ogaza, 1981: 501; Kozár, 1986: 172; Kozár et al., 2013: 55;
Kozár & Walter, 1985: 74; Lindinger, 1912: 133; Lindinger, 1933b: 43; Marchal, 1909:
198; MacGillivray, 1921: 145; Martin-Mateo, 1985: 93; Pellizzari & Kozár, 2011: 66;
Quvrard & Kozár, 2009: 102; Signoret, 1875: 32; Stoetzel & Miller, 1979: 16; Tang
& Hao, 1995: 519; Williams, 1985a: 382; Williams & Ben-Dov, 2009: 45; Yang, 1982:
105 (Miller et al., 2013).
Acanthococcidae 167
Eriococcus thymi; Signoret, 1875: 32. Change of combination.

Eriococcus thymi; Williams, 1985a: 382. Revived combination.
Nidularia thymi; Lindinger, 1957: 548. Change of combination.
Acanthococcus thymi; Kozár & Walter, 1985: 74. Change of combination.
Rhizococcus thymi; Tang & Hao, 1995: 541. Change of combination.
Acanthococcus thymi; Köhler, 1998: 384. Revived combination.
Eriococcus thymi; Miller & Gimpel, 2000: 360. Revived combination.
Acanthococcus thymi; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Body broadly oval, nodulose, largest specimens 1.9–2.5 mm long, 1.35–1.45 mm wide.
Mounted female
Antennae with 6 (rearly 7) segments, length of segments: I: 30, II: 20, III: 70, IV: 20,
V: 20, VI: 36 µm long; the third segment often divided, without hair-like setae, other
segments with a few hair-like setae; two preapical segments with 1 sensory falcate seta, 17
µm long, sixth 27.3 µm long, apical segment with 3 sensory falcate setae, 28.7–43.7 µm
long. Eyes situated on venter near margin. Frontal tubercle present, seta-like, just anterior
to basal antennal segment. Anal lobes well developed, sclerotized, with 3 conical spines,
34–50 µm long, ventrally with two flagellate setae, 50 µm long, anal lobe with 280 µm
long apical seta.
Venter: Labium 130–160 µm long, slightly shorter than clypeolabral shield, basal segment
with 2 pairs of setae, apical segment with 5 pairs of hair-like setae, apical seta shorter and
thicker. Stylet loop twice longer than labium. Legs well developed; lengths of segments
of prothoracic legs; coxa: 41, trochanter: 40, femur: 110, tibia: 79; tarsus: 100 and claw:
22 µm long; lengths of segments of mesothoracic legs; coxa: 58, trochanter: 38, femur:
115, tibia: 88, tarsus: 94, claw: 36 µm long, lengths of segments of metathoracic legs;
coxae: 56, trochanter: 38, femur: 120–125, tibia: 90, tarsus: 105; claw: 33µm long, tarsal
digitules: 40 µm long, tarsal and claw digitules slightly knobbed, longer than claw; hind
coxa with a few translucent pores and spinulae on anterior surface; claw with a denticle;
legs with a few hair-like setae, and with one sensory pore on tarsus; tibia with 4 setae.
Spiracles heavily sclerotized, the sclerotisation continuing around the atrium in varying
degrees. Anterior spiracles 28 µm in diameter. Quinquelocular pores 4 µm in diameter;
distributed in sparse bands across abdominal sternites, a few on thoracic segments and
head region. Macrotubular ducts of three sizes: larger macrotubular ducts 10 µm wide
and 22 µm long, situated on margin; smaller macrotubular ducts in a submarginal band on
the thorax, scattered in the head region and on abdominal segments. Microtubular ducts
sparse marginally, and submarginally on abdomen and head. A few cruciform pores in
submarginal area. A few hair-like setae scattered on submedian and submarginal venter,
about 40 µm long. Enlarged setae conical, forming a row on margin. Cruciform pores
few, in submarginal areas of anterior abdominal segments and thorax, and in median
areas of thorax and head.
Dorsum: Enlarged setae in two sizes, setae conical, 28–60 µm long, in a row on margins,
two on segments, and a few medially on the head; shorter setae more numerous, about
12–28, on thorax setae 12–40, and on abdomen 10–20 µm long, forming rows on
segments. 8 dorsal setae on abdominal segment VII and absent abdominal segment on
VIII. Macrotubular ducts one size, 7 µm wide and 25 µm long, scattered among spines.
Microtubular ducts with inner sclerotised ampulla, sclerotised tube and collar and with
bifid orifice, scattered, 5–8 µm long, elongated. Anal ring situated on margin of dorsum;
53 µm wide, 72 µm long, oval, sclerotized, with 8 setae, each 120–130 µm long. Suranal
seta much shorter than anal ring setae. Cauda visible, membranous, rounded, narrow,
margin irregular, nodulose (after Williams (1985a), with modifications based on slides
from Goux collection).
Other stages
Male described briefly by Goux (1936).
Ecology
Host plant: Anthusa sp., Artemisia herba-alba, A. nana, Calluna sp., Centaurea solstitialis,
Daphne gnidium, Erica arborea, E. carnea, E. scoparia, E. tetralix, Salicornia sp., Santolina
rosmarinifolia, Suaeda vera, Teucrium sp., Thymelaea villosa, Thymus mastichina, T. vulgaris.
Distribution: France, Germany, Hungary, Israel, Italy, Portugal, Spain, Switzerland,
Turkey.
Biology: On the roots, twigs, and leaves of its host plants.
Comments:
According to Williams (1985a) there is no original material available, there seems to be
no doubt that the specimens examined and identified by Signoret and others identified by
Marchall and Goux represent the species described by Schrank on Thymus as C. thymi and
on Daphne gnidium as R. gnidii. The characters on some specimens of D. gnidium tend to
be larger but the specimens themselves are larger and represent the same species. In well-
stained specimens the microducts clearly show the bifid orifice, a feature difficult to see
in some specimens. The illustration is based on specimens from Thymus vulgaris, identified
as E. thymi by Signoret. Further collectings may show a much wider host plant range.
Acanthococcidae 169
Figure 64. Acanthococcus thymi (Schrank,1801), female. After Williams (1985) with
modifications.
Acanthococcus tokaedae (Kuwana, 1932) (Fig. 65)

Eriococcus tokaedae Kuwana, 1932: 146.
Syntypes: Female. Japan (Honshu, Tokyo City), on Acer trifidum, 13.v.1932, by K.

Muramatsu. Deposited in ITLJ.
Common name: Acer scale.
Lit.: Hoy, 1963: 121; Hua, 2000: 137; Kawai, 1972: 4; 1980: 129; Kwon & Han, 2003:
151; Köhler, 1998: 385; Paik, 1978: 167; Takahashi, 1957: 7; Tang, 1984; Tang & Hao,
1995: 450; Wang, 1980: 115; 1982b: 143; 1982: 41; 2001: 208 (Miller et al., 2013).
Acanthococcus tokaedae; Kozár & Walter, 1985: 74. Change of combination.

Eriococcus tokaedae; Miller & Gimpel, 2000: 363. Revived combination.
Acanthococcus tokaedae; Kozár, 2009: 94. Revived combination.
Description
Unmounted female
Ovisac 3 mm long, 2 mm wide, the female somewhat smaller, grayish white, nearly oval.
Female is oval. Male sac is similar to female, but smaller (Kuwana, 1932).
Mounted female
Antennae 7 segmented, second and third segments subequal, and the longest, all
segments except third with few rather long spine-like setae, fifth and sixth segment with
one falcate sensory setae, and the seventh with three. Anal lobes protruding, apically
rounded, moderately sclerotized, serrate on inner margin, with 3 dorsal equal long conical
setae and with three short hair-like setae and a long apical setae.
Venter: Labium small, well developed, 3 segmented; stylet loop rather long, much longer
than labium. Legs subequal, comparatively small, tarsus longer than tibia, tarsal digitules
longer than claw, claw with minute digitules. Quinquelocular pores distributed in sparse
bands across abdominal segment V to VIII, and on thoracic segments. Macrotubular
ducts of two sizes: larger macrotubular ducts, situated on margin; smaller macrotubular
ducts in a submarginal band on the thorax, scattered in the head region and on abdominal
segments. With flagellate setae in median areas, short and slender but stouter and stiff in
lateral areas, with rather large pores especially in abdominal region.
Dorsum: Enlarged setae cylindrical, sides straight, apices rounded of 2 or 3 sizes, those
on the margin somewhat larger, some setae slightly curved, with minute pointed tubercles
between spines; no enlarged setae on abdominal segment VIII, anterior of anal ring;
largest setae form sparse, transverse segmental bands; number of spines on abdominal
segment VII 24, on the abdominal margin 3-4 setae present. Macrotubular ducts of one
size: scattered through. Anal ring oval, sclerotized, with 8 setae. Cauda not mentioned
(after Kawai (1980); Kuwana (1907); Tang & Hao (1995), with modifications).
Acanthococcidae 171
Figure 65. Acanthococcus tokaedae (Kuwana, 1932), female. After Wu (1982) with
modifications.
Ecology
Host plant: Acer trifidum.
Biology: On twigs and stems.
Acanthococcus transversus (Green, 1922) (Fig. 66)

Eriococcus transversus Green, 1922: 351.
Syntypes: Female. Sri Lanka (Maskeliya), on Arundinaria sp.?, ?/08/1902, by E. E. Green.

Deposited in BMNH. Notes: There are three adult female syntypes on 1 slide in the
BMNH (Williams, personal communication, May 15, 1996).
Lit.: Ali, 1970: 77; Fang et al., 2001: 104; Hoy, 1963: 121; Hua, 2000: 137; Köhler, 1998:
385; Tang & Hao 1995: 451, 648; Wang, 1982b: 143; 2001: 208; Yang, 1982: 105 (Miller
et al., 2013).
Acanthococcus transversus; Kozár & Walter, 1985: 74. Change of combination.

Rhizococcus transversus; Xu & Wu, 1989: 117-119. Change of combination.
Eriococcus transversus; Miller & Gimpel, 2000: 365. Revived combination.
Acanthococcus transversus; Kozár, 2009: 94. Revived combination.
Description
Unmounted female
Female sac is ochreous white, laterally compressed and curved into the shape of a
horseshoe. Adult female is olivaceous brown, darker beneath, also curved into a loop.
Ovisac 2.5 mm long, 1.25 mm wide (Green, 1922).
Mounted female
Antennae 7 segmented, second and third segments subequal, and the longest, all
segments except third with few rather long hair-like setae, fifth and sixth segment with
one falcate sensory setae, and the seventh with three. Anal lobes protruding, apically
rounded, moderately sclerotized, with 3 dorsal equally long conical setae and with three
short hair-like setae and a long apical setae.
Venter: Labium small, well developed, 3 segmented; stylet loop rather long, much longer
than labium, reaching median coxae. Legs well developed, tarsus markedly longer than
tibia, tarsal and claw digitules longer than claw, claw with minute denticle. Quinquelocular
pores distributed in sparse bands across abdominal segment V to VIII, and on thoracic
segments. Cruciform pores forming a submarginal band. Macrotubular ducts of two
sizes: larger macrotubular ducts, situated on margin; smaller macrotubular ducts in a
submarginal band on thorax, scattered in the head region and on abdominal segments.
With normal flagellate setae in median areas, short and slender but stouter and stiff in
Acanthococcidae 173
Figure 66. Acanthococcus transversus (Green, 1922), female. Original.

lateral areas, with rather large pores especially in abdominal region.

Dorsum: Slide-mounted adult female with: enlarged conical setae, sides straight, apices
acute, setae of 2 sizes, larger size present around body margin, smaller size abundant
over remainder of dorsum, with two enlarged setae on segment 8 anterior of anal ring;
largest setae form sparse, transverse segmental bands; number of spines 20 on abdominal
segment VII, on the abdominal margin 4 setae present. Macrotubular ducts of one size:
scattered throughout. Anal ring oval, sclerotized, with 8 setae. Cauda well developed
(Green, 1922).
Ecology
Host plant: Arundinaria debilis?, Arundinaria sp., Bambusa dissemulator var. hispida, B. textilis,
B. vulgaris.
Distribution: China, China (Taiwan), India, Sri Lanka.
Biology: Across the axils of leaves, in the angles of branches.
Acanthococcus ulmarius Danzig, 1975 (Fig. 67)

Acanthococcus ulmarius Danzig, 1975: 66.
Holotype: Female. Russia (Southern Primor'ye, Komarovo-zapovednoye), on Ulmus

davidiana var. japonica, 16.vii.1969, by E. Danzig. By original designation. Deposited
in ZMAS.
Lit.: Danzig; 1975: 43; 1977b: 57; 1980: 205; 1986: 240; 1988: 708; Koteja & Zak-
Ogaza, 1981: 514; Kozár & Walter, 1985: 74; Kwon & Han, 2003: 151; Tang & Hao,
1995: 449; Tao, 1999: 33; Wang, 2001: 208 (Miller et al., 2013).
Eriococcus ulmarius; Tang & Hao, 1995: 499. Change of combination.

Acanthococcus ulmarius; Köhler, 1998: 386. Revived combination.
Eriococcus ulmarius; Miller & Gimpel, 2000: 370. Revived combination.
Acanthococcus ulmarius; Kozár, 2009: 94. Revived combination.
Description
Unmounted female
Adult female is oval, violet, up to 2.5 mm long. Ovisac is oval, convex, dirty white,
smooth. Eggs are pink.
Mounted female
Antennae 7 segmented. Frontal lobe well developed. Anal lobes protruding, nodulose
apically acute, lobes dorsally with 3 enlarged setae.
Venter: Apical labial segment with 5 pairs of long setae, the median setae much shorter
than others. Legs medium size; coxae with spinulae on anterior surface, each tibia with 4
Acanthococcidae 175
Figure 67. Acanthococcus ulmarius Danzig, 1975, female. After Tang & Hao (1995) with
modifications.
strong setae, except anterior legs, where median setae present; tarsus longer than tibiae,
claws with denticle near tip; tarsal and claw digitules longer than claw, slightly knobbed.
Quinquelocular pores numerous, present over entire venter. Macrotubular ducts of two
sizes, larger ones on margin and submarginal and scattered on head, narrower ones on
medial abdominal segments. Microtubular ducts absent. Hair-like setae short. Enlarged
setae of small size only, present along body margin. Cruciform pores present along lateral
margin.
Dorsum: Enlarged setae cylindrical, elongate, sides straight except concave basally,
apices rounded, large size form longitudinal band medially, setae abundant over surface;
enlarged setae form wide bands including big groups in middorsum of abdomen and
thorax; number of enlarged setae smaller on the head part. Macrotubular ducts present
in small numbers all over dorsum. Microtubular ducts numerous, narrow, long. Anal
ring, with 10 setae, with one row of pores. Cauda present (after Danzig (1975), with
modifications based on a paratype).
Ecology
Host plant: Ulmus davidiana, U. davidiana var. japonica, U. pumila.
Distribution: China, Mongolia, North Korea, Russia.
Biology: Eggs are laid in mid June (Danzig, 1975).
Acanthococcus uvaeursi (Linnaeus, 1761) (Fig. 68)

Coccus uvae-ursi Linnaeus, 1761: 266.
Holotype: Deposited in LSUK.

Common name: Linnaeus' Felt Scale.
Lit.: Borchsenius, 1938: 135; 1949: 45; 1950: 121; 1963: 214; 1973: 215; Danzig, 1959:
443; 1964: 632; 1971a: 821; 1975: 43; 1977b: 53; 1978a: 13; 1978a: 77; 1980: 215; 1986:
240; 1988: 708; 1994: 47; Foldi, 2000: 81; 2001: 305; Gertsson, 2001: 125; 2008: 56;
Goux, 1936: 299; 1948: 69; Kosztarab & Kozár, 1988: 277; Kozár & Walter, 1985: 74;
Lindinger, 1912: 74; Ossiannilsson, 1951: 3; Ouvrard & Kozár, 2009: 102; Pellizzari &
Kozár, 2011: 66; Tao, 1999: 33; Tereznikova, 1959a: 683; 1959b: 796; 1959c: 92; 1963:
187; 1963a: 47; 1963b: 1527; 1966: 25; 1966: 963; 1967: 475; 1975: 28; 1981: 40; Wang,
2001: 209; Williams, 2007: 42; Williams & Ben-Dov, 2009: 9; Williams & Gertsson,
2005: 3419 (Miller et al., 2013).
Coccus arbuti; Fabricius, 1775: 744. Unjustified replacement name for Coccus uvae-ursi
Linnaeus, 1767: 742.
Coccus arbuti; Turton, 1802: 714. Turton gave a description thus validating the name as C.
arbuti Turton which is a primary homonym of C. arbuti Fabricius and a junior synonym
of C. uvae-ursi Linnaeus (Williams and Ben-Dov, 2009).
Eriococcus bezzii; Lindinger, 1912: 367. Incorrect synonymy. Lindinger (1912) incorrectly
Acanthococcidae 177
considered E. bezzii to be a junior synonym of E. uvaeursi.

Eriococcus uvae-ursi; Lindinger, 1912: 74. Change of combination.
Nidularia uvae-ursi; Lindinger, 1933a: 117. Change of combination.
Eriococcus bahiae; Goux, 1948: 69. Incorrect synonymy.
Acanthococcus uvae-ursi; Borchsenius, 1949: 349. Change of combination.
Acanthococcus uvaeursi; Borchsenius, 1963: 215. Justified emendation.
Eriococcus uvaeursi; Hoy, 1963: 123. Change of combination.
Acanthococcus uvaeursi; Kosztarab & Kozár, 1978: 76. Revived combination.
Eriococcus uvaeursi; Tang & Hao, 1995: 452; Revived combination.
Acanthococcus uvaeursi; Köhler, 1998: 386; Revived combination.
Eriococcus uvaeursi; Miller & Gimpel, 2000: 371; Revived combination.
Acanthococcus uvaeursi; Kozár, 2009: 94. Revived combination.
Description
Unmounted female
Adult female covered in a greyish-white felted sac, often 3 mm long, almost covering
entire insect.
Mounted female
Adult female almost elliptical, sometimes sides sub-parallel, 1.25–1.80 mm long, 0.65–
1.00 mm. wide. Antennae with 7 segmented, 260–290 µm long. Eyes situated on venter
near margin. Frontal lobe present, just anterior to basal antennal segment. Anal lobes well
developed, sclerotized on dorsum and venter, sclerotised on venter and also laterally, with
sclerotization extending antero-medially; each lobe terminating in a long, flagellate apical
seta 150–170 µm long, also with 3 enlarged setae on dorsum and a flagellate seta about
52 µm long on venter.
Venter: Labium 120–149 µm long, shorter than clypeolabral shield. Legs well developed,
hind trochanter+femur 160–180 µm long, hind tibia+tarsus 200–220 µm long. Claw
curved about 25 µm long, with minute denticle towards distal end. Ratio of lengths of
hind tibia+tarsus to hind trochanter + femur 1.15–1.29. Ratio of lengths of hind tibia to
tarsus 0.8–0.9. Hind tarsus with 4 flagellate setae, tarsus of each foreleg with 3 flagellate
setae. Distal trochanteral seta about 70 µm long. Hind coxa without translucent pores.
Quinquelocular pores present about 5 µm in diameter, fairly numerous across abdominal
segments, medial areas of head and thorax and near each spiracle, extending to lateral
margins from each anterior spiracle. Macrotubular ducts of 3 main sizes; a large type, 30
µm long, cup about 11 µm wide present around margins; a smaller type, similar to those
on posterior dorsal segments, present mainly across middle of abdominal segments,
scattered on head and thorax; a minute type of duct, 10–16 µm long, with cup about
5 µm wide, present across abdominal segments only. Microtubular ducts same as on
dorsum, sparse, a few situated on abdominal margins. Ventral surface with short flagellate
setae, except near middle of venter where each seta thicker, mostly about 50 µm long.
Marginal enlarged setae in band about 25 µm long. Cruciform pores, about 4 µm long,
present submarginally.
Dorsum: Marginal enlarged setae, mostly pointed with rounded sides, present on margins
forming a single row, normally 2 present on margin of each abdominal segment VII,
50–55 µm long, other marginal setae 35–50 µm long. Enlarged setae distributed across
posterior abdominal segments, each seta usually about 12.5–15.0 µm long, others across
anterior segments, each about 35–45 µm long except some on mid-thorax where each
about 50 µm long. Macrotubular ducts normally of 2 sizes. A large type, 30 µm long,
cup about 11 µm wide, fairly abundant across segments except on posterior abdominal
segments with smaller ducts about 20 µm long and cup about 7.5 µm wide. Microtubular
ducts slender, 10 µm long, sclerotised except near outer end, scattered through. Anal ring
about 60 µm long, sometimes folded as base of anal lobes, normally ventral in position,
with a single row of cells except double towards posterior end, bearing also 6 setae each
110–115 µm long (Williams, 2007).
Other stages
First instar described by Leonardi (1920) under name of A. bezzii.
Ecology
Host plant: Artemisia sp., Arbutus sp., Arctostaphylos uva-ursi, Azalea sp., A. indica, Erica sp.,
Eriophyllum sp., Eubotryoides sp., E. grayana, Rhododendron sp., R. tolmachevii, Vaccinium hirtum,
V. myrtillus, V. praestans, V. uliginosum, V. vitis-idaea, Crithmum sp., C. maritimum.
Distribution: Austria, China, France, Germany, Italy, Russia, Sweden, Switzerland,
Ukraine.
Biology: On the twigs, often found on soil-covered stem parts of host, Schmutterer
(1955) considered to be a rare montane species, has been observed at elevations of 2.600
m (Kosztarab & Kozár, 1988). This species could have two generations per year (Danzig,
1986). Males unknown.
Acanthococcidae 179
Figure 68. Acanthococcus uvaeursi (Linnaeus, 1761), female. After Williams (1985) with
modifications.
Genus:
Anophococcus Balachowsky, 1954
Type sp.: Eriococcus inermis (Green, 1915), by original designation.
Lit.: Boratynski, 1962: 55; Borchsenius, 1948: 502; 1949: 22; Danzig, 1962a: 860;
1975: 42; 1980: 205; Ferris, 1957b: 85; Gómez-Menor Ortega, 1937: 322; Green,
1922: 20; Hodgson & Miller, 2010: 45; Hoy, 1962: 510; 1963: 132; Kawecki, 1985: 27;
Kosztarab & Kozár, 1978: 76; 1988: 286; Koteja, 1974a: 248; 1974b: 77; Koteja &
Zak-Ogaza, 1981: 501; Kozár & Konczné Benedicty, 2008a: 142, 2008b: 247; Kozár
& Walter, 1985: 73; Leonardi, 1920: 426; Miller & Gimpel, 1996: 605; Morrison &
Morrison, 1966: 10; Newstead, 1903: 195; Ouvrard & Kozár, 2009: 101; Tang &
Hao, 1995: 448; Tereznikova, 1981: 13; Williams, 1969b: 325, 1985a: 347 (Miller et
al., 2013).
Eriococcus; Synonymy by Ferris, 1955: 94. Notes: Synonymy of this genus was a subjective
decision (Miller & Gimpel, 2000: 106). Some have treated it as valid (Kosztarab & Kozár,
1988; Koteja & Zak-Ogaza, 1981), while others have treated it as a junior subjective
synonym of Eriococcus (Hoy, 1963; Williams, 1985a).
Anophococcus; Koteja & Zak-Ogaza, 1981. Revived combination.
Erioococcus; (Miller & Gimpel, 2000: 106). Revived combination.
Anophococcus; Kozár, 2009: 111. Revived combination.
Description
Ovisac ovoid, felt-like, white or grey completely enclosing female body. Adult female
elongate-oval, narrowed posteriorly, with anal lobes conical, slightly sclerotised, antennae
6 usually 7 segmented; frontal lobes usually absent, frontal tubercles present, labium 3
segmented, with 18 setae, of these 12 on apical segment, apical setae of labium about
half length, or less of the subapical setae, with well developed segments and a weakly
developed basal segment with two pairs of hair-like setae, the outer one about half
length of the inner ones. Stylets usually short, as long as labium; legs well-developed,
midcoxae and hind coxae often with spinulae on anterior surfaces, translucent sensory
pores generally present or absent, inner side of hind tibia with 5 setae, claw usually with
a denticle; tarsal and claw digitules mostly longer than claw, knobbed, in some cases claw
digitules shorter than claw, spine-like. Spiracles often with a few associated discodial pores;
discodial pores on venter only, usually quinquelocular, but the number of loculi vary
between 3 to 9, generally double rows of pores; cruciform pores absent from dorsum,
but often on prosomal venter in a marginal band; not sclerotized, not grouped; tubular
ducts of 2 types: micro- and macrotubular ducts; microtubular ducts short, 3–4 um long
and 2 um wide, scattered or form transverse rows or bands on dorsum, often among
dorsal conical setae; macrotubular ducts often of 2 or 3 sizes, usually form transverse
rows or bands on body surfaces; larger ducts with inner ductile shorter than main ductile
Acanthococcidae 181
and with a flower-shaped terminal gland; smaller ducts with short filamentous ductile.
Enlarged conical setae normally present only on margin, sometimes only on anal lobes
and on margin of head, dorsal setae minute needle-like. Hair-like setae on venter only;
anal ring well developed, sclerotized with one or partly double row of pores and with 8,
rarely 6, anal ring setae, latter often as long as apical seta on anal lobes; each anal lobe
with a long apical seta and usually with 3 short dorsal conical setae, seldom with 4, ventral
hair-like setae, suranal setae hair-like, Cauda usually absent, or weakly developed.
In a comparative table (Fig. 69) we studied differences of enlarged setae variations in
females and first instar nymph of some species belonging to genus Anophococcus. The type
of enlarged setae which represent the unity of these species in one genus quite similar.
an exception An. parvisetus differs from others by absence of marginal setae in both
immature and adult stages. In this genus the number of spines, size and shape of studied
species quite similar and for the identification of species other characters should also be
needed.
Figure 69. Anophococcus setae range on adult female and first instar.
Distribution
Kozár (2009) cited only four species from the Palaearctic Region in his World list. Here
we transferred several of the grass living species which were regarded in genus Rhizococcus
sensu lato. Because of the instability of the system, several species from the Nearctic
Region might belong to this genus. Similarly species in the genus Eriococcus sensu lato
from other regions may also belong to this genus. The composition of species in several
regions needs to be revised. In the Palearctic Region, we recognize 24 species in total. It
seems that the genus has a wide distribution in the Palaearctic and probably some live in
the Nearctic Region, considered under a different genus name.
Biology
They feed mostly on upper side of leaves of different grasses, especially on Agropyron
and Cynodon. Some species prefer different herbaceous plants, but on the other hand,
we assume that some host plant records could be erroneous, because the females when
egglaying often choose dry plants, like A. confusus on Pinus needles (Danzig, 1962a).
Generally found in large colonies. Most species have one yearly generation; usually
overwinters in egg, or second nymphal stage; males normally present.
Comments
Here we follow mostly the concept of Koteja & Zak-Ogaza (1981) and Koteja (1990).
Key to species
1. – Body margin without enlarged setae................................................................................2

– Body margin with at least one enlarged setae on each segment..................................4
2. – Anal lobe without enlarged setae..................................................................... An. salsolae
– Anal lobe with enlarged setae............................................................................................3
3. – Antennae 6 segmented, without frontal lobe................................................. An. inermis
– Antennae 7 segmented, with frontal lobe................................................ An. parvispinus
4. – Margin of abdominal segments with more than five enlarged setae..........................5
– Margin of abdominal segments with less than five enlarged setae.............................6
5. – Enlarged marginal setae blunted, without submarginal enlarged setae on venter,
some of ventral setae slightly blunted................................................................An. abaii
– Enlarged marginal setae sharp ended, with submarginal enlarged setae on venter,
none of ventral setae slightly blunted.....................................................An. oxyacanthus
6. – Only one enlarged setae on each margin of body segments..................An. pannonicus
– Two or more enlarged setae on margin of abdominal segments................................7
Acanthococcidae 183
7. – Number of enlarged setae on margin of abdominal segment

VII two, with frontal lobe ................................................................................................8
– Number of enlarged setae on margin of abdominal segment
VII more than two, without frontal lobe........................................................................9
8. – Marginal enlarged setae long and blunted, dorsal setae thin and
sharp ended.....................................................................................................An. cingulatus
– Marginal enlarged setae short and stout, dorsal setae wide and
blunted................................................................................................................An. confusus
9. – On dorsum of abdominal segment VIII four enlarged setae
present................................................................................................................An. lerzanae
– On dorsum of abdominal segment VIII enlarged setae absent................................10
10 – The number of marginal enlarged setae on seventh abdominal segment three.....11
– The number of marginal enlarged setae on seventh abdominal segment four.......18
11. – Marginal enlarged setae sharp pointed...........................................................................12
– Marginal enlarged setae blunted or cut ended..............................................................14
12. – With 6 segmented antennae, smaller macrotubular duct on venter not in
submarginal longidual rows, rare.............................................................An. kotejai sp.n.
– With 7 segmented antennae, smaller macrotubular duct on venter in
submarginal longidual rows with high number............................................................13
13. – Enlarged setae on dorsum restricted on thorax and head, dorsal setae on thorax
and head as long as marginal enlarged setae........................................... An. kondarensis
– Enlarged setae on dorsum all over dorsum, no of setae on dorsum as long as
marginal enlarged setae.............................................................................An. pseudinsignis
14. – Marginal enlarged setae blunted......................................................................................15
– Marginal enlarged setae cut ended..................................................................................17
15. – With 6 segmented antennae, dorsal derm granulated...............................An. granulatus
– With 7 segmented antennae, dorsal derm not granulated..........................................16
16. – Each abdominal segment with two enlarged setae (except abdominal segment
VII), multilocular pores on venter with five or seven loculars.................. An. agropyri
– Each abdominal segment with three enlarged setae, multilocular pores on venter
with fiveloculars.................................................................................................. An. iljiniae
17. – The size of enlarged setae on dorsum almost half of the marginal enlarged setae
and curved....................................................................................................... An. herbaceus
– The size of enlarged setae on dorsum small, almost one third of the marginal
enlarged setae and curved.....................................................................An. sanguinairensis
18. – Venter without cruciform pores.......................................................................An. socialis
– Venter with cruciform pores...........................................................................................19
19. – With 6 segmented antennae............................................................................................20
– With 7 segmented antennae............................................................................................22
20. – With longudial submarginal enlarged setae row on venter..................... An. adzharicus
– Without longudial submarginal setae row on venter...................................................21
21. – Marginal setae narrow, 3-4 times longer than wide.................................. An. formicicola
– Marginal setae wide, strong, two times longer than wide..............................An. selmae
22. – With cruciform pores on middle part of head on venter, claw
without denticle.................................................................................................An. oblongus
– Without cruciform pores on middle part of head on venter, claw
with denticle......................................................................................................................23
23. – With a group of enlarged setae on head on dorsum, without quinquelocular
pores on middle part of head on venter........................................................An. evelinae
– Without a group of enlarged setae on head, with quinquelocular pores
on middle part of head..................................................................................... An. insignis
Anophococcus abaii (Danzig, 1990) (Fig. 70) Combination nova

Acanthococcus abaii Danzig, 1990: 373.
Holotype: Female. Iran (143 km NW of Tehran), on Haloxylon sp., 25.ix.1986, by M.

Abai. By original designation. Deposited in ZMAS.
Lit.: Germain, 2008: 77; Kozár, et al., 1996: 64; Miller & Gimpel, 1999: 213; Miller &
Gimpel, 2000: 111; Tang & Hao, 1995: 520 (Miller et al., 2013).
Rhizococcus abaii; Tang & Hao, 1995: 521. Change of combination.

Eriococcus abaii; Miller & Gimpel, 1999: 213. Change of combination.
Rhizococcus abaii; Kozár, 2009: 105. Revived combination.
Description
Unmounted female
Adult female is up to 6 mm long.
Mounted female
Antennae 7 segmented. Frontal lobe and frontal tubercle well developed. Anal lobe
slightly sclerotised, with four enlarged setae, three of which on inner side.
Venter: Apical labial segment with 6 pairs of short setae, the median setae almost as
long as the others. Stylet loop twice longer than labium. Legs medium sized; posterior
coxae with large pores and spinulae on anterior surface, some pores on femur, tibia with
4 strong setae, tarsus shorter than tibiae, claws with denticle near tip. Tarsal and claw
digitules longer than claw, slightly knobbed. Quinquelocular pores and septalocular pores
with double ring, numerous present over entire venter. Macrotubular ducts of two sizes,
scattered all over the venter, form a band around margin. Microtubular ducts present
only on body margin. Cruciform pores absent. Hair-like setae short. Enlarged setae of
small size only, present along body margin.
Acanthococcidae 185
Figure 70. Anophococcus abaii (Danzig, 1990), female. After Danzig (1990) with
modifications.
Dorsum: Slide-mounted adult female with: 7-10 enlarged setae on margin of most
abdominal segments; marginal setae much longer than dorsal setae. Macrotubular ducts
numerous all over dorsum. Microtubular ducts numerous, narrow, medium size. Anal
ring dorsal, with 4 pairs of short setae with one row of pores. Cauda present (after
Danzig (1990), with modifications based on paratypes).
Ecology
Host plant: Haloxylon sp.
Distribution: Iran.
Biology: Unknown.
Anophococcus adzharicus (Hadzibejli, 1960) (Fig. 71) Redescription,

Combination nova
Acanthococcus adzharicus Hadzibejli, 1960: 306.
Holotype: Female. Georgia (Adzharia, Zelenyi Mys 18.ix.1953), on Gramineae, by

Z. Hadzibejli. By original designation. Deposited in GPPT. Notes: 4 paratypes on 1
slide with same data in ZMAS
Lit.: Danzig, 1975: 54; Hadzibejli, 1960: 269; Kozár & Walter, 1985: 73; Köhler, 1998:
372; Miller & Gimpel, 1999: 213; Miller & Gimpel, 2000: 119 (Miller et al., 2013).
Eriococcus adzharicus; Miller & Gimpel, 1999: 213. Change of combination.

Acanthococcus? adzharicus; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female is oval, cream-coloured, latter becomes to violet, up to 2.0 mm long. Ovisac
is oval, convex, dirty white, smooth, up to 3.5 mm. Eggs are pink.
Mounted female
Adult female oval, 1.5–1.8 mm long, 1 mm wide. Antennae 6 rarely 7 segmented, length
of segment: I: 34.5, II: 29.5, III: 74, IV: 25, V: 24 and VI: 34.5 µm long; segments with a
few hair-like setae; apical seta on apical segment 37.5 µm long; apical segment also with
3 sensory falcate setae, longest 26 µm long; two preapical segments also with 1 sensory
falcate seta: on forth segment 15.5, fifth segment 29.5 µm long. Frontal lobe absent,
frontal tubercle 3 µm wide and 2 µm high. Eyes situated on venter near margin. Diameter
of anterior spiracles 27 µm. Anal lobes well developed, sclerotized, with 3 slightly blunt
conical spines, 33.6–38.2 µm long, ventrally with two flagellate setae, the longest is 33.6
µm long; anal lobe with 202.5 apical seta.
Acanthococcidae 187
Figure 71. Anophococcus adzharicus (Hadzibejli, 1960), female. Oriiginal.

Venter: Labium 3 segmented, joint length of two apical segments 67 µm; basal segment
with 2 pairs of setae, apical segment with 6 pairs of hair-like setae, the apical seta
shorter; stylet loop short, as long as labium. Legs well developed; length of segments
of prothoracic legs; coxa: 58, trochanter: 32.5, femur: 125, tibia: 101.5, tarsus: 102.5 and
claw: 27 µm long, tarsal digitules: 45.5, claw digitules: 31 µm long; length of segments
of mesothoracic legs; coxa: 55, trochanter: 40.5, femur: 126.5, tibia: 106.5, tarsus: 113.5
and claw: 28 µm long, tarsal digitules: 47, claw digitules: 32 µm long; length of segments
of metathoracic legs; coxae: 70, trochanter: 41.5, femur: 142, tibia: 119, tarsus: 121.5
and claw: 29.5 m long, tarsal digitules: 47.5, claw digitules: 33.5 µm long, tarsal and
claw digitules slightly knobbed, longer than claw; metathoracic coxae with spinulae on
anterior surface; metathoracic coxae and femur also with small pores on both anterior
and posterior surfaces; claw with a denticle; legs with a few hair-like setae, and tibia with 5
setae. Multilocular pores with 5-7 loculi, mostly 7; quinquelocular pores 4 µm in diameter,
septalocular pores 4–5 µm in diameter and distributed in sparse rows on the whole venter.
Macrotubular ducts of two sizes: larger macrotubular ducts 5 µm wide and 19 µm long,
situated on margin; smaller macrotubular ducts 3 µm wide and 19 µm long scattered
throughout the venter. Microtubular ducts situated marginally and submarginally on head
and thorax. Cruciform pores 4 µm long; a few number in submarginal area, marginally on
head. A few hair-like setae present on submedian and submargianal venter, submedian
ones longer. Slightly blunted enlarged setae in two sizes; larger 34 µm long on head
margin; smaller 8–19 µm long marginally on thorax and abdomen. Suranal setae hair-like.
Dorsum: Enlarged setae in two sizes; pointed conial spines 34–41 µm long in a group
of head margin; slightly blunt conical spines in two sizes, longer setae 34–44 µm long
in a row on margins, 3 or 4 on margin of segments; shorter setae numerous, 8–19 µm
long, forming sparse bands on thoracic and abdominal segments, absent from abdominal
segment VIII; twelve dorsal setae on abdominal segment VII. Macrotubular ducts 19 µm
long and 5 µm wide, scattered among spines. Microtubular ducts 4 µm long, scattered.
Anal ring situated on margin of dorsum; oval, sclerotized, 52 µm wide, 54.5 µm long,
with one row of pores and with 8 setae 113 µm long. Cauda not seen (after Hadzibejli
(1960), with modifications and a redescription and new drawing from the holotype and a
paratype, where the sizes are somewhat different).
Ecology
Host plant: Festuca montana.
Distribution: Georgia.
Biology: Overwinters in stage of eggs, in eggsacs, the females lay about 20–38 eggs.
Egglaying ended at end of November. Hatching of nymph starts from April, adults
appeared in August, with one generation in a year (Hadzibejli, 1960).
Acanthococcidae 189
Anophococcus agropyri (Borchsenius, 1949) (Fig. 72) Revived combination

Rhizococcus agropyri Borchsenius, 1949: 359.
Lectotype: Female. Kazakhstan (Alga, Aktyubinsk Oblast), on Agropyron sp.,

18.viii.1936, by N. Borchsenius. By subsequent designation Danzig, 1962a: 852
(type no. 151-36). Deposited in ZMAS. Notes: Single slide contains 5 specimens. A
specimen was selected by Danzig (1962a) as the "holotype"; which must be considered
the lectotype.
Common name: Borchsenius' felt scale
Lit.: Bazarov, 1968b: 77; Danzig, 1962a: 841; 1964: 634; 1971a: 823; 1985: 111;
Dziedzicka & Koteja, 1971: 575; Fetykó et al., 2010: 296; Gertsson, 2001: 126; Hoy,
1963: 68; Kawecki, 1985: 29; Kaydan et al., 2002: 255; Kosztarab & Kozár, 1978: 69;
1988: 298; Koteja, 1974b: 76; Koteja & Zak-Ogaza, 1966: 310; Kozarzhevskaya &
Reitzel, 1975: 7; Kozár et al., 1996: 61; Kozár et al., 1991: 63; Kozár et al., 2004: 55;
Kozár et al., 2013: 55; Kozár & Walter, 1985: 75; Köhler, 1998: 396; Longo et al., 1999:
121; Marotta, 1993: 155; Matesova, 1968: 115; Milonas et al., 2008: 143; Myartseva,
1981: 97; Ossianilson, 1985: 146; Ouvrard & Kozár, 2009: 102; Pellizzari, 1993: 51;
Pellizzari & Kozár, 2011: 66; Rogojanu, 1966: 432; Tang & Hao, 1995: 519; Tao, 1999:
34; Tereznikova, 1981: 17; Ter Grigorian, 1983: 881; Tsalev, 1968: 207; Ülgentürk et
al., 2003, Wang, 2001: 225 (Miller et al., 2013).
Rhizococcus graminicola; Borchsenius, 1949: 359.

Syntypes: Female. Type data: Uzbekistan (Tashkent). Deposited in ZMAS. Notes:
Danzig (1962a: 856) first suggested the possibility that R. graminicola is a junior
synonym of Eriococcus agropyri, stating that presence of three additional enlarged
setae on abdominal segments other than the 7th and the ratio of length of anal setae
and preanal hairs did not seem to be reliable characters for an independent species.
Synonymy formalized by Köhler (1998): 396.
Rhizococcus obscurus; Borchsenius, 1949: 360.
Holotype: Female. Tadjikistan (Ramit District, Jabroz). By original designation.
Notes: Danzig (1962a: 858), first suggested the possibility that R. obscurus was a
junior synonym of Eriococcus agropyri, one adult female on 1 slide in ZMAS (personal
correspondence, Danzig, 1996). Deposited in ZMAS. Synonymy formalised by
Kosztarab & Kozár (1978: 69).
Eriococcus obscurus; Hoy, 1963: 104. Change of combination.
Eriococcus agropyri; Hoy, 1963: 68. Change of combination.
Eriococcus graminicola; Hoy, 1963: 92. Change of combination.
Acanthococcus agropyri; Kosztarab & Kozár, 1978: 69. Change of combination.
Anophococcus agropyri; Koteja, 1990: 120. Change of combination.
Acanthococcus graminicola; Miller & Gimpel, 1996: 601. Change of combination.
Eriococcus agropyri; Miller & Gimpel, 2000: 119. Revived combination.
Rhizococcus agropyri; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Adult female up to 4 mm long, 1.2–1.7 mm wide, Ovisac gray, white or yellowish, with
almost flat dorsum and carinate submargin. Adult female elongate-oval, yellow.
Mounted female
Antennae 7 rarely 6 segmented, 260–279 µm long, length of segments: I: 37.2, II: 35.0,
III: 50.6, IV: 45.5, V: 27.7, VI: 27.1 and VII: 37.3 µm long. Frontal tubercle present. Eyes
situated on margin. Anal lobe apex and inner margin distinctly sclerotized, apical seta
234–315 µm long, subapical seta 150 µm long, suranal setae 105 µm long, anal lobes with
3 dorsal setae, 30 and 52 µm long, the lateral seta truncated, 25 µm long.
Venter: Labium about 125 µm long, 80 µm wide; stylet loop 1.5 times as long as labium.
Apical labial segment with 6 pairs of long setae, the median setae much shorter than
others. Legs 570–645 µm long, hind coxae and femur with a group of translucent pores;
hind femur 137–180, tibia 125–143, tarsus 170 µm long, claw with denticle; tarsal and
claw digitules longer than claw, slightly knobbed. Quinquelocular and septalocular pores
double rimmed, numerous present over entire venter. Cruciform pores present on
margin. Macrotubular ducts of one size, scattered all over the venter. Microtubular ducts
present on margin. Hair-like setae short. Enlarged setae of small size only, present along
body margin.
Dorsum: Enlarged setae, conical with truncate or rounded apex, form a row along body
margin, with 2 setae on margin of abdominal sternites I-VII; other enlarged setae on
dorsum small, 4.5 µm long, form rows along body margin. Macrotubular ducts numerous
all over dorsum. Microtubular ducts numerous, narrow, short. Anal ring dorsal, with 8
setae, each 100–129 µm long, partly with double row of pores. Cauda present (after
Danzig (1990), with some modifications based on lectotypes).
Other stages
Mounted first instar nymph (Fig.72, top right).

with strong sensory setae as in adult female. Dorsum with six rows of equal long, spines.
Tubular ducts absent, on dorsum with four sparse pairs of microtubular ducts. Venter
with transverse rows of six small hair-like setae on each abdominal segment; median
setae longer than others. Cruciform pores present on thoracic segments. With one pair
of quinquelocular pores on each thoracic segment, one near each spiracle, plus one pair
on frons. Stylet loop reaching the third abdominal segment. Anal ring normal, with 6
hair-like setae.
Acanthococcidae 191
Figure 72. Anophococcus agropyri (Borchsenius, 1949), female, after Danzig (1962) with
modifications. First instar on top right, original.
Ecology
Host plant: Agropyron sp., A. cristatum, A. fragile, A. repens, A. sibiricum, Agrostis sp.,
Aneurolepidium sp., Cissus sp., Cynodon dactylon, Elymus sp., E. angustus, E. giganteus, Kalidium
gracile, Lolium sp., and Scabiosa sp.
Distribution: Bulgaria, China, Greece, Hungary, Italy, Kazakhstan, Moldova, Poland,
Romania, Russia, Sweden, Tajikistan, Turkey, Ukraine, Uzbekistan.
Biology: On the shoots, at the base, and on the upper surface of the leaves. Often in
high numbers. They form compact colonies often overlaying each other on leaf sheaths.
Females collected from May through October, laid 98-211 eggs (Matesova, 1968).
Probably has more than one generation per year. Often becoming a pest.
Acanthococcidae 193
Anophococcus cingulatus (Kiritchenko, 1940) (Fig. 73) Redescription,

Combination nova
Eriococcus cingulatus Kiritchenko, 1940: 131.
Lectotype: Female. Ukraine (Crimea, Tuak), on Astragalus sp., 02.ix.1928. By

subsequent designation Danzig, 1996: 521 (type no. 311-58). Deposited in ZMAS.
Notes: Type series consists of one paralectotype on same slide as lectotype. The
remaining paralectotypes are from several localities. Danzig (1996) states "Odessa, on
Achillea, 26 females at 2 preparations; same locality on Artemisia roots, 28.v.1936, Yu.
Shuvalova, 3 females at one preparation; the Crimea, Tuak, on Astragalus, 16 females
at one preparation."
Common name: Kiritchenko's felt scale.
Lit.: Borchsenius, 1949: 50; Danzig, 1962a: 841; 1964: 634; 1971a: 823; 1985: 111;
1986 241; 1988: 709; 1996: 521; Fetykó et al., 2010: 296; Hua, 2000: 138; Kaydan et
al., 2007: 90; Kosztarab & Kozár, 1978: 69; 1988: 299; Kozár et al., 2013: 55; Kozár &
Walter, 1985: 75; Köhler, 1998: 397; Matesova, 1955: 202; 1957: 169; 1968: 116; 1971:
27; Ouvrard & Kozár, 2009: 102; Tang & Hao, 1995: 519; Tang & Li, 1988: 66–67, 68;
Tao, 1999: 35; Tereznikova, 1981: 22; Ter-Grigorian, 1969: 101, 1983: 878; Ülgentürk
et al., 2003: 442; Wang, 1982b: 143; 2001: 225 (Miller et al., 2013).
Rhizococcus cingulatus; Borchsenius, 1949: 356. Change of combination.

Eriococcus cingulatus; Hoy, 1963: 118. Revived combination.
Acanthococcus cingulatus; Danzig, 1975: 62. Change of combination.
Eriococcus cingulatus; Miller & Gimpel, 2000: 163. Revived combination.
Rhizococcus cingulatus; Kosztarab & Kozár, 1988: 300. Revived combination.
Acanthococcus cingulatus orientalis; Danzig, 1975: 79. Synonym nova
Holotype: Female. Russia (Southern Primor'ye, Lazvoskiy Reserve, shores of the Sea
of Japan, close to Glazkovka) (type no. 87-70). By original designation. Deposited
in ZMAS. Notes: There are 2 paratypes on the same slide as the holotype and 3
paratypes on a second slide. There are 4 other paratype slides with 8 females from
Primorskiy Kray and 4 slides with 9 females from Vladivostok.
Acanthococcus cingulatus orientalis; Synonymy by Kosztarab & Kozár (1988), according to
Ben-Dov et al. (2013). Notes: It was not a formal new synonymy by Kosztarab & Kozár
(1988), only a remark for literature comments on similarity).
Rhizococcus orientalis; Tang & Hao, 1995: 521. Change of status and combination.
Rhizococcus terrestris; Matesova, 1957: 169. Synonym nova
Holotype: Female. Kazahstan (Usek River), on Clematis songorica, 20/08/1951.
By original designation (type no. 101). Deposited in ZMAS. Notes: In Matesova's
description the terms type and ecotypes are used for the holotype and paratypes.
Paratypes in AAKA.
Eriococcus terrestris; Hoy, 1963: 118. Change of combination.
Acanthococcus cingulatus terrestris; Danzig, 1975: 54. Change of status and combination.
Acanthococcus cingulatus terrestris; Synonymy by Kosztarab & Kozár (1988) according to

Ben-Dov et al. (2013). Notes: It was not a formal new synonymy by Kosztarab & Kozár
(1988), only a remark for literature comments on similarity).
Acanthococcus terrestris; Kozár, 2009: 94. Combination nova.
Description
Unmounted female
Adult female egg-shaped and of a yellow or rusty dark reddish color. Ovisac grey, white
or yellowish, up to 3.6 mm long, 2.2 mm wide.
Mounted female
220–251 µm, length of segments: I: 45.0–67.5, II: 30–40, III: 42.5–50.0, IV: 35.0–37.5, V:
20–25, VI: 15–17.5, and VII: 37.5–40 µm long; each segment covered with a few, strong
hair-like setae (except third segment); apical segment with apical seta 45.0–47.5 µm long;
apical segment also with 3 sensory falcate setae, 17.5–30 µm long; segment VI with 1
sensory falcate seta 25–35 µm long, segment V with 1 sensory falcate seta, 15.0–17.5
µm long. Frontal lobe and frontal tubercle present. Eyes situated on venter near margin.
Anal lobes strongly developed, each with 3 enlarged setae, each 55–63 µm plus 0 or 1
microtubular duct on dorsal surface; apical seta 285–333 µm; ventral hair-like subapical
seta 118–135 µm long, suranal setae 55 µm long.
short, 7.5–12.5 µm long, stylet loop extends to mesothoracic coxae. Legs well developed;
lengths of segments and digitules of prothoracic legs; coxa: 70–80, trochanter: 60–70,
femur: 120–125, tibia: 100–105, tarsus: 95–105 and claw: 27.5–30.0, tarsal digitules: 45–
50, claw digitules: 27.5–32.5 µm long; lengths of segments and digitules of mesothoracic
legs; coxa: 62.5–75.0, trochanter: 60.0–67.5, femur: 120–127.5, tibia: 107.5–112.5,
tarsus: 95.0–102.5 and claw: 32–35, trochanther + femur: 185, tibia + tarsus: 210, tarsal
digitules: 45.0–47.5, claw digitules: 32–35 µm long; lengths of segments and digitules of
metathoracic legs; coxa: 85–90, trochanter: 60–65, femur: 125–135, tibia: 112.5–117.5,
tarsus: 107.5–112.5 and claw: 30–35, trochanther + femur: 190–195, tibia + tarsus: 225–
230, tarsal digitules: 45.0–47.5, claw digitules: 27.5–35 µm long, claw thick, with denticle;
metathoracic coxae and femur with many pores, tibiae each with 5 setae (median seta
present), tarsi each with 5 setae. Length of spiracles 60–70 µm; diameter of spiracular
peritreme 30.0–37.5 µm, posterior spiracles slightly larger than anterior. Derm with a
sparse covering of scattered flagellate hair-like setae, each 12.55–80.0 µm long. Enlarged
setae, situated on submargin of last abdominal segments in 1 row, each 10–13 µm
long. Multilocular pores each 5–8 µm in diameter with 3, 5 or 8 loculi (triloculars 3–5
µm, quinqueloculars and octoloculars 5–8 µm), double rimmed, distributed in sparse
bands on all abdominal and thoracic segments and head. Macrotubular ducts of two
sizes, larger ducts, each 7–8 µm wide and 21–23 µm long, present on submargin on on
Acanthococcidae 195
Figure 73. Anophococcus cingulatus (Kiritchenko, 1940), female. Original.

abdomen, thorax and head, smaller ducts, each 4–6 µm wide and 20.0–27.5 µm long,
present on median area on abdomen, thorax and head. Microtubular ducts, each 5–6 µm
long scattered on submargin, sometimes on the median area of thorax. Cruciform pores
few, each, 3–4 µm in diameter, distributed on the submargin of thorax and head.
Dorsum: Marginal enlarged setae slightly truncated, long, each 40.0–72.5 µm, forming a
marginal row of 2 or 3 on each segment, submarginal setae conical, long, each 12.5–55.0
µm, longest on thorax and head, dorsal setae, conical, small 5–15 µm long, arranged
in transverse rows across each body segment, rows irregular on head. Macrotubular
ducts, each 7–9 µm wide and 20–27 µm long, scattered throughout dorsum, generally in
segmental bands. Microtubular ducts with 2 sclerotized areas, each 5–6 µm long scattered
over dorsum. Anal ring sclerotized, with on each inner side, 67.5–70.0 µm in diameter,
with 8 setae, each 105–135 µm long; anal ring situated on margin of dorsum. Cauda
present.
Ecology
Host plant: Artemisia absinthium, A. austriaca, A. dracunculus, A. ordosica, A. schrenciana, A.
sublissingiana, Achillea millefolium, Astragalus sp., Centaurea cyanus, C. orientalis, Clematis sp., C.
songorica, Camphorosma sp., Dianthus sp., Galium sp., Koeleria sp., K. macrantha, Medicago sativa,
Potentilla sp., Pyrethrum sp., Russelia sp., Salix sp., Scabiosa ucranica, Stipa sp., Taraxacum sp.,
Trigonella sp., T. ruthenica, Trinia sp., Vitis sp.
Distribution: China, Hungary, Italy, Iran, Kazakhstan, Mongolia, Romania, Russia,
South Korea, Turkey, Ukraine.
Biology: A xerophilous species known only from steppic regions, on the roots of host
plants. Males are known. Females present from May to September and each lays about 70
eggs. Young females and molting last-stage nymph were found in the last part of August.
Many of the specimens that Kiritchenko collected were parasitized. This species has one
generation per year (Kiritchenko, 1940).
Comment
Rhizococcus terrestris Matesova, 1957 and R. orientalis (Danzig, 1975), and cited by Tang &
Hao (1995) are here treated as Synonym nova. The descriptions, drawings of Danzig
(1962a, 1975), Matesova (1957), Tang & Hao (1995) and the study of type materials
and additional collections from Kazahstan, Tatarstan, Moldova and Hungary, does not
give enough base to treat them as separate species. In addition it was recognized that R.
orientalis described by Tang & Hao (1995) is different from other specimens that we have
observed in this study by having two enlarged marginal setae on abdominal segments.
Acanthococcidae 197
Anophococcus confusus (Danzig, 1962) (Fig. 74) Revieved combination

Rhizococcus confusus Danzig, 1962a: 854.
Holotype: Female. Russia (Leningrad Oblast, Priozersk district), on dead needles on

soil, 06.viii.1956, by E. Danzig. By original designation. Deposited in ZMAS. Notes:
There also are 2 paratypes on 1 slide in ZMAS.
Lit.: Dziedzicka, 1977: 59; Dziedzicka & Koteja, 1971: 575; 1974a: 322; 1974b: 76;
Koteja & Zak-Ogaza, 1981: 514; Kozár & Walter, 1985: 75; Köhler, 1998: 397-398;
Miller & Gimpel, 1999: 213; 2000: 185-186; Lagowska, 2002: 243; Lagowska &
Koteja, 1996: 31; Tang & Hao, 1995: 519, 525, 653 (Miller et al., 2013).
Eriococcus confusus; (Danzig, 1962a); Kawecki, 1985: 29. Change of combination.

Anophococcus confusus; Koteja, 1990: 120. Change of combination.
Acanthococcus danzigae; Miller & Gimpel, 1996: 605. Change of combination and replacement
name for Eriococcus confusus (Danzig, 1962a). Notes: Though the combination Acanthococcus
confusus was not published by Miller & Gimpel (1996), this change of combination was
implicit in their work, which made necessary to introduce their proposed replacement
name as Acanthococcus danzigae Miller & Gimpel, 1996.
Eriococcus danzigae; Miller & Gimpel, 1999: 213. Change of combination. Acanthococcus
confusus Danzig (1962a) is a junior secondary homonym of Acanthococcus confusus (Maskell,
1892). However R. confusus Danzig have been transferred to Anaphococcus, and treated here
as it is, consequently the homonymy does not persist any more and the original species
name as An. confusus is reestablished.
Rhizococcus confusus; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Adult female is up to 2 mm long.
Mounted female
Antennae 7 segmented, with short segments, length of segments: I: 35, II: 26, III: 35, IV:
25, V: 22, VI: 22 and VII: 35 µm long; segments with a few hair-like setae. Frontal lobe
well developed. Anal lobes sclerotized, with three strong enlarged setae.
Venter: Labium well developed, basal segment with 2 pairs of almost equal long setae.
Legs medium sized; posterior coxae with pores; tarsus longer than tibiae, claws with
denticle near tip; femur 121 µm, tibia 125 µm and tarsus 164 µm; tarsal and claw digitules
slightly knobbed, longer than claw. Multilocular pores with 3–6 loculi; numerous present
over entire venter. Macrotubular ducts of two sizes; larger one same as those on dorsum,
on body margin; narrower one 15 µm long, and 4 µm long µm wide, scattered all over
the venter. Microtubular ducts present on margin. Hair-like setae short, 6–12 µm long.
Suranal setae 62 µm long. Enlarged setae of small size only, present along body margin.
Few cruciform pores present on margin.
Dorsum: Enlarged setae wide conical, apices blunted, marginal setae much larger than
other dorsal setae, 2 or 3 setae on lateral margin of abdominal segments; larger 25–
30 µm long, the third very short, only 9–12 µm long. Microtubular ducts short with
two sclerotized areas. Slide-mounted adult female with: 7-10 enlarged setae on margin
of most abdominal segments; marginal setae larger than dorsal setae; anal lobes with
3 enlarged setae. Macrotubular ducts numerous, present all over dorsum 15 µm long
and 6 µm wide. Microtubular ducts numerous, narrow, short, with two sclerotized areas.
Anal ring dorsal, with 8 setae with partly two rows of pores with cupola-like spinulae.
Anal ring setae 110 µm long. Cauda weakly developed (after Danzig (1962a), with some
modifications based on two paratypes).
Ecology
Host plant: Poaceae and egg laying females were found on dry needles of Pinus sp.
Distribution: Poland, Netherlands, Russia.
Biology: On grasses and dry pine needle on soil (Danzig, 1962a).
Acanthococcidae 199
Figure 74. Anophococcus confusus (Danzig, 1962), female. After Danzig (1962) with
modifications.
Anophococcus evelinae (Kozár, 1983) (Fig. 75) Combination nova

Rhizococcus evelinae Kozár, 1983: 144.
Holotype: Female. Slovenia (Portoroz), on Bromus sp., 25.iv.1981, by F. Kozár.

By original designation (type no. 1562. Deposited in HNHM. Notes: Paratype in
ZMAS.
Lit.: Kozár & Walter, 1985: 75; Kozár, 1985: 202; Miller & Gimpel, 1996: 600; 1999:
213; 2000: 206; Lagowska, 2002: 239; Lagowska & Koteja, 1996: 29; Milonas et al.,
2008: 143; Tang & Hao, 1995: 520 (Miller et al., 2013).
Acanthococcus evelinae; Miller & Gimpel, 1996: 600. Change of combination.

Rhizococcus evelinae; Köhler, 1998: 398. Revived combination.
Eriococcus evelinae; Miller & Gimpel, 1999: 213. Change of combination.
Rhizococcus evelinae; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Adult female is up to 2 mm long, yellow in life.
Mounted female
Adult female, 3 mm long, 1 mm wide. Antennae 7 segmented, 310 µm long; segments
with a few hair-like setae. Anal lobes with three strong spines.
Venter: Labium 3 segmented, 90 µm long; stylet loop as long as labium. Legs medium
sized; prothoracic legs 650–660, mesothoracic legs 690–700, metathoracic legs 710–
720 µm long, metathoracic coxae with some translucent pores; tarsus as long as tibiae,
claws with denticle near tip; tarsal and claw digitules slightly knobbed, longer than claw.
Multilocular pores with 5 and 7 loculi, double rimmed, numerous, on abdominal segment
in rows, a few on thorax and head, 3-5 pores associated with spiracles. Macrotubular
ducts of two sizes, larger ones on margin; narrower ones scattered all over the venter.
Microtubular ducts present only on margin. Few cruciform pores present on margin.
Hair-like setae short, forming groups and rows, suranal setae 62 µm long, and with 1-3
submarginal enlarged setae.
Dorsum: Enlarged setae conical, apices truncate or slightly rounded, marginal setae
conspicuously larger than other dorsal setae, 4 or 5 setae on lateral margin of abdominal
segments; larger 25–30 µm long, the third very short, only 9–12 µm long. Microtubular
ducts short with two sclerotized areas. Slide-mounted adult female with: 7-10 enlarged
setae on margin of most abdominal segments; marginal setae larger than dorsal setae;
anal lobes with 4 setae. Macrotubular ducts numerous, present all over dorsum 15 µm
long and 6 µm wide. Microtubular ducts numerous, short, with 2 sclerotized areas. Anal
ring dorsal, with 4 pairs of setae with partly a row of pores. Anal ring setae 120–130 µm
long (after Kozár (1983)).
Acanthococcidae 201
Figure 75. Anophococcus evelinae (Kozár, 1983), female. After Kozár (1983) with
modifications.
Ecology
Host plant: Bromus sp., Cynodon dactylon.
Distribution: Greece, Slovenia.
Biology: On the leaves.
Anophococcus formicicola ( Newstead, 1897) (Fig. 76) Redescription,

Combination nova
Eriococcus formicicola Newstead, 1897a: 102.
Lectotype: Algeria (Constantine), on wooded slope of the Mansourah), on Cynodon

dactylon, 1896, by A. E. Eaton. Designated here (BMNH-1945-121); (No 7721 PPI).
Paralectotype (BM 1940, 180); (No 7720 PPI). Deposited in BMNH. Notes: There
are pieces of possibly 4 adult female syntypes on 1 slide in the BMNH.
Common name: Bermuda-grass felt scale.
Lit.: Balachowsky, 1927: 189; Borchsenius, 1949: 357; Danzig, 1962a: 840; Fernald,
1903: 74; Gómez-Menor Ortega, 1960: 201; Hoy, 1963: 90; Kosztarab & Kozár,
1978: 69; Koteja, 1974a: 76; Kozár et al., 2013: 55; Kozár & Walter, 1985: 75; Köhler,
1998: 398; Lindinger, 1912: 130; Martin-Mateo, 1985: 92; Miller & Gimpel, 1996: 600;
Newstead, 1907a: 16; Trabut, 1911: 71; Tsalev, 1968: 53 (Miller et al., 2013).
Nidularia formicicola; Lindinger, 1933a: 108. Change of combination.

Eriococcus formicicola; It is a valid name, but Eriococcus formicicola, cited by Borchsenius
(1937:182), is a misidentification of An. cynodontis.
Acanthococcus formicicola; Miller & Gimpel, 1996: 600. Change of combination.
Rhizococcus formicicola; Kozár, 2009: 106. Change of combination.
Eriococcus cynodontis; Kiritchenko, 1940: 133. Synonym nova.
Lectotype: Female. Ukraina (Crimea, Kekeneis). Designated by Danzig (1996).
Notes: Two paralectotype were studied. Deposited in ZMAS.
Rhizococcus cynodontis; (Kiritchenko, 1940), Danzig, 1962a: 844. Change of combination.
Acanthococcus cynodontis; Tereznikova, 1981: 26. Change of combination.
Rhizococcus cynodontis; Kosztarab & Kozár, 1988: 301. Revived combination.
Eriococcus cynodontis; Miller & Gimpel, 2000: 208. Revived combination.
Rhizococcus cynodontis; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Adult female is elongate oval. Sac of female is short, ovate, white and closely felted. Sac
of male has same colour and texture as that of the female.
Acanthococcidae 203
Figure 76. Anophococcus formicicola, (Newstead, 1897), female, first instar on top left.
Original.
Mounted female
Adult female oval, 2.1 mm long, 1.3 mm wide. Antennae 6 segmented; length of
segments: I: 32–43, II: 36–43, III: 89–103, IV: 20–24, V: 22–24 and VI: 36 µm long;
segments with a few hair-like setae; apical segment with 31–34 µm long apical seta; apical
segment also with 3 sensory falcate setae, 19–21 µm long; two preapical segments also
with 1 sensory falcate seta: on segment IV:13–15, segment V: 17–21 µm long. Frontal
lobe absent, frontal tubercle present. Eyes situated on venter near margin. Anal lobes well
developed, sclerotized, with 3 blunted conical enlarged setae, ventrally with two flagellate
setae; anal lobe with 98 µm long apical seta.
Venter: Labium 3 segmented, joint length of two apical segments 72–77 µm; basal
segment with 2 pairs of almost equal long setae, apical segment with 6 pairs of hair-
like setae, apical seta short, spine-like. Stylet loop longer than labium. Width of anterior
spiracles 26–33 µm. Legs normal: length of segments of prothoracic legs; coxa: 61–67,
trochanter: 40–46, femur: 138–154, tibia: 103–113; tarsus: 108–111 and claw. 28–30
µm long, tarsal digitules: 48–53, claw digitules: 34–41 µm long; length of segments of
mesothoracic legs; coxa: 67–79, trochanter: 43–48, femur: 139–154, tibia: 112–132,
tarsus: 112–129, claw: 29–33 µm long, tarsal digitules: 48–51 µm, claw digitules: 36–
38 µm long; length of segments of metathoracic legs; coxa: 77–86, trochanter: 38–43,
femur: 144–163, tibia: 115–132, tarsus: 137; claw: 32 µm long, tarsal digitules: 53 µm
long, claw digitules: 41 µm long, tarsal and claw digitules slightly knobbed, longer than
claw; meso- and metathoracic coxae with spinulae on anterior surface; metathoracic
coxae and femur also with small pores on both anterior (ventral) and posterior (dorsal)
surfaces; claw with a denticle; legs with a few hair-like setae, and with one sensory pore
on tarsus; tibia with 5 setae. Multilocular pores 6 µm in diameter and with 5 or 7 loculi,
double rimmed, pores more abundant near spiracles, septalocular pores more abundant
on abdomen. Macrotubular ducts of two sizes: larger macrotubular ducts 3–6 µm wide
situated on margin; smaller macrotubular ducts 3 µm wide and scattered throughout the
venter. Microtubular ducts sparse marginally and submarginally, sparse on submedian
thorax. Cruciform pores few in submarginal area, sparse on head and thorax margin.
A few short, hair-like setae present on venter; no enlarged setae; suranal setae hair-like.
Dorsum: Enlarged setae of two kinds; pointed conical setae 34–38 µm long in a group
of head margin; blunt conical spines in 3 sizes, longer setae, 31–36 µm long and medium
sized setae, 22–25 µm long, in a row on margins, four on margin of segments; shorter
setae more numerous, 4–6 µm long, forming sparse rows on dorsum, but absent from
abdominal segment VIII; 12 dorsal setae on abdominal segment VII. Macrotubular ducts
6 µm wide and about 15 µm long scattered among enlarged setae. Microtubular ducts
scattered, 4 µm long. Anal ring situated on margin of dorsum; oval, sclerotized, 50 µm in
diameter, with one row of pores and with 8 setae, 78–80 µm long. Cauda well developed
(Redescription is based on lectotype designed in this study).
Acanthococcidae 205
Other stages
Mounted first instar nymph (Fig. 76, top left)

with strong sensory setae as in adult female. Stylet loop reaching the abdominal segment
III. Dorsum with small spines of equal length. With six longitudinal rows of spines on
dorsum. Microtubular ducts form four sparse rows on dorsum. Venter with transverse
rows of six small hair-like setae on each abdominal segment; median setae longer than
others. Cruciform pores present on thoracic segments. With one pair of quinquelocular
pores on each thoracic segment, one near each spiracle, plus one pair on frons. Anal ring
normal, with 6 hair-like setae. Cauda well developed.
Ecology
Host plant: Cynodon dactylon, Elymus sp., Festuca sp., Hyparrhenia hirta, Sedum sp., Setaria sp.
Distribution: Algeria, Bulgaria, Croatia, Cyprus Island, Greece, Hungary, Italy, Malta,
Russia, Slovenia, Spain, Switzerland, Turkey, Ukraine.
Biology: On wooded slope of the Mansourah. Based on information taken by the
collector the species lives underground in the company of ants Camponotus sp. After rains
the ants bring the species above ground (Newstead, 1897a). Females collected from July
through September found on blades of grass. Males unknown.
Anophococcus granulatus (Green, 1931), (Fig. 77) Redescription,

Combination nova
Eriococcus granulatus (Green, 1931): 263.
Holotype: Female. Iceland (Haukstaoir), on Festuca rubra, by C. H. Lindroth. By

original designation. Deposited in Goteborg: Notes: There are several slides of
this species in the BMNH, but no types. Six slides of this species from the original
collection by C. H. Lindroth deposited in the BMNH, were studied.
Lit.: Danzig, 1968: 304; Foldi, 2001: 305; Goux, 1935: 92; 1940: 65; 1948: 69;
1989: 21; Green, 1931: 263; Hoy, 1963: 93; Jancke, 1955: 289; Kozár et al., 2013: 55;
Kozár & Walter, 1985: 75; Kozarzhevskaya, 1986: 307; Kozarzhevskaya & Reitzel,
1975: 7; Köhler, 1998: 398; Lindinger, 1933a: 116; 1936: 156; Lindroth, 1931: 154;
Ossiannilsson, 1955: 4; Ouvrard & Kozár, 2009: 102 (Miller et al., 2013).
Nidularia granulatus; Lindinger, 1933a: 116. Change of combination.

Acanthococcus granulatus; Ossiannilsson, 1955: 4-5. Change of combination.
Rhizococcus granulatus; Kozár & Walter, 1985: 75. Change of combination.
Eriococcus granulatus; Miller & Gimpel, 2000: 221. Revived combination.
Rhizococcus granulatus; Kozár, 2009: 106, Revived combination.
Description
Unmounted female
Ovisac is grey or ochreous, closely felted, ovate and rounded at both extremities, 2.5–2.75
mm long, 1. 25 mm wide. Adult female ovate. Adult male has well developed wings and
a pair of long white filaments (Green, 1931).
Mounted female
Adult female oval, 1.9–2.6 mm long, 1.0–1.4 mm wide. Antennae 6 segmented, length
of segments: I: 38–43, II: 34–36, III: 84, IV: 19–24, V: 19–20 and VI 31–36 µm long;
with a few hair-like setae; apical segment with 33–39 µm long apical seta; apical segment
also with 3 sensory falcate setae, 30–36 µm long; two pre-apical segments also with 1
sensory falcate seta: on fourth segment 19–22, fifth segment 30–36 µm long. Frontal
lobe absent, frontal tubercle 3 µm wide. Eyes situated on venter near margin. Width
of anterior spiracles 27–29 µm. Nodulation on segment margins both on venter and
dorsum. Anal lobes well developed, sclerotized, with 3 blunted enlarged setae, ventrally
with two flagellate setae; anal lobe with 182–194 µm long apical seta.
segment with 2 pairs of setae, apical segment with 6 pairs of equal length hair-like setae;
stylet loop very short, about half of length of labium. Legs well developed; length of
segments of prothoracic legs; coxa: 65–72, trochanter: 58–60, femur: 132–144, tibia:
120–129; tarsus: 120 and claw: 26–29 µm long, tarsal digitules: 43, claw digitules: 34;
length of segments of mesothoracic legs; coxa: 62–72, trochanter: 56–62, femur: 120–
142, tibia: 110–120, tarsus: 121–126, claw: 26–27 µm long, tarsal digitules: 46 µm long,
claw digitules: 34–36 µm long; length of segments of metathoracic legs; coxa: 74–84,
trochanter: 58–62, femur: 130–158, tibia: 110–134, tarsus: 134–151; claw: 29, tarsal
digitules: 41–48, claw digitules: 31–33 µm long, tarsal and claw digitules slightly knobbed,
longer than claw; meso- and metathoracic coxae with spinulae on anterior surface;
metathoracic coxae and femur with large pores on both anterior (ventral) and posterior
(dorsal) surfaces (not mentioned by Green); claw with denticle; legs with a few hair-like
setae, tibia with 5 setae. Multilocular pores 6 µm in diameter and with 5-9 loculi, mostly
7; distributed in transverse bands across abdominal sternites, in sparse rows on thoracic
segments and head. Macrotubular ducts of two sizes: larger macrotubular ducts 5 µm
wide and 20 µm long, situated on margin; smaller macrotubular ducts 3 µm wide and
20 µm long scattered throughout the venter. Microtubular ducts sparse throughout the
venter. Cruciform pores in a band on submarginal area, and throughoth the head region
5 µm long. A few long hair-like setae present on submedian venter and a few shorter hair-
like setae. Enlarged setae blunted, on margin, 8–10 µm long, form a sparse submarginal
band. Suranal setae short, hair-like.
Dorsum: The cuticula surface with nodulations. Enlarged setae of two kinds; pointed
conical spines 41–43 µm long in a group of head margin; blunt conical spines of two
sizes, marginal setae noticeably longer than other setae on dorsum, longer setae 36–
Acanthococcidae 207
Figure 77. Anophococcus granulatus (Green, 1931), female. Original.

41 µm long in a row on margins, 3 on segments; shorter setae more numerous, 8–10

µm long, forming sparse rows on dorsum, but absent on abdominal segment VIII; ten
enlarged setae on abdominal segment VII. Macrotubular ducts 5 µm wide and 20 µm
long, scattered among spines. Microtubular ducts scattered, 5 µm long. Anal ring situated
on margin of dorsum; oval, sclerotized, 55–60 µm wide, 60–67 µm long, with one row
of pores and with 8 setae each, 107 µm long. Cauda present, with serrated distal margin
(after Green (1931), with modifications based on four slides from the original collection
deposited in BMNH).
Other stages
There is a description of nymphs and adult males in Green (1931) and Jancke (1955).
Ecology
Host plant: Festuca ovina, Festuca rubra.
Distribution: France, Iceland, Hungary.
Acanthococcidae 209
Anophococcus herbaceus ( Danzig, 1962) (Fig. 78) Revived combination

Rhizococcus herbaceus Danzig, 1962b: 22.
Holotype: Female. Russia (St. Petersburg Oblast=Leningrad Obl., Zelenogorsk), on

undetermined Gramineae, 23.viii.1955, by E. Danzig. Deposited in ZMAS.
Common name: Danzig's felt scale.
Lit.: Danzig, 1962a: 840; 1964: 633; 1971a: 823; 1975: 42; 1988: 709; Dziedzicka,
1977: 59; Dziedzicka & Koteja, 1971: 557, Foldi, 2001: 303; Komosinska & Podsiadlo,
1967: 684; Kosztarab & Kozár, 1988: 301. Koteja, 1974a: 322; 2000: 172; Koteja &
Liniowska, 1976: 666; Koteja & Zak-Ogaza, 1969: 310, 1981: 514; 1983: 477; Kozár
et al., 2013: 55; Lagowska, 2002: 243; Miller & Gimpel, 1999: 213; Milonas, et al.,
2008: 143; Tang & Hao, 1995: 520; Tereznikova, 1981: 29; Ülgentürk, et.al., 2003: 442
Acanthococcus herbaceus; Kosztarab & Kozár, 1978: 68. Change of combination.

Eriococcus herbaceus; Kawecki, 1985: 5. Change of combination.
Rhizococcus herbaceous; Kozár & Walter, 1985: 75. Revived combination.
Anophococcus herbaceus; Koteja, 1990: 120, in Nast et al., (1990). Change of combination.
Rhizococcus herbaceous; Köhler, 1998: 398. Revived combination.
Eriococcus herbaceus; Miller & Gimpel, 2000: 229. Revived combination.
Rhizococcus herbaceous; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Ovisac is elongate-oval, white. Adult female is elongate-oval, greenish. 2.4–3.0 mm long,
1.0–1.4 mm wide.
Mounted female
Antennae 6 or 7 segmented, 223–250 µm long, third segment distinctly longest; length of
segments: I: segment 47, II: 34, III: 94, IV: 21, V: 21 and VI: 34 µm long; segments with
a few hair-like setae. Frontal lobe absent, tubercle present. Anal lobes sclerotized, with
three strong enlarged setae.
Venter: Labium 3 segmented, basal segment with 2 pairs of almost equal long setae,
apical segment with 6 pairs of hair-like setae, the apical seta shorter, subapical setae twice
longer slightly capitated. Labium 140 µm long, 90 µm long; stylet loop reaching median
coxae. Hind coxa with spinulae and translucent pores, 180 µm long, femur 137 µm long,
tibia 180 µm long; tarsus longer than tibiae, claws with denticle near tip; tarsal and claw
digitules slightly knobbed, longer than claw. Multilocular pores with 5 or 7 loculi, pores
double rimmed, numerous, form bands on abdominal sternites, scattered on prosoma.
Macrotubular ducts of 2 sizes, 19 µm long, and 6 µm wide, scattered all over the venter.
Microtubular ducts present on margin, microtubular ducts 7 µm long, common on entire
body, except median area of venter, forming a band along ventral margin. A band of
cruciform pores present on margin. Hair-like setae short, 6–12 µm long. Suranal setae 43
µm long. Enlarged setae of small size only 13–21 µm long, present in a row along body
margin.
Dorsum: Enlarged setae conical, apices truncate or rounded, marginal setae much larger
than other dorsal setae, 2 or 3 setae on lateral margin of abdominal segments, 26–52 µm
long. Dorsal enlarged setae 13–26 µm long. Macrotubular ducts scattered, present all
over dorsum. Microtubular ducts short with two sclerotized areas, anal lobes with 3 setae.
Anal ring dorsal, with 8 setae, with partly two rows of pores with cupola-like spinulae.
Anal ring setae 140 µm long. Cauda weakly developed (after Danzig (1962a), with some
modifications based on two paratypes).
Ecology
Host plant: Brachypodium sp., Calamagrostis sp., Carex sp., Cynodon dactylon, Luzula sp., L.
nemorosa, Piptatherum miliaceum.
Distribution: Greece, Hungary, Poland, Russia, Turkey, Ukraine.
Biology: On the leaves. This is a rare boreal species. Females have been collected from
June through September in Poland.
Acanthococcidae 211
Figure 78. Anophococcus herbaceus (Danzig, 1962), female. After Danzig (1962) with
modifications.
Anophococcus iljinae (Danzig, 1972) (Fig. 79) Combination nova

Rhizococcus iljinae Danzig, 1972b: 339.
Holotype: Female. Mongolia (Bayan-Hongor Aymag, 110 km S. Shine-Djinst), on

Iljinia regelii roots, 30.viii.1970, E. Nartshuk. By original designation. (type no. 7-71)
ZMAS. Notes: There is 1 female paratype on the same slide as the holotype and a
second slide with 2 females and the same data. A third slide contains 2 additional
paratypes from different a location in Mongolia.
Lit.: Danzig, 1972b: 339; Hua, 2000: 138; Kozár & Walter, 1985: 75; Köhler, 1998:
399; Miller & Gimpel, 2000: 233; Tang & Hao 1995: 520; Tang & Li, 1988: 75 (Miller
et al., 2013).
Acanthococcus iljiniae; Miller & Gimpel, 1996: 601. Change of combination.

Rhizococcus iljinae; Köhler, 1998: 399. Revived combination.
Eriococcus iljiniae; Miller & Gimpel, 1999: 214. Change of combination.
Rhizococcus iljinae; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Adult female oval, globose.
Mounted female
Up to 4.5 mm long. Antennae 7 segmented, with short segments, segments with a few
hair-like setae. Frontal lobe and tubercle well developed. Anal lobes sclerotized, with
three strong enlarged setae.
Venter: Labium narrow, very long, 3 segmented, basal segment with 2 pairs of equal
short setae, apical segment with 6 pairs of hair-like setae, the apical seta shorter, spine-
like; stylet loop reach behind the median coxae. Legs medium sized, posterior coxae with
pores and spinulae, median coxae with spinulae; tarsus longer and tibiae equal long, claws
with denticle near tip; tarsal digitules slightly knobbed, longer than claw; claw digitules
shorter than claw, spine-like. Multilocular pores with 3-5 loculi, numerous present over
entire venter, pores double rimmed. Macrotubular ducts of two sizes, scattered all over the
venter. Microtubular ducts present on margin. Few cruciform pores present on margin.
Hair-like setae short. Enlarged setae of small size only, present along body margin.
Dorsum: Enlarged setae cylindrical, apices rounded, or truncate, marginal setae much larger
than other dorsal setae, 3 setae on lateral margin of abdominal segments; almost equal long;
marginal setae much longer than dorsal setae. Macrotubular ducts few in sparse rows, two sizes,
short, present all over dorsum. Microtubular ducts short with two sclerotized areas, numerous.
Anal ring dorsal, with 8 long setae, with two rows of pores in some parts. Suranal setae long.
Cauda well developed (after Danzig (1972b), with some modifications based on two paratypes).
Acanthococcidae 213
Figure 79. Anophococcus iljiniae (Danzig, 1972), female. After Danzig (1972) with
modifications.
Ecology
Host plant: Iljinia regelii, Kalidium gracile.
Distribution: Mongolia. Notes: Tang & Li (1988) included this species in their book on
Inner Mongolian Chinese Coccoidea. Tang & Hao (1995) realized that the specimens
that they considered to be R. iljiniae were a misidentification of An. agropyri.
Biology: On the roots.
Anophococcus inermis (Green, 1915) (Fig. 80) Revived combination

Eriococcus inermis Green, 1915: 176.
Lectotype: Female. England (Camberley, Surrey, Virginia Water), on Agrostis curtisii,

?/viii/1914, by E. E. Green. By subsequent designation Williams, 1985a: 370.
Deposited in BMNH. Notes: In addition to the lectotype there are five adult female
paralectotypes on the same slide and an additional slide contains six adult females
paralectotypes in the BMNH.
Common name: Harmless felt scale.
Lit.: Balachowsky, 1934c: 130; 1937b: 6; 1954: 61; Boratynsky & Williams, 1964:
91; Borchsenius, 1949: 47; Borchsenius & Danzig, 1966: 41; Danzig, 1962a: 854;
1964: 634; 1971a: 823; 1975: 43; 1980: 226; 1986: 241; 1988: 79; Dziedzicka, 1977:
5; Dziedzicka & Koteja, 1971: 576; Ferris, 1957b: 86; Fetykó, et al., 2010: 296; Foldi,
2001: 305; Goux, 1937: 93; 1948: 16; Green, 1915: 176; 1921: 149; 1922: 20; 1923:
211; 1928a: 9; Hoy, 1963: 133; Kiritchenko, 1935: 1; Kosztarab & Kozár, 1978: 67;
Koteja, 1974b: 76; Koteja & Zak-Ogaza, 1969: 363; Köhler, 1998: 399; Lagowska
& Koteja, 1996: 31; Malumppy, et al., 2010: 258; Ouvrard & Kozár, 2009: 102;
Schmutterer, 1956b: 66; Tang & Hao, 1995: 518; Tereznikova, 1966: 27; Williams,
1985a: 370 (Miller et al., 2013).
Nidularia inermis; Lindinger, 1933a: 116. Change of combination.

Greenisca inermis; Borchsenius, 1948: 502. Change of combination.
Anophococcus inermis; Balachowsky, 1954: 61. Change of combination.
Rhizococcus inermis; Danzig, 1962a: 854. Change of combination.
Acanthococcus inermis; Danzig, 1975: 64. Change of combination.
Eriococcus inermis; Williams, 1985a: 370. Revived combination.
Rhizococcus inermis; Kozár & Walter, 1985: 75. Revived combination.
Anophococcus inermis; Kosztarab & Kozár, 1988: 286. Revived combination.
Acanthococcus inermis; Miller & Gimpel, 1996: 601. Revived combination.
Eriococcus inermis; Miller & Gimpel, 2000: 235. Revived combination.
Anophococcus inermis; Kozár, 2009: 94. Revived combination.
Acanthococcidae 215
Figure 80. Anophococcus inermis (Green, 1915), female. After Williams (1985) with
modifications.
Description
Unmounted female
Adult female elongate-oval, tapering toward both end; 2.6–3 mm long, 1.3 mm wide.
Adult female is elongate oval. Ovisac is white, slightly tinged with pale ochreous, closely
felted, comparatively smooth. Eggs are yellow. Male puparium similar in color and
substance, but smaller and flatter.
Mounted female
Adult female up to 4.5 mm long. Antennae 6 segmented, with short segments, segments
with a few hair-like setae 150–210 µm long. Frontal tubercle well developed. Anal lobes
sclerotized, with two well develped enlarged setae inner side, seta on outer side very small
spinelike.
Venter: Labium narrow, 3 segmented, basal segment with 2 pairs of setae, apical
segment with 6 pairs of hair-like setae, the apical seta shorter, labium 95–110 µm long;
stylet loop reach behind the median coxae. Legs well developed, hind trochanter + femur
140–170 µm long, hind tibia 90–110 µm long, hind tarsus 110–130 µm long, the tibia
+ tarsus conspicuously longer than trochanter + femur, claw curved, 30 µm long, with
a denticle near apex; hind coxa with a few large translucent pores on outer half and
hind femur sometimes with one or two on anterior edge. Discodial pores varying in size,
numerous on abdomen where, on the posterior segments, many have 7 or occasionally
more loculi but anteriorly on the abdomen, the median area of thorax and around the
spiracles, quinquelocular. Macroducts of two sizes; a larger type, the same as dorsal ducts,
on margins only; a narrower type in median areas as far forward as head margins. Setae
slender in median areas and minute setae, similar to those on dorsum, around margins and
submargins. Cruciform pores present in a submedian zone from head to about abdominal
segment V.
Dorsum: Dorsal setae very small and few, macrotubular ducts about 20 µm long and
microtubular ducts 4 µm long, numerous. Microtubular ducts small with 2 sclerotized
areas, scattered through the body (after Williams (1985a).
Ecology
Host plant: Agrostis curtisii, Deschampsia flexuosa, Festuca ovina, Koeleria askoldensis.
Distribution: France, Georgia, Germany, Lithuania, Netherlands, Poland, Romania,
Russia, Ukraine, United Kingdom (England).
Biology: Ovisacs are nearly always attached to dry and dead blades, seldom on the green
parts. On the upper surface of the leaves. Feed on leaves of grasses. Females collected
from July to September.
Acanthococcidae 217
Comment:
This species was originally the type of the genus Greenisca Borchsenius, 1948. Later,
Borchsenius and Danzig (1966) replaced the misidentified type species with An. gouxi.
Anophococcus insignis (Newstead, 1891) (Fig. 81) Revived combination

Eriococcus insignis (Newstead, 1891): 164.
Lectotype: Female. England (Cheshire, Ince), on Agrostis sp., 1890. By subsequent

designation Williams, 1985a: 372. Deposited in BMNH. Notes: the lectotype has
been selected and clearly marked, from four specimens on a single slide labelled
“Cotype” by Newstead. In addition to the lectotype adult female, there are three
paralectotype adult females on the same slide in the BMNH.
Common name: Conspicuous felt scale, remarkable eriococcin, remarkable felt scale.
Lit.: Bazarov, 1962: 77; Borchsenius, 1949: 47; Borchsenius & Danzig, 1966: 41;
Boratynsky & Williams, 1964: 92; Danzig, 1962a: 841; 1964: 633; 1971a: 822; 1975:
43; 1980: 223; 1986: 241; 1988: 709; Dziedzicka, 1977: 59; Dziedzicka & Koteja,
1971: 561; Fetykó, et al., 2010: 296; Foldi, 2001: 305; Gill, 1993: 157; Goux, 1931: 332;
1935: 92; 1940: 65; 1948: 69; Green, 1915: 176; 1920: 116; 1922: 20; 1923: 211; 1925:
517; 1926: 182; 1927: 28; 1928a: 8; 1928b: 30; Hadzibejli, 1983: 269; Hoy, 1963: 95;
Hua, 2000; Kiritchenko, 1928: 112; 1931: 311; 1940; Kosztarab & Kozár, 1978: 69;
1988: 303; Koteja, 1974b: 76, Koteja & Zak-Ogaza, 1969: 363; Kozár et al., 2013: 55;
Lagowska & Koteja, 1996: 31; Miller & Miller, 1992: 5; 1993: 7; Ouvrard & Kozár,
2009: 102; Schmutterer, 1952: 378; Tang & Hao, 1995: 520; Tereznikova, 1981: 29;
Ter-Grigorian, 1983: 879; Williams, 1985a: 372 (Miller et al., 2013).
Eriococcus greeni; Lindinger, 1912: 367. Incorrect synonymy. Notes: Lindinger (1912)
incorrectly treated Eriococcus greeni as a junior synonym of E. insignis. These taxa are
considered as distinct by Hoy (1963) and Williams (1985a).
Nidularia insignis; Lindinger, 1933a: 116. Change of combination.
Eriococcus socialis; Lindinger, 1936: 156. Incorrect synonymy. Notes: Lindinger (1936)
incorrectly synonymized E. socialis with E. insignis (Hoy, 1963; Williams, 1985a).
Eriococcus saratogensis Rau, 1938: 157-159.
Holotype: Female. United States (New York, Saratoga Springs), on Hystrix patula,
09/11/1936, by Rau. By original designation. Synonymy by Miller & Miller, 1992:
48–51. Notes: The location of type material is unknown, but Miller & Miller (1993)
have studied topotypes.
Rhizococcus insignis; Borchsenius, 1949: 357-358. Change of combination.
Acanthococcus insignis; Tereznikova, 1975: 39. Change of combination.
Rhizococcus insignis; Kozár & Walter, 1985: 75. Revived combination.
Anophococcus insignis; Koteja, 1990: 120, in Nast et al., 1990: 120. Change of combination.
Acanthococcus insignis; Miller & Miller, 1992: 236. Revived combination.
Rhizococcus insignis; Köhler, 1998: 399. Revived combination.

Eriococcus insignis; Miller & Gimpel, 2000: 236. Revived combination.
Rhizococcus insignis; Kozár, 2009: 106. Revived combination.
Eriococcus chaoticus; Goux, 1991: 49. Synonym nova.
Holotype: Female. France (Hautes-Pirénées, Artigues), on Gramineae, 12. viii. 1953,
by L. Goux (MNHN 14448). By original designation. Deposited in MNHN.
Acanthococcus chaoticus; Miller & Gimpel, 1996: 599. Change of combination.
Rhizococcus chaoticus; Kozár, 2009: 106. Change of combination.
Description
Unmounted female
Body of adult female elongate-oval, the sides often subparallel, nodulose, largest specimens
2.9 mm long, 1.2 mm wide, sac is oval or elongate and white or cream colored. Adult
females are dark red, elongate-oval. Ovisac is tough, yellowish-white; produced on leaves
of host.
Mounted female
Up to 4.5 mm long. Antennae 7 segmented, 200–270 µm long, segments with a few
hair-like setae. Frontal tubercle well developed. Anal lobes conical, pointed, about twice
as long as wide, moderately sclerotised. Each lobe with an apical seta 280–320 µm long, on
the dorsum 1 outer and 2 inner enlarged setae, on the venter 2 slender setae and a suranal
setae shorter than anal ring setae.
Venter: Labium narrow, about 100 µm long, 3 segmented, basal segment with 2 pairs
of setae, apical segment with 6 pairs of hair-like setae, the apical seta shorter; stylet
loop short. Legs well developed, hind trochanter + femur: 200–250 µm long, hind
tibia: 120–160, hind tarsus: 130–150 µm long the tibia + tarsus conspicuously longer
than trochanter + femur, claw curved, 30 µm long, with a denticle near apex; hind coxa
with a few large translucent pores on outer half and hind femur sometimes with one
or two on anterior edge. Discodial pores varying in size, numerous on abdomen where,
on the posterior segments, many have 7 or occasionally more loculi but anteriorly on the
abdomen, the median area of thorax. Macroducts of two sizes; a larger type, the same
as dorsal ducts, on margins only; a narrower type in median areas as far forward as head
margins. Microtubular duct few on marginal area. Ventral surface with normal slender setae
in median areas, a few enlarged setae on head margin and stiff setae in a marginal zone from
head to anterior abdominal segments, these setae usually larger than the small dorsal setae
but smaller than dorsal marginal setae. Cruciform pores present in a submedian zone from
head to about abdominal segment V.
Dorsum: Dorsal enlarged setae cylindrical, apices narrow, truncate posteriorly, acute or
slightly rounded on thorax and head very small and few; marginal setae conspicuously
larger than other setae on dorsum, 4 or 5 lateral setae on margin of abdominal segments;
40–65 µm long; elsewhere on the dorsum there are small setae in moderate numbers,
Acanthococcidae 219
Figure 81. Anophococcus insignis (Newstead, 1891), female. After Williams (1985) with
modifications.
6–8 µm long; on segments 5-7 these setae are truncate, but anteriorly they are bluntly
pointed and on head they extend to submarginal area near midline; the setae number on
the abdominal segment VII always four. Macrotubualar ducts about 25 µm long, scattered
through. Microtubular ducts are more numerous about 4 µm wide. Cauda rounded,
narrow, sometimes nodulose and lightly sclerotised. Anal ring with 8 setae, each 135 µm
long, each with slightly concave sides (after Williams, 1985a).
Other stages
First instar described by Gómez-Menor Ortega (1968).
Ecology
Host plant: Agropyron cristatum, A. repens, Agrostis sp., A. capillaris, A. curtisii, A. rupestris,
Andropogon sp., Anthoxanthum sp., A. odoratum, Arrhenatherum elatius, Avenastrum sp.,
A. pubescens, Brachypodium sp., Bromus sp., B. inermis, B. mollis, Calamagrostis arundinacea,
Carex sp., Cichorium sp., C. intybus, Cynodon dactylon, Dactylis glomerata, Deschampsia sp.,
D. flexuosa, Elymus sp., Festuca sp., F. ovina, F. pratensis, F. rubra, Hieracium pilosella, Holcus
lanatus, Hypericum sp., Hystrix patula, Koeleria sp., K. askoldensis, Linnaea borealis, Luzula
sp., Malus sylvestris ?, Nardus sp., Phalaris sp., Phleum phleoides, P. pratense, Poa sp., Pteridium
sp., Pteris sp., Rumex sp., Saxifraga sp., Silene sp., Spiraea douglasii, Stipa sp., Thymus sp.,
Trifolium medium, Ulex sp., Ulmus sp., Urtica sp., U. dioica.
Distribution: Armenia, Austria, Bulgaria, former Czechoslovakia, Denmark, France,
Georgia, Germany, Hungary, Iraq, Italy, Kazakhstan, Netherlands, Norway, Poland,
Romania, Russia, Sweden, Ukraine, United Kingdom, USA.
Biology: A common steppe inhabiting mesophilous species; feeds on leaves of
grasses; host genera. Adults have been observed in June. First instars appeared during
the first part of June. Adult females present again in August and September. Based on
this information, it is seem to be two generations per year, at least in Central Europe.
Males known in the United States. Schmutterer (1952) observed one yearly generation
and eggs overwintering. This species damages timothy grass in the northwestern
United States.
Acanthococcidae 221
Anophococcus kondarensis (Borchsenius, 1949) (Fig. 82) Combination

nova
Rhizococcus kondarensis Borchsenius, 1949: 353.
Lectotype: Female. Tajikistan (Gissar Ridge, Kondara River), 21.vii.1940, by N.

Borchsenius (type no. 183-42). Deposited in ZMAS. Notes: One female on same
slide as lectotype.
Lit.: Babaeva, 1980: 57; Bazarov, 1968a: 63; Borchsenius, 1949: 57; Danzig, 1962a:
840; Hoy, 1963: 98; Hua, 2000: 138; Köhler, 1998: 400; Miller & Gimpel, 1996: 601;
Ouvrard & Kozár, 2009: 102; Ter-Grigorian, 1983: 880; Wang, 1974: 333 (Miller et
al., 2013).
Eriococcus kondarensis; Hoy, 1963: 98. Change of combination.

Acanthococcus kondarensis; Ter-Grigorian, 1983: 880. Change of combination.
Rhizococcus kondarensis; Kozár & Walter, 1985: 75. Revived combination.
Acanthococcus kondarensis; Miller & Gimpel, 1996: 601; Revived combination.
Eriococcus kondarensis; Miller & Gimpel, 2000: 247. Revived combination.
Rhizococcus kondarensis; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Adult female elongate-oval. Ovisac gray, white or yellowish, up to 3.5 mm long, 2 mm
wide.
Mounted female
Adult female elongate-oval. Antennae 7 rarely 6 segmented, 260–279 µm long, length
of segment: I: 36, II: 35, III: 46, IV: 56, V: 22, VI: 19 and VII: 32 µm long. Frontal
tubercle developed. Anal lobes with 3 dorsal setae, up to 55 µm long, and the lateral seta
truncated, 25 µm long; anal lobe subapical seta 95 µm long, suranal setae 65 µm long.
Venter: Labium about 125 µm long, 80 µm wide; stylet loop 1.5 times as long as labium.
Apical labial segment with 6 pairs of long setae, the median setae much shorter than
others. Legs medium sized, hind coxae and femur with a group of translucent pores and
spinulae; hind femur: 190, tibia: 147, tarsus: 165 µm long, claw with denticle; tarsal and
claw digitules longer than claw, slightly knobbed. Quinquelocular and septalocular pores
double rimmed, numerous, present over entire venter. Macrotubular ducts of two sizes,
larger ones on marginal band; narrower ones scattered all over the venter. Microtubular
ducts present on margin. Hair-like setae short. Enlarged setae of small size only, present
along body margin. Cruciform pores few, present on margin.
Dorsum: Slide-mounted adult female with enlarged setae with sharp-pointed apex, form
a row along body margin, with 3 setae on margin of abdominal sternites I-VII, 49–55
µm long; the median sized setae 27–33 µm long, small setae, 11–17 µm long, form rows
on dorsum and along body margin, some setae on thorax 16–23 µm long. Macrotubular
ducts numerous all over dorsum 19 µm long and 8 µm wide. Microtubular ducts short,
with 2 sclerotized areas, scattered on entire dorsum. Anal ring dorsal, with 8 setae, each
100–129 µm long, partly with double row of pores. Cauda present (after Danzig (1962a),
with some modifications based on paralectotypes).
Ecology
Host plant: Agropyron sp., A. repens, Bromus sp., Hordeum sp.
Distribution: China, Iran, Russia, Tajikistan, Turkey, Uzbekistan.
Anophococcus kotejai Kozár & Kaydan Species nova (Fig. 83)
Holotype: Female. Greece (Voula), on Cynodon dactylon, F. Kozár, 30.vii.1983

(Kozár’collection number 2262a). Paratypes: 1 adult female, same data as holotype
(PPI: 2262a), in the same slide with holotype. Deposited in PPI.
Description
Unmounted female
Unknown.
Mounted female
232.5–235.0 µm, length of segments: I: 35–45, II: 35, III: 82.5–85.0, IV: 25–30, V: 22.5–
25.0, and VI: 32.5–35.0 µm long, the segments covered with a few, strong hair-like setae;
apical segment with apical seta 32.5–35.0 µm long; apical segment also with 3 sensory
falcate setae, 22.5–27.5 µm long; segment V with 1 sensory falcate seta 27.5–30.0 µm long,
segment IV with 1 sensory falcate seta 15.0–17.5 µm long. Frontal lobe absent, frontal
tubercle present. Eyes situated on venter near margin. Anal lobes strongly developed,
with 3 enlarged setae, 37.5–60.0 µm, plus 3-4 microtubular ducts on dorsal surface; apical
seta 400 µm; ventral hair-like subapical seta 75–95 µm long.
Venter: Labium 102.5–105.0 µm long, 85 µm wide, basal segment with two pairs of equal
long setae, with six pairs of setae on apical segment, median setae on apex of labium
12.5 µm long; stylet loop short. Legs well developed; lengths of segments and digitules
of prothoracic legs; coxa: 60–65, trochanter: 65, femur: 145.0–152.5, tibia: 107.5–120,
tarsus: 110.0–122.5 and claw: 32.5 µm long; tarsal digitules: 40.0–57.5, claw digitules:
35.0–37.5 µm long; lengths of segments and digitules of mesothoracic legs; coxa: 80–85,
trochanter: 60.0, femur: 145–150, tibia: 130.0–132.5, tarsus: 127.5–132.5, claw: 32.5–35.0
µm long, tarsal digitules: 52.5, claw digitules: 35.0–37.5 µm long; lengths of segments
and digitules of metathoracic legs; coxa: 90.0–95.0, trochanter: 60–65, femur: 150–160,
Acanthococcidae 223
Figure 82. Anophococcus kondarensis (Borchsenius, 1949), female. After Williams (1985)
with modifications.
tibia: 137.5–145.0, tarsus: 135.0–142.5 and claw: 35 µm long, tarsal digitules: 52.5–55.0,
claw digitules: 35.0–42.5 µm long; meso- and metathoracic coxae with spinulae on ventral
surface, and metathoracic coxae with a few pores on dorsal surface surface; tibiae with 5
setae (median seta present), tarsi with 5 setae. Length of spiracles 45.0–57.5 µm; diameter
of spiracular peritreme 22.5–30.0 µm, posterior spiracles slightly larger than anterior.
Multilocular pores 3–5 µm in diameter and with 3 or 5 loculi, distributed in sparse bands
on all abdominal segments and scattered on thorax and head; three locular pores mainly
on head. Macrotubular ducts of two sizes, smaller macrotubular ducts 2.5–4 µm wide and
15.0–17.5 µm long, present on median areas of abdominal segments, larger macrotubular
ducts 4.0–6.0 µm wide and 17.5–20.0 µm long, sparse in band on last abdominal segments,
and on submarginal band on abdomen, thorax and head. Microtubular ducts very small,
1.0–2.0 µm long, situated on marginal. Setae in median areas flagellate, long, 15–52.5
µm long. Enlarged setae, situated on submargin in a row, 17.5–37.5 µm long. Cruciform
pores on thorax and head on generally on submedian area in band on thorax and first for
abdominal segments and a few on head, 3.0–4.0 µm in diameter.
Dorsum: Marginal dorsal setae spine like, long, pointed, 35.0–62.5 µm forming a
marginal row and differentiated from dorsal setae, other dorsal setae, small, conical, seta
7.5–12.5 µm long; arranged in transverse rows across body segments, rows irregular on
head. Macrotubular ducts, 5.0–7.5 µm wide and 17.5–20.0 µm long, scattered throughout
dorsum, generally in segmental bands. Microtubular ducts very small, 1.0–2.0 µm long,
scattered over dorsum. Anal ring strongly sclerotized, with 14-19 pores on side, 65 µm in
diameter, with 8 setae, each 135–140 µm long; anal ring situated on margin of dorsum.
Cauda present.
Etymology
The new species is named after Professor Dr. Jan Koteja, who made great works on scale
insects.
Ecology
Host plant: Cynodon dactylon.
Distribution: Greece.
Biology: Unknown.
Comments
The adult female of An. kotejai differs from nearest species (An. cingulatus, An. confusus)
by absence of frontal lobes, from An. kondarensis by six segmented antennae and strong,
long marginal enlarged setae.
Acanthococcidae 225
Figure 83. Anophococcus kotejai Kozár & Kaydan sp. n., female.
Anophococcus lerzanae Kaydan & Kozár, Species nova (Fig. 84)
Holotype: Female. Turkey (Van-Gürpınar road), on Bromus sp., M. B. Kaydan,

24.vi.2010. Deposited in Kaydan’s collection (KPCT: 4712).
Description
Unmounted female
Unknown.
Mounted female
Body elongate oval, 3.54 mm long, 1.98 mm wide. Antennae 7 segmented, 250–270
µm, length of segments: I: 55, II: 30–32.5, III: 45, IV: 40–50, V: 27.5–32.5, VI: 27.5 and
VII: 40.0–45.0 µm long, the segments covered with a few, strong hair-like setae; apical
segment with apical seta 37.5 µm long; apical segment also with 3 sensory falcate setae,
30.0–32.5 µm long; segment VI with 1 sensory falcate seta 35.0–37.5 µm long, segment
V with 1 sensory falcate seta 22.5 µm long. Frontal lobe absent, frontal tubercle present.
Eyes situated on venter near margin. Anal lobes strongly developed, with 3 enlarged
setae, 40.0–62.5 µm plus 2 or 3 microtubular ducts on dorsal surface; apical seta 150 µm
long.
Venter: Labium 120 µm long, 85 µm wide, basal segment with two pairs of setae one
of them two times longer than others, with six pairs of setae on apical segment, median
setae on apex of labium 10 µm long; stylet loop short. Legs well developed; lengths
of segments and digitules of prothoracic legs; coxa: 70–75, trochanter: 60, femur: 135,
tibia: 110–120, tarsus: 125–135 and claw: 32.5–35.0 µm long, tarsal digitules: 35–40, claw
digitules: 32.5–35.0 µm long; lengths of segments and digitules of mesothoracic legs;
coxa: 85–105, trochanter: 55, femur: 130, tibia: 130, tarsus: 135–140, claw: 35.0–37.5
µm long, tarsal digitules: 55, claw digitules: 35.0–37.5 µm long; lengths of segments
and digitules of metathoracic legs; coxa: 100, trochanter: 55, femur: 140, tibia: 135,
tarsus: 130, claw: 35 µm long, tarsal digitules: 60, claw digitules: 35 µm long; meso- and
metathoracic coxae with spinulae on ventral surface, and metathoracic coxae with 25–27
pores on dorsal surface surface; tibiae with 5 setae (median seta present), tarsi with 6
setae. Length of spiracles 62.5–70.0 µm; diameter of spiracular peritreme 32.5–35.0 µm,
posterior spiracles slightly larger than anterior. Multilocular pores 5.0–7.5 µm in diameter
and with 5 or 7 loculi, double rimmed, distributed in sparse bands on all abdominal
segments and scattered on thorax and head. Macrotubular ducts of three sizes, smallest
macrotubular ducts 2.5–3 µm wide and 12.5–14.0 µm long, present on median areas of
abdominal segments II and III, medium size macrotubular ducts 10–12 µm wide and
20.0–22.5 µm long, sparse in band on last abdominal segments, thorax and head, largest
macrotubular ducts 12.5–15.0 µm wide and 30.0–32.0 µm long on submarginal band on
abdomen head and thorax. Microtubular ducts very small, 6.0–7.5 µm long, situated on
margin. Setae in median areas flagellate, long, 22.5–72.5 µm long. Enlarged setae, situated
Acanthococcidae 227
Figure 84. Anophococcus lerzanae Kaydan & Kozár sp. n., female.
on submargin in a row, 25.0–30.0 µm long. Cruciform pores on thorax and head on

generally on submedian area in band on thorax and first for abdominal segments and a
few on head, 4.0–5.0 µm in diameter.
Dorsum: Marginal dorsal setae spine like, long, pointed, 35–60 µm forming a marginal
row and differentiated from dorsal setae, other dorsal setae, conical, some of them
curved, seta 25.0–37.5 µm long; arranged in transverse rows across body segments,
rows irregular on head, last abdominal segment with 3 or 4 enlarged setae. Macrotubular
ducts, 12.5–15.0 µm wide and 30–32 µm long, scattered throughout dorsum, generally in
segmental bands. Microtubular ducts, 6.0–7.5 µm long, scattered over dorsum. Anal ring
strongly sclerotized, with 16 pores on side, 60 µm in diameter, with 8 setae, each 85–95
µm long; anal ring situated on margin of dorsum. Cauda present, small.
Etymology
The new species is named after Dr. Lerzan Erkılıç, who made great works on scale
insects in Turkey.
Ecology
Host plant: Bromus sp.
Biology: Unknown.
Anophococcus oblongus (Borchsenius, 1949) (Fig. 85) Combination nova

Rhizococcus oblongus Borchsenius 1949: 358.
Lectotype: Female. Tajikistan (Shaartuz District), on Cynodon dactylon leaves,

12.vi.1944, by N. Borchsenius (type no. 13-44). Deposited in ZMAS. Notes: Lectotype
to be designated by Danzig (1996). 2 paralectotype females on same slide as lectotype;
1 additional adult female paralectotype on 1 slide all in ZMAS.
Lit.: Bazarov, 1968b: 67; Borchsenius, 1949: 54; Danzig, 1962a: 840; Hoy, 1963:
104; Hua, 2000: 138, Matesova; 1968: 116; 1971: 27; Tao, 1999: 35; Wang, 2001: 225
Eriococcus oblongus; Hoy, 1963: 104. Change of combination.

Acanthococcus oblongus; Ter-Grigorian, 1983: 880. Change of combination.
Rhizococcus oblongus; Kozár & Walter, 1985: 75. Revived combination.
Acanthococcus oblongus; Miller & Gimpel, 1996: 283. Revived combination.
Rhizococcus oblongus; Köhler, 1998: 400. Revived combination.
Eriococcus oblongus; Miller & Gimpel, 2000: 283. Revived combination.
Rhizococcus oblongus; Kozár, 2009: 106. Revived combination.
Acanthococcidae 229
Figure 85. Anophococcus oblongus (Borchsenius, 1949), female. After Tang & Hua (1995)
with modifications.
Description
Unmounted female
Adult female elongate-oval, about 2.8 mm long, 1.2 mm wide. Ovisac gray, white or
yellowish, up to 4.0 mm long, 1.6 mm wide.
Mounted female
Antennae generally 7 rarely 6 segmented, 260–279 µm long, length of segment: I: 44.2,
II: 36, III: 65.5, IV: 54.7, V: 30.6, VI: 25.2, VII: 41.3 µm long. Frontal tubercle developed.
Anal lobe setae 310 µm long, subapical seta 80 µm long, suranal setae 60 µm long, anal
lobes with 3 blunted dorsal setae, up to 60 µm long.
Venter: Labium about 125 µm long, 80 µm wide, stylet loop 1.5 times as long as labium;
apical labial segment with 6 pairs of long setae, the median setae much shorter than
others. Legs medium sized, hind coxae and femur with a group of translucent pores and
spinulae; hind femur 210, tibia 170, tarsus 175 µm long, claw with denticle; tarsal and claw
digitules longer than claw, slightly knobbed. Quinquelocular pores numerous, present
over entire venter. Macrotubular ducts of two sizes, larger ones as those on dorsum,
narrower one 16 µm long and 4–6 µm wide, scattered all over the venter. Microtubular
ducts present on margin. Hair-like setae short. Enlarged setae of small size only, present
along body margin. Cruciform pores few, present on margin, some on midthorax.
Dorsum: Enlarged setae conical blunted, form a row along body margin, with 4 or 5 setae
on margin of abdominal sternites I-VII, 50–60 µm long, marginal setae conspicuously
larger than other setae on dorsal surface, the median sized setae 30–45 µm long, small
setae, 6–12 µm long, form rows on dorsum and along body margin, some setae on thorax
16–23 µm long. Macrotubular ducts in small number, in two rows on segments, about
12–16 on the last abdominal segments, 16–19 µm long and 8–11 µm wide. Microtubular
ducts scattered on entire dorsum. Anal ring dorsal, with 8 setae 135 µm long, partly with
double row of pores. Cauda present (after Borchsenius (1949), with some modifications
based on paralectotypes).
Ecology
Host plant: Agropyron sp., Avena fatua, Cissus sp., Cynodon dactylon, Festuca sp., Vitex
sampsoni.
Distribution: China, Tajikistan.
Comments
Danzig (1962a) mentioned the similarity of An. oblongus with An. insignis, and supposed
that it could be a subspecies. Here we treat it as a separate species on the base of very big
difference in number of macrotubular ducts on abdominal tergites.
Acanthococcidae 231
Anaphococcus oxyacanthus (Danzig, 1975) (Fig. 86) Redescription,

Combination nova
Acanthococcus oxyacanthus Danzig 1975: 80.
Holotype: Female. Russia (Vladivostok, Akademgorodok, railway embankment), on

Melilotus sp., 17.viii.1949, by N. Borchsenius. By original designation. Deposited in
ZMAS.
Lit.: Danzig, 1975: 43; 1980: 206; 1986: 241; 1988: 709), Kozár & Walter, 1985: 75;
Köhler, 1998: 400; Miller & Gimpel, 1999: 215; 2000: 290; Tang & Hao, 1995: 535
Rhizococcus oxyacanthus; Kozár & Walter, 1985: 75. Change of combination.

Eriococcus oxyacanthus; Miller & Gimpel, 1999: 214. Change of combination.
Rhizococcus oxyacanthus; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Female is oval, brown and the ovisac is grey, up to 2.5 mm long.
Mounted female
Adult female oval, 2.7 mm long, 1.7 mm wide. Antennae 7 segmented, length of
segments: I: 43, II: 48, III: 60, IV: 55, V: 24, VI: 20 and VII: 38 µm long; third and
fifth segments without hair-like setae, other segments with a few hair-like setae; apical
segment with apical seta; apical segment also with 3 sensory falcate setae, longest 34 µm
long; two preapical segments also falcate seta: on fifth segment 28 µm long, on sixth
segment 43 µm long. Frontal lobe absent, frontal tubercle 5 µm long. Eyes situated on
venter near margin. Width of anterior spiracles 43 µm. Anal lobes well developed, slightly
sclerotized, with 3 pointed conical spines, ventrally with three flagellate setae; anal lobe
with 240 µm long apical setae.
Venter: Labium 3 segmented, joint length of two apical segments 125 µm; basal
segment with 2 pairs of equally long setae, apical segment with 6 pairs of hair-like setae,
apical seta shorter, thicker. Legs well developed; length of segments of prothoracic legs,
coxa; 84, trochanter: 72, femur: 163, tibia: 113; tarsus: 112 and claw: 38 µm long, tarsal
digitules: 48, claw digitules: 40 µm long; length of segments of mesothoracic legs; coxa:
84, trochanter: 79, femur: 156, tibia: 120, tarsus: 124, claw: 38 µm long, claw digitules: 42
µm long; length of segments of metathoracic legs; coxa: 96, trochanter: 79, femur: 168,
tibia: 120, tarsus: 134; claw: 41 µm long, tarsal digitules: 55, claw digitules: 46 µm long,
tarsal and claw digitules slightly knobbed, longer than claw; all coxae with spinulae on
anterior surface, more abundant on metathoracic coxa; metathoracic coxae and femur
with large pores with cribriform surface on both anterior and posterior surfaces; claw
with a denticle; legs with a few hair-like setae, tibia with 5 setae. Quinquelocular pores
5 µm in diameter distributed in transverse bands across abdominal sternites, in sparse

rows on thoracic segments and head region. Macrotubular ducts of two sizes: larger
macrotubular ducts 7 µm wide medium long, situated on margin; smaller macrotubular
ducts 4 µm wide and medium long scattered throughout the venter. Microtubular ducts
few throughout the venter. Cruciform pores 5 µm wide; numerous in submarginal and
submedial areas on thoracic segments, and a few on submedial part of head. A few hair-
like setae present on submedian venter, some of them capitated. Enlarged setae pointed
in two sizes; larger 53 µm long on margin; smaller ones about half-size of marginal ones,
submarginally. Suranal setae hair-like.
Dorsum: Enlarged setae pointed, conical; longer setae, 91 µm long and medium sized
setae, 53 µm long, in a row on margins, 3-5 on segments; a few furcate setae and amoeboid
patches on margin might be aberrations; shorter setae few, 7–12 µm long, forming sparse
rows on dorsum, but absent from last abdominal segment; ten dorsal setae on abdominal
segment VII. Macrotubular ducts 7 µm wide and medium long, scattered among enlarged
setae. Microtubular ducts scattered, 7 µm long, elongate, with 2 sclerotized areas. Anal
ring situated on margin of dorsum; oval, sclerotized, 67 µm wide, 96 µm long, with one
row of pores and with 8 setae, each 130 µm long. Cauda plate-like, triangular shaped with
serrated distal margin (after Danzig (1975) with modifications. Drawing made based on
type material).
Ecology
Host plant: Melilotus sp.
Biology: On the roots. Ovisacs constructed from late July to mid August, but last stage
nymphs were still seen.
Acanthococcidae 233
Figure 86. Anophococcus oxyacanthus (Danzig 1975), female. Original.

Anophococcus pannonicus Kozár & Konczné Benedicty, Species nova

(Figs. 87, 88),
Holotype: Female (marked by red). Hungary, Fehérszék, 15.viii. 2002. Festuca sp. (PPI:
6538), with a paratype female on the same slide. Paratypes: 3 females on two slides,
2nd instar male nymph were also present, with the same data as holotype; Fehérszék,
10.vi. 2003, 5 females on two slides, from D-Vac (PPI: 6838); Fehérszék, 28.v. 2002,
female, Agropyron sp. (6379); Fehérszék, 28.v. 2002, female, and first instar nymph,
D-Vac (PPI: 6381); Fehérszék, 17.ix.2002, female, Cynodon sp. (PPI: 6572); Fehérszék,
22.v.2002, female, in leafsheets of Cynodon sp. (6616); Fehérszék, 20.iii.2003, 2
females, D-Vac (PPI: 6723); Orgovány, 21.x. 2002, female, D-Vac (PPI: 6663);
Jászboldogháza, 19.ix.2002, Agropyron sp., Cynodon sp., Festuca sp., female in eggsac on
the leaves (PPI: 6590, 6591, 6592); Kunszentmiklós, 03.ix.2002, female, Poaceae (PPI:
6563); Kunszentmiklós, 01.x.2002, female, Cynodon sp. (PPI: 6606); Kunszentmiklós,
19.xi. 2002, female, D-Vac (PPI: 6649); Kunszentmiklós, 03.vi.2003, 2 females, D-Vac
(PPI: 6834); Kunpeszér, 01.viii.2002, 12 females on five slides, 1st and 2nd instar female
and male nymph in 5 slides, Cynodon dactylon (PPI: 6488); Kunpeszér, 01.viii.2002, 2
females on a slide, D-Vac (PPI: 6494); Csobánc, 13.vi.2003, female D-Vac (PPI: 6839);
Érsekhalma, 01.ix.2006, female, Poaceae (PPI: 7625); Érsekhalma, 29.ix.2006, female,
D-Vac (PPI: 7648); Érsekhalma, 23.iv.2008, female, male second instar nymph, in the
leafsheets of Dactylis glomerata (PPI: 7625); Ecséd, 29.vii.2008, 4 females on two slides,
Agropyron sp., Bromus sp. (PPI: 8322, 8323); Ecséd (highway M3), 09.ix.2009, 4 females
on two slides, Bromus sp., (PPI: 9005); Töreki (highway M7), 09.ix.2009, 2 females,
Agropyron sp., (PPI: 8381); Tatabánya (Turul) (highway M7), 08.v.2009, 3 female on
two slides and a nymph, Artemisia sp., (PPI: 8593).
Description
Unmounted female
Unknown.
Mounted female
Adult female oval, 1.6–2.0 mm long, 0.7–1.2 mm wide. Antennae 6 (rarely 7) segmented,
length of segments: I: 31–40, II: 24–31, III: 77–98, IV: 20–25, V: 20–25 and VI: 36–
39 µm long; segments with a few hair-like setae; apical segment with 34–38 µm long
apical seta; apical segment also with 3 sensory falcate setae, longest 26–31 µm long; two
preapical segments also with 1 sensory falcate seta: on segment IV: 17–20, on segment V:
31 µm long. Frontal lobe absent, frontal tubercle 5 µm wide, 5 µm high. Eyes situated on
venter near margin. Width of anterior spiracles 29–34 µm. Cuticula with spinulae both
on dorsum and venter.
Venter: Labium 3 segmented, joint length of two apical segments 70–79 µm long; basal
segment with 2 pairs of short setae, apical segment with 6 pairs of equal length hair-like
Acanthococcidae 235
Figure 87. Anophococcus pannonicus Kozár & Konczné Benedicty sp. n., female, first instar
on top right.
setae, and 1 pair short apical setae; stylet loop longer than labium. Legs well developed:
length of segments of prothoracic legs; coxa: 40–58, trochanter: 38–50, femur: 115–139,
tibia: 84–101; tarsus: 103–115 and claw: 26–33, tarsal digitules: 43–50, claw digitules: 26–
35 µm long; length of segments of mesothoracic legs; coxa: 54–62, trochanter: 40–48,
femur: 110–134, tibia: 95–120, tarsus: 110–121, claw: 25–33 µm long, tarsal digitules:
46–50, claw digitules: 32–38 m long; length of segments of metathoracic legs; coxa:
66–79, trochanter: 45–55, femur: 128–144, tibia: 103–115, tarsus: 120–130; claw: 26–33
µm long, tarsal digitules: 46–52, claw digitules: 26–38 µm long, tarsal and claw digitules
slightly knobbed, longer than claw, meso- and metathoracic coxae with spinulae on
anterior surface; metathoracic coxae and femur also with small pores on both anterior
and posterior surfaces; claw with a denticle; legs with a few hair-like setae, and with one
sensory pore on tarsus; tibia with 5 setae. Multilocular pores 5–6 µm in diameter and with
5-9 loculi, mostly 7; distributed in transverse bands across abdominal sternites, in sparse
rows on thoracic segments and head region. Macrotubular ducts of two sizes: larger
macrotubular ducts 6 µm wide and 24 µm long, situated on margin; smaller macrotubular
ducts 4 µm wide and 20 µm long scattered throughout the venter. Microtubular ducts
sparse marginally and submarginally. Cruciform pores 6 µm long; a few in submarginal
area of thorax and throughout the head. A few hair-like setae present on submedian
venter, short hair-like setae submarginally and marginally. Suranal setae hair-like.
Dorsum: Enlarged setae of 3 kinds; pointed conical setae 32–39 µm long on anal lobe;
blunt conical spines of 3 sizes, longer setae 29–36 µm long, in a row on margin, 1 on
segment, medium sized setae 9–15 µm long, in a row on margin, 0-2 on each segment and
a few on dorsal segments, shorter setae 6–8 µm long, in a row on margin, 0 or 1 on each
segment and a few on each dorsal segment; slightly blunt conical spines each 5–11 µm
long, forming sparse rows on dorsum; no enlarged setae on abdominal tergite VIII; ten
dorsal setae on abdominal segment VII. Macrotubular ducts 6 µm wide and 24 µm long,
scattered among spines. Microtubular ducts scattered, each 6 µm long. Anal ring situated
on margin of dorsum; oval, sclerotized, 47–53 µm wide, 58–73 µm long, with one row of
pores and with 8 setae 79–94 µm long. Anal lobes well developed, membranouos, with 3
pointed conical spines, ventrally with two flagellate setae; anal lobe with 216–265 µm long
apical seta. Cauda wide, plate-like.
Other stages
Mounted L2 female (Fig. 88a)

Body oval, 1.1–1.2 mm long, 0.6 mm wide. Antennae 6 or 7 segmented, length of segments:
I: 10–25, II: 22–28, III: 60, IV: 23, V: 19–20, VI: 33–34 µm long; segments with a few hair-like
setae; apical segment with 29–31 µm long apical seta; apical segment also with 3 sensory falcate
setae, 21–23 µm long; two preapical segments also with 1 sensory falcate seta: on segment IV:
16, V: 26 µm long. Frontal lobe absent, frontal tubercle 3 µm wide. Eyes situated on venter near
margin. Width of anterior spiracles 16–19 µm. Cuticula with spinulae both on dorsum and
venter. Anal lobes well developed, membranouos, with 3 pointed enlarged setae, ventrally with
two flagellate setae; anal lobes with 190–210 µm long apical setae.
Acanthococcidae 237
segment with 2 pairs of setae, apical segment with 6 pairs of equal length hair-like setae,
and 1 pair short apical setae; stylet loop longer than labium. Legs well developed: length
of segments of prothoracic legs; coxa: 30–32, trochanter: 30–33, femur: 95–99, tibia:
60–62; tarsus: 77–80, claw: 23–24 µm long, tarsal digitules: 36 µm long, claw digitules:
24–26 µm long; length of segments of mesothoracic legs; coxa: 33–35, trochanter: 29,
femur: 97–102, tibia: 72–74, tarsus: 88–90, claw: 24–25 µm long, tarsal digitules: 33–34,
claw digitules: 24–26 m long; length of segments of metathoracic legs; coxa: 40–44,
trochanter 37–38, femur: 100, tibia: 70–74, tarsus: 96–98; claw: 25–28 µm long, tarsal
digitules: 38–39, claw digitules: 27–28 µm long, tarsal and claw digitules slightly knobbed,
longer than claw; meso- and metathoracic coxae with spinulae on anterior surface; claw
with a denticle; legs with a few hair-like setae; tibia with 5 setae. Multilocular pores 5–6
µm in diameter with 5-7 loculi, mostly 5; few on the whole venter. Microtubular ducts
sparse marginally and submarginally. Cruciform pores 4 µm long; a few in marginal
area of thorax. A few hair-like setae present on submedian venter, short hair-like setae
submarginally and marginally. Suranal setae hair-like.
Figure 88. Anophococcus pannonicus Kozár & Konczné Benedicty sp. n., second instar: a
female, b male.
Dorsum: Cuticular surface with nodulations. Enlarged setae pointed, conical; longest setae
28 µm long, in a row on margin, 1 on each sement, and a group on head margin, shorter
setae 5–8 µm long, in a row on margin, no enlarged setae on abdominal tergite VIII. Four
dorsal setae on abdominal segment VII. Macrotubular ducts absent. Microtubular ducts
scattered, 3 µm long. Anal ring situated on margin of dorsum; oval, sclerotized, 42–48
µm wide, 41–42 µm long, with one row of pores and with 6 setae, each 68 µm long.
Cauda wide, plate-like.
Mounted L2 male (Fig. 88b)

Body oval, 0.9–1.1mm long, 0.4–0.5 mm wide. Antennae 7 segmented, length of segments:
I: 20–28, II: 18–22, III: 25–30, IV: 22–25, V: 16–20, VI: 18–20 and VII: 28–32 µm long;
segments with a few hair-like setae; apical segment with 24–29 µm long apical seta; apical
segment also with 3 sensory falcate setae, 20–23 µm long; two preapical segments also
with 1 sensory falcate seta: on fifth segment 11–16, on sixth segment 21–24 µm long.
Frontal lobe absent, frontal tubercle 3 µm wide. Eyes situated on venter near margin.
Width of anterior spiracles 15–17 µm. Cuticula with spinulae both on dorsum and venter.
Mounted L1 (Fig. 87, top right)

Body oval. Antennae 6 segmented, all segments with a few hair-like setae; apical segment
also with 3 sensory falcate setae, two preapical segments also with 1 sensory falcate
seta. Frontal lobe absent, frontal tubercle 3 µm wide. Eyes situated on venter near
margin. Cuticula with spinulae both on dorsum and venter. Anal lobes well developed,
membranouos, with 3 pointed conical spines, ventrally with two flagellate setae; and a
long apical seta.
apical segment with 6 pairs of equal length hair-like setae, and 1 pair short apical setae,
stylet loop longer than labium. Legs normal: tarsal and claw digitules slightly knobbed,
longer than claw; meso- and metathoracic coxae with spinulae on anterior surface; claw
with a denticle; legs with a few hair-like setae; tibia with 5 setae. Multilocular pores with
5-7 loculi, mostly 5; few on the whole venter. Microtubular ducts sparse marginally and
submarginally. Cruciform pores; a few in marginal area of thorax. A few hair-like setae
present on submedian venter, short hair-like setae submarginally and marginally. Suranal
setae hair-like.
Dorsum: Cuticular surface with nodulations. Enlarged setae pointed, conical; in a row on
margin, 1 on segments, and a group on head margin, shorter setae in a row on margin, no
enlarged setae on abdominal tergite VIII. Four dorsal setae on abdominal segment VII.
Microtubular ducts scattered. Anal ring situated on margin of dorsum; oval, sclerotized,
with one row of pores and with 6 setae. Cauda wide, plate-like.
Acanthococcidae 239
Etymology
The species named after the province Pannonia of the Roman Empire, of which territory
is now Hungary.
Ecology
Host plant: Agropyron sp., Bromus sp. Cynodon sp., Dactylis glomerata, Artemisia sp.
Distribution: Hungary.
Biology: Females were found from April till October, probably it has two generations.
Comments
An. pannonicus differs from all species included in Anophococcus genus by having only one
enlarged spine on margin of segments.
Anophococcus parvispinus (Goux, 1948) (Fig. 89) Redescription

Eriococcus parvispinus Goux, 1948: 5.
Holotype: Female. France (Camargue, Riéges). By original designation. On Artemisia

gallica, 26.vi.1946, by L. Goux. Paratypes: 31 slides are with the same data as holotype
(MNHN 14498-02-32). Deposited in MNHN.
Lit.: Balachowsky, 1954: 61; 1989a: 21; Foldi, 2001:305; 2002: 245; Goux, 1948: 5;
1948: 67; Hoy, 1963: 107; Koteja & Zak-Ogaza, 1981: 512; Kozár & Walter, 1985: 75;
Lindinger, 1957: 550; Ouvrard & Kozár, 2009: 110 (Miller et al., 2013).
Anophococcus parvispinus; Balachowsky, 1954: 61, 64. Change of combination.

Nidularia parviseta; Lindinger, 1957: 550. Change of combination and replacement
name for Eriococcus parvispinus; Notes: Eriococcus parvispinus Goux 1948 is preoccupied
by Eriococcus parvispinus Chaffin 1923. Lindinger (1957) proposed Nidularia parvisetus as a
replacement name.
Eriococcus parvisetus; Hoy, 1963: 107. Change of combination.
Greenisca parvispinus; Kozár & Walter, 1985: 75. Reestablishment a change of combination.
Acanthococcus parvisetus; Miller & Gimpel, 1996: 603. Change of combination.
Greenisca parvispinus; Köhler, 1998: 390. Revived combination.
Eriococcus parvisetus; Miller & Gimpel, 2000: 296. Revived combination.
Anophococcus parvispinus; Kozár, 2009: 94; Revived combination.
Description
Unmounted female
Adult female at time of sac formation is reddish orange and varies throughout the life
cycle.
Mounted female
Body elongate oval. 1.82–2.05 mm long, 1.07–1.27 mm wide. Antennae 7 segmented,
sometimes on one antennae of a specimen third and fourth parallel sided; length of
segments: I: 29–49, II: 20–32, III: 58–76, IV: 48–58, V: 26–30 and VI: 20–26 µm long;
apical setae of antenna 41–55 µm; on apical segment three sensory falcate setae are
found, the longest 26–29 µm long; on the preapical segment the falcate sensory seta
20–35 µm long; the segments of the antenna are covered with few hair-like setae. Frontal
lobes present. Eyes visible, situated on venter. Anal lobes slightly sclerotized, not fused
at the base, with two setose spines along inner margin, the posterior one longer, and
one much shorter spine on base of the outer margin, apical seta 320–380 µm long. The
subapical seta spinose, situated near to base of apical seta.
Venter: Labium 3 segmented, 142–160 µm long; stylet loop long reaching the posterior
legs. Legs well developed; length of segments of prothoracic legs; coxa: 55–77, trochanter:
41–62, femur: 131–149, tibia: 100–112, tarsus: 121–134, claw: 34–42 µm long; length of
segments of mesothoracic legs; coxa: 65–75, trochanter: 45–65, femur: 128–139, tibia:
112–125, tarsus: 123–135, claw: 33–38 µm long; length of segments of metathoracic
legs: coxa: 71–95, trochanter: 40–67, femur: 135–144, tibia: 118–129, tarsus: 134–170,
tarsal digitules knobbed, 45–52, claw 30–42 µm, claw digitules, 32–41 µm long, slightly
knobbed; posterior coxae with some longitudinal spinulae and small pores; claw with
denticle; legs with few hair-like setae. The diameter of anterior spiracles 36–39 µm.
Quinquelocular pores found around spiracles, and band on last abdominal segments
and scattered elsewhere, 5–6 µm in diameter. Macrotubular ducts about 3–7 µm wide
and 20–22 µm long, scattered throught. Microtubular duct present, scattered on surface.
Cruciform pores present in a submarginal band. Venter with a small number of scattered,
short hair-like setae.
Dorsum: Enlarged setae few, 4–9 µm long; on body margin row of spines absent.
Macrotubular ducts similar to those on venter, sparse, present throughout, 7–8 µm wide
and 24-26 µm long. Microtubular ducts 4–5 µm long, scattered throughout. Anal ring
64–72 µm wide, 72–86 µm long, with pores and with 8 hair-like setae, 110–140 µm long.
Cauda not seen (Redescription based on type materials).
Other stages
Detailed description and illustrations of all stages, including the winged male, by Goux
(1948).
Acanthococcidae 241
Figure 89. Anophococcus parvispinus (Goux, 1948), female. Original.

Ecology
Host plant: Artemisia gallica.
Biology: In June all stages were found.
Comments
While most of the related species are living on different grasses, this species was collected
from Artemisia sp.
Anophococcus pseudinsignis (Green, 1921) (Fig. 90) Revived combination

Eriococcus pseudinsignis Green, 1921, 149.
Lectotype: United Kingdom (England, Kent, Thurnham), on Festuca?, ?.ix.1920.

female, by subsequent designation Williams, 1985a: 378-380. Deposited in BMNH.
Notes: In addition to the lectotype adult female, there are two paralectotype adult
females on the same slide in the BMNH.
Common name: Boreal felt scale.
Lit.: Boratynsky & Williams, 1964: 92; Borchsenius, 1949: 354; Danzig 1962: 24;
1971a: 823; Dziedzicka & Koteja, 1971: 569; Goux, 1940: 65; Green, 1921: 149;
Hodgson, 2005: 39; Kaydan et al.; 2005: 400; Kiritchenko, 1940: 135; Kosztarab &
Kozár, 1978: 68. Koteja, 1971: 322; 1974b: 76; 1976: 272; 1983: 675; Koteja & Zak-
Ogaza, 1969: 364; 1983: 477; Kozár et al., 2013: 55; Ouvrard & Kozár, 2009. 102;
Schmutterer, 1952: 68; Tang & Hao, 1995: 520; Tereznikova, 1981: 35; Wang, 1982a:
442; Williams, 1985a: 378; Zahradník, 1959: 540; 1977: 121 (Miller et al., 2013).
Rhizococcus pseudinsignis; Borchsenius, 1949: 354. Change of combination.

Nidularia pseudinsignis; Lindinger, 1933a: 116. Change of combination.
Eriococcus pseudinsignis; Hoy, 1963: 110. Change of combinaítion.
Acanthococcus pseudinsignis; Tereznikova, 1975. Change of combination.
Rhizococcus pseudinsignis; Kozár & Walter, 1985: 75. Revived combination.
Anophococcus pseudinsignis; Koteja, 1990. Change of combination.
Acanthococcus pseudinsignis; Miller & Gimpel, 1996. Revived combination.
Rhizococcus pseudinsignis; Köhler, 1998: 400. Revived combination.
Eriococcus pseudinsignis; Miller & Gimpel, 2000: 307. Revived combination.
Rhizococcus pseudinsignis; Kozár, 2009: 106. Revived combination.
Acanthococcidae 243
Description
Unmounted female
Adult female at time of sac formation is reddish orange and varies throughout the life
cycle. Ovisacs are elongate-oval, white to light yellow and usually placed at the junction
of the blade with the stem of the grass, where they are practically concealed from view
(Green, 1921).
Mounted female
Adult female is elongate, rather narrow, yellow. Slide-mounted specimens, elongate-oval,
the largest 2.8–3 mm long, 1.5 mm wide, nodulose. Anal lobes conical, about twice as
long as wide, sclerotised. Each lobe with an apical seta 280 µm long; dorsally 1 outer and
2 inner enlarged setae and ventrally 2 slender setae and a suranal seta shorter than anal
ring setae.
Venter: Labium 100–120 µm long, shorter than clypeolabral shield, basal segment with
2 pairs of setae. Stylet loop twice longer than labium. Antennae 180–300 µm long with 7
segments. On apical segment three sensory falcate setae are found, the longest 26–29 µm
long. On the preapical segment the falcate sensory seta 20–35 m long. The segments of
the antenna are covered with few hair-like setae. Frontal tubercle present just anterior to
basal segment. Legs well developed, slender, hind trochanter + femur 230–260 µm long,
hind tibia 130–150 µm long, hind tarsus 150–160 µm long, claw 35 µm long, curved, with
a small denticle near apex. Coxa with a few translucent pores on outer half. Hind coxa
with a large group of translucent pores; hind leg 720 µm long, hind femur 138–187, tibia
130–160, tarsus 135–160 µm long.
Dorsum: Dorsal setae few about three on each margial segment, 28–60 µm long.
Enlarged setae conical, sides slightly concave, apices slightly rounded anteriorly, rounded
or truncate posteriorly, setae of 2 sizes, marginal setae largest, smaller setae scattered
over remainder of dorsum, 3 or 4 lateral setae on margin of each abdominal segment;
ventral multilocular pores predominantly with 7 loculi; microtubular ducts short, with
2 sclerotized areas. Cauda sclerotised, narrow, rounded and slightly nodulose. Anal ring
with 8 setae, each about 140 µm long (after Williams, 1985a).
Other stages
First instar described by Schmutterer (1952).
Ecology
Host plant: Achillea sp., A. millefolium, Agropyron sp., A. ponticum, A. repens, Agrostis
capillaris, Alopecurus sp., A. pratensis, Ammophila sp., Andropogon sp., Anthoxanthum sp.,
Arrhenaterum elatius, Brachypodium pinnatum, B. sylvaticum, Bromus sp., B. inermis, Calamagrostis
sp., Carex sp., Cynodon dactylon, Deschampsia flexuosa, Dianthus crinitus, Festuca sp., Holcus sp.,
H. lanatus, H. mollis, Koeleria glauca, Luzula sp., Phragmites sp., Setaria sp., Taraxacum sp.,
Veronica sp., V. chamaedrys.
Distribution: Austria, Bulgaria, former Czechoslovakia, France, Germany, Hungary,

Italy, Poland, Slovenia, Sweden, Tunisia, Turkey, Ukraine, United Kingdom, former
Yugoslavia (Slovenia).
Biology: This species has one generation, eggs overwinter. Adults appear during the first
part of June, start laying 29-78 eggs per female from the end of June.
Primarily on grasses; infests leaves of the plants in the host genera. According to
Schmutterer (1952) it has one generation; eggs overwinter; adults appear during first part
of June, start laying 29-87 eggs per female from end of June. Dziedzicka & Koteja (1971)
collected adult females from April through September. Green (1921) found most ovisacs
concealed at base of upper surface of leaves.
Comments
Green's original slide contains three specimens, one of which is selected as lectotype and
clearly marked, the other two are labelled paralectotypes.
It is sometimes difficult to distinguish this species from An. insignis. At present
the main differences are the 3 marginal setae on the abdominal segment VII in An.
pseudinsignis compared with 4 in An. insignis. The dorsal setae on the head and thorax
in An. pseudinsignis are quite robust and are much longer than the setae on the posterior
abdominal setae, whereas all the dorsal setae in An. insignis are short and usually slender,
although those on the head and thorax are often wider than the abdominal setae. At present
both species are here regarded as distinct, but intermediates may be found to warrant
further research.
Acanthococcidae 245
Figure 90. Anophococcus pseudinsignis (Green, 1921), female. After Williams (1995), with
modifications.
Anophococcus salsolae (Borchsenius, 1949) (Fig. 91) Revived combination

Rhizococcus salsolae Borchsenius, 1949: 363.
Lectotype: Female. Tajikistan (Shaartuz District, Kuyuk Ridge), to be designated

by Danzig (1996). Designated by Danzig & Gavrilov-Zimin (2011: 272). 1 female
paralectotype on same slide as lectotype, additional paralectotype females on the
other slide. Deposited in ZMAS. Notes: Paralectotype were studied.
Lit.: Balachowsky, 1954: 61; Borchsenius, 1949: 53; Danzig, 1962a: 841; Danzig &
Gavrilov-Zimin, 2011: 272; Hoy, 1963: 114; Köhler, 1998: 401; Koteja & Zak-Ogaza,
1981: 512; Kozár, 2009: 94; Kozár & Walter, 1985: 75; Miller & Gimpel, 1996: 604;
2000: 327; Myartseva, 1982: 41; Tang & Hao, 1995: 519 (Miller et al., 2013).
Anophococcus salsolae; Balachowsky, 1954: 61. Change of combination.

Eriococcus salsolae; Hoy, 1963: 114. Change of combination.
Rhizococcus salsolae; Kozár & Walter, 1985: 75. Revived combination.
Acanthococcus salsolae; Miller & Gimpel, 1996: 604. Change of combination.
Rhizococcus salsolae; Köhler, 1998: 401. Revived combination.
Eriococcus salsolae; Miller & Gimpel, 2000: 327. Revived combination.
Anophococcus salsolae; Kozár, 2009: 94. Revived combination.
Description
Unmounted female
Adult female is oval. Sac is greyish or white, 3.0 mm long, 1.8 mm wide.
Mounted female
Slide-mounted adult female, 2.4 mm long, 1.3 mm wide. Antennae 7 segmented, I: 43,
II: 22, III: 35, IV: 43, V: 20, VI: 17 and VII: 28 µm long; third segment without hair-like
setae, other segments with a few hair-like setae. Frontal lobe present. Eyes situated on
venter near margin. Anal lobes well developed, sclerotized, with 3 pointed, conical spines,
24–27 µm long, ventrally with two flagellate setae; anal lobe with 140 µm long apical and
with 60 µm long subapical setae.
Venter: Labium 3 segmented, narrow, basal segment with 2 pairs of equal long setae,
apical segment with 5 pairs of hair-like setae, apical seta shotrer, thicker. Stylet loop about
one third longer than labium. Legs normal: metathoracic legs: femur 180 µm, tibia 125
µm, tarsus 165 µm; tarsal and claw digitules not knobbed, longer than claw. Posterior
coxae with spinulae on anterior surfaceand with pores. Claw with a denticle. Legs with a
few hair-like setae, and with one sensory pore on tarsus; tibia with 4 setae. Multilocular
pores with 3-5 loculi; distributed in sparse transverse bands across abdominal sternites,
few on thoracic segments and head region. Macrotubular ducts of two sizes: larger
macrotubular ducts situated on margin 5 µm wide and 20 µm long; smaller macrotubular
ducts scattered throughout the venter. Microtubular ducts around margin. Cruciform
Acanthococcidae 247
Figure 91. Anophococcus salsolae (Borchsenius, 1949), female. After Danzig (1962), with
modifications.
pores absent. A few hair-like setae present on submedian venter. Pointed conical spines
in two sizes on submargin. Suranal setae hair-like 60 µm long.
Dorsum: Enlarged setae absent. Anal ring situated on margin of dorsum; oval, sclerotized,
67 µm wide, 96 µm long, with one row of pores and with 6 setae 50 µm long. Cauda
not seen (after Borchsenius (1949); Danzig (1962a), with modifications based on type
material).
Ecology
Host plant: Salsola sp.
Distribution: Tajikistan.
Biology: On the branches, and roots.
Anophococcus sanguinairensis (Goux, 1993) (Fig. 92) Redescription,

Combination nova
Eriococcus sanguinairensis Goux, 1993: 68.
Holotype: Female. (MNHN 14510-01). France (Corse, Ajaccio rte des Sanguinaires).
Paratypes: 5 slides are with the same data as holotype.
Lit.: Foldi, 2001: 305; Goux, 1993: 68; Kozár, 2009: 106; Miller & Gimpel, 1996: 604;
2000: 327; Ouvrard & Kozár, 2009: 102 (Miller et al., 2013).
Acanthococcus sanguinairensis; Miller & Gimpel, 1996: 604. Change of combination.

Rhizococcus sanguinairensis; (Goux, 1993). Kozár (2009): 106. Combination nova.
Description
Unmounted female
Two mm long, 1.2 mm wide.
Mounted female
the size of the segments: I: 40–43, II: 30–38, III: 81–101, IV: 21–29, V: 19–25 and VI:
30–40 µm long; the third segment is almost parallel sided; apical setae of antenna 36–40
µm; on apical segment three sensory falcate setae are found, the longest 25–33 µm long;
on the preapical segment the falcate sensory seta 27–31 µm long; the segments of the
antenna are covered with few hair-like setae. Frontal lobes, or tubercles not seen. Eyes
visible, situated on venter. Anal lobes slightly sclerotized with two spinose setae along
inner margin, and one spinose seta on outer margin, similar in size to those on margin,
apical seta 180–210 µm long.
Acanthococcidae 249
Figure 92. Anophococcus sanguinairensis (Goux, 1993), female. Original.

Venter: Labium apparently 3 segmented, 45–82 µm long, basal segment with two pairs
of equal long setae; stylet loop long as long as the labium. Legs well developed: length
of segments of prothoracic legs; coxa: 34–63, trochanter: 40–53, femur: 130–145, tibia:
92–118, tarsus: 102–125, claw: 25–30 µm long. length of segments of mesothoracic legs;
coxa: 45–67, trochanter: 32–62, femur: 135–165, tibia: 115–128, tarsus: 105–126, claw:
30–35 µm long; length of segments of metathoracic legs; coxa: 61–86, trochanter: 45–59,
femur: 115–146, tibia: 122–128, tarsus: 126–138, claw: 32–36 µm long, tarsal digitules
knobbed, 40–51 µm long, claw digitules, 31–40 µm long, slightly knobbed; posterior
coxae without spinulae, with medium number of large pores; claw with denticle; legs
with few hair-like setae, tibia with 5 setae. Multilocular pores with 5-7 loculi, mostly 7,
distributed in sparse bands and rows on all segments of abdomen and thorax, 5–7 µm
in diameter. The diameter of anterior spiracles 30–36 µm. Macrotubular ducts about
3–5 µm wide and 17–20 µm long, scattered. Microtubular duct present in a submarginal
band. Venter with a small number of scattered, hair-like setae and a submarginal row of
enlarged setae. Few submarginal cruciform pores present.
Dorsum: Enlarged setae cylindrical, sides concave basally, apices truncate posteriorly and
rounded anteriorly, marginal setae conspicuously larger than all other dorsal setae Dorsal
setae few, 6–7 µm long. On margin of segments three spines present, 31–43 µm long.
and 23–25 µm long. Microtubular ducts 4–5 µm long, scattered throughout. Anal ring
50–56 µm wide, 45–69 µm long, with pores and with 8 hair-like setae, about 100 µm long.
Cauda not seen. (Redescription based on type materials).
Ecology
Host plant: Poaceae.
Distribution: France, Iran, Turkey.
Comments
According to drawings and descriptions this species is similar to R. pseudinsignis (Green,
1921), and needs further studies.
Turkey is a new record of distribution of this species.
Acanthococcidae 251
Anophococcus selmae Kaydan & Kozár, Species nova (Fig. 93)
Holotype: Female. Turkey (Erzincan-Tercan road), on Phalaris sp., N: 39°38’384’’,

E: 39°47’864’’, M. B. Kaydan, F. Kozár, 08.vii.2010. Deposited in Kaydan’s collection
(KPCT: 4790).
Description
Unmounted female
Unknown.
Mounted female
Body elongate oval, 1.82 mm long, 0.80 mm wide. Antennae 6 segmented, 250–260
µm, length of segments: I: 40, II: 35, III: 90, IV: 27.5, V: 25 and VI: 37.5 µm long, the
segments covered with a few, strong hair-like setae; apical segment with apical seta 25
µm long; apical segment also with 3 sensory falcate setae, 15–25 µm long; segment V
with 1 sensory falcate seta 22.5–25.0 µm long, segment IV with 1 sensory falcate seta
12.5–15.0 µm long. Frontal lobe absent, frontal tubercle present. Eyes situated on venter
near margin. Anal lobes strongly developed, with 3 enlarged setae, 25.0–27.5 µm plus 2
or 3 microtubular ducts on dorsal surface; apical seta 195 µm long.
Venter: Labium 95 µm long, 100 µm wide, basal segment with two pairs of setae one of
them two times longer than others, with six pairs of setae on apical segment, median setae
on apex of labium 5 µm long; stylet loop short. Legs well developed; lengths of segments
and digitules of prothoracic legs; coxa: 62.5–75.0, trochanter: 50–60, femur: 122.5–135.0,
tibia: 112.5, tarsus: 105 and claw: 22.5–25.0 µm long, tarsal digitules: 35.0–42.5, claw
digitules: 30 µm long; lengths of segments and digitules of mesothoracic legs; coxa: 70.0–
82.5, trochanter: 60–65, femur: 130–135, tibia: 120–125, tarsus: 115–125, claw: 25.0–30
µm long, tarsal digitules: 45, claw digitules: 25 µm long; lengths of segments and digitules
of metathoracic legs; coxa: 85–90, trochanter: 55–60, femur: 135, tibia: 125, tarsus: 120,
claw: 20.0–27.5 µm long, tarsal digitules: 35–40 µm long, claw digitules: 22.5 µm long;
meso- and metathoracic coxae with spinulae on ventral surface, and metathoracic coxae
with 6–8 pores on dorsal surface surface; tibiae with 5 setae (median seta present), tarsi
with 5 setae. Length of spiracles 55–60 µm; diameter of spiracular peritreme 22.5–
30.0 µm, posterior spiracles slightly larger than anterior. Multilocular pores 5–6 µm in
diameter and with 5 or 7 loculi, distributed in sparse bands on all abdominal segments
and scattered on thorax and head. Macrotubular ducts of two sizes, smaller macrotubular
ducts 5–6 µm wide and 12.5–14.0 µm long, sparse in band on last abdominal segments,
thorax and head, larger macrotubular ducts 7–8 µm wide and 15.0–17.5 µm long on
submarginal band on abdomen head and thorax. Microtubular ducts very small, 2–3 µm
long, situated on margin. Setae in median areas flagellate, long, 12.5–50.0 µm long, setae
on submargin in a longitudinal band, 10.0–17.5 µm long. Cruciform pores on thorax and
head on generally on submedian area in band on thorax and first for abdominal segments
and a few on head, 3–4 µm in diameter.
Dorsum: Marginal dorsal enlarged setae wide at the base, pointed, 20.0–27.5 µm long
on abdominal segments, spine like, longer than abdominal marginal setae, 35.0-37.5 µm
long on thorax and head, forming a marginal row and differentiated from dorsal setae,
other dorsal setae, conical, spine like (A. cingulatus type), seta 7–8 µm long; arranged in
transverse rows across body segments, rows irregular on head. Macrotubular ducts, 7–8
µm wide and 15–18 µm long, scattered throughout dorsum, generally in segmental bands.
Microtubular ducts, 3–4 µm long, scattered over dorsum. Anal ring strongly sclerotized,
60 µm in diameter, with 8 setae, all setae are broken; anal ring situated on margin of
dorsum. Cauda present, small.
Etymology
The new species is named after Prof. Dr. Selma Ülgentürk, who made great works on
scale insects in Turkey.
Ecology
Host plant: Phalaris sp.
Biology: Unknown.
Anophococcus socialis (Goux, 1931) (Fig. 94) Redescription, Combination

nova
Eriococcus socialis Goux, 1931: 64.
Lectotype: Female (MNHN 4951-26). France (Var, La Seyne, Tamaris-sur-Mer).

Designated here. Paralectotypes: 29 slides are with the same data as lectotype (MNHN
4951-02-30). Deposited in MNHN. Notes: In the syntype series there are 6 slides
MNHN 14513-01-06, same data as holotype, but collected in 09. VIII. 1930, one
of them MNHN 14513-03 contains one female of R. pseudinsignis and some larval
stages, other slides from this series contains only nymphs, so these six slides were not
included into the paralectotype series.
Lit.: Balachowky, 1934a: 38; 1934b: 101; Foldi, 2001: 305; Goux, 1931: 63; 1933: 116;
1940: 65; 1944: 137; Hoy, 1963: 116; Kozár, 2009: 106; Lindinger, 1936: 156; Miller
& Gimpell, 1996: 604; Miller & Gimpell, 2000: 336; Ouvrard & Kozár, 2009: 102
Eriococcus insignis; Newstead, 1891, Lindinger (1936) erroneously considered Eriococcus

socialis as a synonym.
Acanthococcus socialis; Miller & Gimpel, 1996: 604. Change of combination.
Acanthococcidae 253
Figure 93. Anophococcus selmae Kaydan & Kozár sp. n., female.
Rhizococcus socialis; Köhler, 1998:399. Change of combination, as synonym of An. insignis.

Eriococcus socialis; Miller & Gimpel, 2000: 336. Revived combination.
Rhizococcus socialis; Kozár, 2009: 106. Revived combination.
Eriococcus franceschinii; (Balachowsky, 1934b: 98). Synonym nova.
Lectotype: France (Corsica, Restonica, Aitone forest). Designated here. Deposited
in MNHN. Notes: The slides under this name are 4940-1 (6-8 1933), 4940-2 (8 1933),
4940-3 (8 1933), 4940-4 (8 1933, 1000 m), 4940-6 (8 1933, 1200 m), 4940-5 (20
8 1933) on Gramineae. They contain two species, five slides (4940-2-6) of them
belong to Anophococcus pseudinsignis, and the only female specimen on 4940-1 slide
belongs to An. francenschini. The original drawing of Balachowsky (1934b) presents
this specimen. Study of characters of this specimens did not show any important
difference from An. socialis (Goux, 1931) and considered here as Synonym nova.
Nidularia franceschinii; Lindinger, 1936: 156. Change of combination.
Rhizococcus franceschinii; Kozár & Walter, 1985: 75. Change of combination.
Acanthococcus franceschinii; Miller & Gimpel, 1996: 601. Change of combination.
Rhizococcus franceschinii; Köhler, 1998: 399. Revived combination.
Eriococcus franceschinii; Miller & Gimpel, 2000: 336. Revived combination.
Rhizococcus franceschinii; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Ovisac is white to cream in color 4 mm long 2-3 mm wide. First instar is yellow and oval.
Adult female is reddish, ovoid.
Mounted female
Body elongate oval. 1.22– 2.24 mm long, 0.62– 1.33 mm wide. Antennae 6 segmented,
the size of the segments: I: 27–36, II: 29–33, III: 83–101, IV: 23–31, V: 22–25, and VI:
32–41 µm. Apical setae of antenna 34–38 µm. On apical segment three sensory falcate
setae are found, the longest 20– 29 µm long. On the preapical segment the falcate sensory
seta 25–31 µm long. The segments of the antenna are covered with few hair-like setae.
Frontal lobes absent. Eyes visible, situated on venter. Anal lobes slightly sclerotized with
two spinose setae along inner margin, and one spinose seta on outer margin, similar in
size to those on margin, apical seta 165–240 µm long.
Venter: Labium 3 segmented, 66–73 µm long basal segment with two pairs of equal long
setae. Stylet loop as long as the labium. Lengths of segments of metathoracic legs; coxa:
48–58, trochanter: 39–42, femur: 118– 130, tibia: 90–105, tarsus: 104–116, claw, 27–35 µm
long; lengths of segments of mesothoracic legs; coxa: 52– 60, trochanter: 35– 40, femur:
128–135, tibia: 95–118, tarsus: 113–120, claw: 27–32 µm long; lengths of segments of
prothoracic legs; coxa: 65–71, trochanter: 35–40, femur: 135–141, tibia:108–132, tarsus:
128–141, tarsal digitules knobbed, 40–46 µm long, claw 26–32 µm long, claw digitules,
29–35 µm long, slightly knobbed. Posterior coxae without spinulae, with medium number
Acanthococcidae 255
Figure 94. Anophococcus socialis (Goux, 1931), female. Original.

of large pores. Claw with denticle. Legs with few hair-like setae. Multilocular pores with 7
or 8 loculi, distributed in sparse bands and rows on all segments of abdomen and thorax,
5–6 µm in diameter. The diameter of anterior spiracles 26–32 µm. Venter with a small
number of scattered, hair-like setae. Microtubular ducts scattered. Macrotubular ducts
about 4–5 µm wide and 13–14 µm long, scattered. Cruciform pores not found.
Dorsum: Enlarged setae conical, sides straight, apices acute, 2 sizes of setae, larger setae
present on margin, smaller setae scattered over rest of dorsum few, 7–12 µm long, with
blunted apex. On margin of segments four truncated spines present, two of them longer
29–35 µm long, two shorter 15–17 µm long. Macrotubular ducts similar to those on
venter, sparse, present throughout, 8–9 µm wide and 24–27 µm long. Microtubular ducts
3–5 µm long, scattered throughout. Anal ring 40–53 µm wide, 50–60 µm long, with pores
and with 8 hair-like setae, 73–88 µm long. Cauda present (Redescription based on type
materials).
Other stages
Male shortly described by Goux (1931).
Ecology
Comments
According to drawings and descriptions this species is very similar to An. formicicola but
differs from this species by absence of cruciform pores on the body. The number of
antennal segments are not entirely visible, on this one specimen there is some division
but not a typical segmentation, as it is usually seen in the related species.
The specimens in the type series of An. franceschinii with well developed 7 segmented
antennae belongs to An. pseudinsignis.
Acanthococcidae 257
Genus:
Borchseniococcus Kaydan & Kozár, 2008
Type sp.: Borchseniococcus duzgunesae Kaydan & Kozár. By monotypy.
Lit.: Kaydan & Kozár, 2008:16; Kozár & Konczné Benedicty, 2008a:148 (Miller et
al., 2013).
Description
Antennae 7 segmented; frontal tubercle present. Multilocular pores with 5–10 loculi, and
macrotubular ducts both present in small numbers, scattered over venter. Legs short, with
tibia shorter than tarsus. Claw with denticle. All coxae with spinulae; posterior coxae also
with small pores. Macrotubular ducts heavily sclerotized, each with inner ductules ending
in a simple sclerotized pore; terminal gland not observed scattered on both surface.
Microtubular ducts short; absent on venter, few on dorsum. Cruciform pores present on
venter situated on submargin of thorax. Ventral setae short and hair-like. Enlarged setae
absent on dorsum. Anal lobes not well developed; dorsal surface of each lobe with two
hair-like setae, ventral surface of each lobe with a short apical seta and shorter subapical
seta. Anal ring sclerotized but not well developed and lacking anal ring pores but with six
strong setae, all shorter than diameter of ring. Cauda absent.
Ecology
Distribution: Palearctic, with 1 species known from Irano-Turanian subregion.
Biology: Feed on the roots of herbaceous plants.
Borchseniococcus duzgunesae Kaydan & Kozár, 2008 (Fig. 95)

Borchseniococcus duzgunesae Kaydan & Kozár, 2008: 16.
Holotype: Female. Turkey, Iğdır-Tuzluca-Gaziler road, N: 40° 06’717’’, E: 043°

34’183, 993 m altitude, on the root of Panderia pilosa, M. B. Kaydan, 28.vi.2005, (KPCT:
1966). Paratypes: One female on a separate slide with the same data as for holotype,
by monotypy and original designation. In addition four females on one slide labelled:
Turkey, Iğdır-Tuzluca-Gaziler road, N: 40° 06’717’’, E: 043° 34’138, 990 m altitude,
on the root of Panderia pilosa, M. B. Kaydan, 12.06.2007, coll. No 3562. Deposited in
Kaydan’s collection, and two females on two slides in the collection of PPI.
Lit.: Erkılıç, et al., 2011: 16; Kozár & Konczné Benedicty, 2008a: 148; Kozár, 2009:
Description
Unmounted female
Adult females dark yellow to dark brown; found on the root under cavities with honeydew
secretion.
Mounted female
Body elongate oval, 1.35–1.83 mm long, 0.75–0.89 mm wide. Frontal tubercle present.
Antennae 7 segmented, length of segments: I: 24–40, II: 18–28, III: 28–34, IV: 28–35,
V: 15–24, VI: 15–26, and VII: 22–32 µm long; segment III with almost parallel sides;
each segment covered with a few, strong hair-like setae; apical segment with apical seta
52–54 µm long; apical segment also with 3 falcate sensory setae, each 18–22 µm long; two
preapical segments each also with 1 sensory falcate seta 12–14 µm long. Frontal tubercle
present. Anal lobes weakly developed, each with two hair-like setae on dorsal surface, one
seta 8 µm long, other 12–14 µm long; apical seta 92–105; subapical setae 28–30 µm long.
Eyes situated on venter near margin.
Venter: Labium 3 segmented, 70–82 µm long; basal segment not well developed, but
with two different sized setae on each side; median setae on apex of labium 7–9 µm long;
stylet loop long, reaching to level with posterior legs. Legs comparatively small: lengths
of segments and digitules of prothoracic legs; coxa: 44–54, trochanter: 30–40, femur:
80–104, tibia: 54–65; tarsus: 74–94, claw: 28–30, tarsal digitules: 36–44, claw digitules:
24–28 µm long; lengths of segments and digitules of mesothoracic legs: coxa: 50–55,
trochanter 27–35, femur 80–85, tibia 62–64, tarsus 85–94, claw: 30, tarsal digitules: 40–
44, claw digitules: 25–28 µm long; lengths of segments and digitules of metathoracic legs:
coxa: 54–60, trochanter 30–38, femur: 80–84, tibia: 60–70, tarsus: 92–94; tarsal digitules
knobbed, 38–40, claw: 28–32 µm, claw digitules 25–28 µm long, each slightly knobbed.
All coxae with spinulae on anterior surface; posterior coxae also with small pores on
posterior surface. Claw with a denticle. Legs with a few hair-like setae; tibia each with
4–5 setae, tarsus with 5 setae. Multilocular pores each 6–8 µm in diameter, with 5–10
Acanthococcidae 259
Figure 95. Borchseniococcus duzgunesae Kaydan & Kozár, 2008, female. After Kaydan &
Kozár (2008) with modifications.
loculi, generally 8–10, distributed in sparse rows on all abdominal and thoracic segments.
Diameter of anterior spiracles 15–22 µm. Derm with a sparse covering of scattered
capitated hair-like setae, each about 28–46 µm long. Macrotubular ducts each 2.5–3.0
µm wide and 12.5–17.5 µm long; scattered throughout venter but more abundant on
submargin. Microtubular ducts absent. Cruciform pores few, present on submargin of
thorax. Suranal setae setose, short.
Dorsum: Dorsal setae hair-like, slightly blunt, each 12–28 µm long, in sparse rows across
all segments, longest on margins. Macrotubular ducts short, broad, each 4–6 µm wide
and 18–20 µm long, few, scattered throughout dorsum. Microtubular ducts each 2.0–2.5
µm wide and 3–4 µm long, few, scattered over dorsum. Anal ring strongly sclerotized,
without pores, circular, 30–34 µm in diameter, with 6 strong short setae, each 7.5–8.0
µm long; anal ring situated on margin of dorsum, anterior part with a gap. Cauda absent
(after Kaydan & Kozár, (2008)).
Ecology
Host plant: Panderia pilosa.
Biology: On the root under cavities with honeydew secretion.
Genus:
Eriokermes Miller & Miller, 1993
This genus was described in Kermesidae, with type species Eriococcus gillettei Tinsley, 1899
by original designation. The genus Eriokermes was erected for three species previously
placed in the Eriococcidae, including Eriococcus juniperi Goux, 1936 from Palaearctic
Region. Recent papers by Kaydan & Kozár (2008: 6), Kozár & Konczné Benedicty
(2008b: 256) treated Eriokermes as an eriococcid. However these works concerned only
the species Eriococcus juniperi Goux, 1936. Kozár (2009: 106) transferred E. juniperi to the
genus Gossypariella in the Acanthococcidae FG.
Acanthococcidae 261
Genus:
Gossyparia Signoret, 1875
Type sp.: Coccus ulmi Linnaeus, 1767: 740. By original designation.
Current status: Coccus spurius Modeer, 1778: 43.
Lit.: Borchsenius, 1949: 327; Cockerell, 1899b: 268; Hoy, 1963: 127; Kosztarab &
Kozár, 1978: 76; 1988: 289; Kozár & Walter, 1985: 74; Leonardi, 1920: 464; Morrison
& Morrison, 1966: 85; Schmutterer, 2008: 84; Tang & Hao, 1995: 501; Tereznikova,
1981: 50; Ferris, 1955: 94; Williams, 2009: 46 (Miller et al., 2013).
Description:
Adult female in live lilac; ovisac felt-like, incomplete; ruptured in middle of dorsum,
dorsum of female uncovered. Adult female oval, narrowed posteriorly with large anal
lobes; eyes circular, laterad from base of antennae; antennae 6 or 7 segmented; frontal
lobe large; stylet loop almost as long as body; legs well-developed, claw with denticle;
cruciform pores on ventral submargin; quinquelocular disc pores only on venter;
microtubular ducts long, narrow, only on dorsum; macrotubular ducts on dorsum and
venter form a band along body margin; dorsum uniformly covered with enlarged setae;
they form a row on the margin with 3-5 setae on each segment; venter with hair-like setae
only; anal ring with pores and 8 setae; anal lobes sclerotized with small teeth on inner
margin, each with long apical seta and usually with 3 enlarged setae on dorsal surface.
Cauda well developed, with teeths on margin.
Host plant
Feed on woody plants.
Distribution
Holarctic genus, with two species in the Palaearctic Region.
Biology
One yearly generation; biology was well studied (Miller & Miller, 1993; Herbert 1924).
Comments
Ferris, (1955: 94) synonymized this genus with Eriococcus, noting that the unique distribution
of the dorsal macrotubular ducts is not considered sufficient to treat Gossyparia as distinct
from ‘Eriococcus’. It was accepted by Hoy (1963), Williams (1985a) and others, but it was
not accepted by Kosztarab & Kozár (1978, 1988), Danzig (1980) and Kozár (2009). Here
we follow the study of Kozár (2009).
Key to species
1. – Venter with wide bands and groups of multilocular pores......................... G. salicicola

– Venter with scattered rows of multilocular pores............................................. G. spuria
Gossyparia salicicola ( Borchsenius, 1949) (Fig. 96)

Gossyparia salicicola Borchsenius, 1949: 328.
Lectotype: Female. Tadjikistan (Gissar Ridge), 15/07/1944, by N. Borchsenius.

Notes: One paralectotype on same slide as lectotype. Designated by Danzig, 1996:
521. Deposited in ZMAS.
Lit.: Afifi, 1968:190-193; Babaeva, 1980: 57; Bazarov, 1962: 53, 61; Dziedzicka, 1968:
141; Hoy, 1962: 22; Hoy, 1963: 128; Kosztarab & Kozár, 1978: 76; 1988: 289; Koteja,
1974a: 152; Köhler, 1998: 388; Lashin, 1956: 114; Matesova, 1967: 1202; Miller &
Gimpel, 2000: 324; Myartseva, 1972: 54; Tang & Hao, 1995: 501; Tao, 1999: 33;
Yasnosh, 1973: 908; Yasnosh, 1978: 491; Yasnosh & Borovkov, 1974: 35 (Miller et
al., 2013).
Eriococcus salicicola; Lindinger, 1958: 368. Change of combination.

Nidularia salicicola; Lindinger, 1958: 368. Change of combination.
Acanthococcus salicicola; Danzig, 1980: 207. Change of combination.
Gossyparia salicicola; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus salicicola; Miller & Gimpel, 1999: 215. Revived combination.
Gossyparia salicicola; Kozár, 2009: 102. Revived combination.
Description
Unmounted female
Adult female eggshaped, reddish-brown, nearly black. Egg sac is grayish, compact and
entirely encloses the body of fertilized young female, later dorsum will be exposing. First
stage nymph are reddish-brown. Up to 2.5 mm long, 1.8 mm wide.
Mounted female
Adult female broadly oval, largest strongly nodulose 3 mm long, 1.8 mm wide. Antennae
7 segmented, I: 30.4, II: 36.9, III: 49.7, IV: 40.8, V: 23.1, VI: 20.8 and VII: 40.0 µm long;
each segment covered with a few, strong hair-like setae (except third segment); apical
segment with apical seta and 3 sensory falcate setae. Frontal lobes well developed. Eyes
situated on venter near margin. Anal lobes strongly developed, about twice as long as
wide, rounded at apex, sclerotised, nodulose, with sclerotized teeth, each with 3 enlarged
setae on dorsal surface, 65–80 µm; apical seta up to 300 µm; ventrally each with two
flagellate setae; subapical setae 155 µm, other setae 50 µm.
Acanthococcidae 263
Figure 96. Gossyparia salicicola Borchsenius, 1949, female. After Matesova (1967) with
modifications.
Venter: Labium wide, stylet loop reaches till sternite of abdominal segment V, with
normal flagellate setae in median areas, each short and slender but stouter and stiff in
lateral areas; apical labial segment with 5 pair of hair-like setae, of what the median is
somewhat shorter and thicker. Legs well developed; hind femur 155 µm, tibia 115 µm,
tarsus with pores, 145 µm long; claw thick, with small denticle. Meso and metathoracic
coxae each with spinulae on ventral surface. Tibiae of meso and metathoracic legs each
with 4 setae (median seta absent), tarsi each with 6 setae. Setae in median areas flagellate,
about 115 µm long, setae in lateral areas spine like, 24–30 µm long. Multilocular pores 5
µm in diameter and with 5 loculi, distributed in sparse bands on all abdominal segments
and scattered on thorax and head, few trilocular pores in median area of thoracic venter.
Cruciform pores generally on submedian area in band on thorax and abdominal segments
and a few on head; macrotubular ducts form a wide band on lateral margins; ventral hair-
like setae in transverse rows on abdominal sternites, some scattered on prosoma.
Dorsum: Strongly nodulose; with numerous enlarged setae cylindrical, slender, sides
straight, apices round, marginal setae slightly longer than remaining dorsal setae, setae
abundant over dorsum. Marginal dorsal setae long blunted each 50–80 µm forming a
marginal row and differentiated from dorsal setae. Enlarged setae form also a longitudinal
band in the midline of body, the setae 20–55 µm long. Macrotubular ducts present
marginally on dorsum and mediolaterally on thorax and first abdominal segment; 24–30
µm long and 8 µm wide, ducts form a wide band on lateral margins. Microtubular ducts
long, each 6 µm long with oval or bifurcated dermal orifice, scattered over dorsum. Anal
ring with partly double row of pores and 8 setae, each 130 µm wide. Cauda triangular,
lightly sclerotised, strongly nodulose (after Borchsenius (1949), with modifications based
on lectotype and paralectotype).
Other stages
First instar described by Borchsenius (1949).
Ecology
Host plant: Salix sp., S. capitata.
Distribution: China (Inner Mongolia), Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan,
Uzbekistan.
Biology: Lives on the trunk and twigs in cracked bark. Female appeared in May, crawlers
in June.
Acanthococcidae 265
Gossyparia spuria (Modeer, 1778) (Fig. 97)

Coccus spurius Modeer, 1778: 43.
Type status: Unknown. Notes: Type has probably been lost (Williams, 1985a).
Common name: Elm scale, Elm-Tree Scale, European Elm-Scale, Elm Bark Louse.
Lit.: Afifi, 1968:182; Cockerell, 1899b: 268; Ferris, 1955: 164; Herbert, 1924:1; Hoy,
1963: 127; Koteja, 1974a: 152; Kozár & Kosztarab, 1982: 204, 201; Kozár & Walter,
1985: 74; Leonardi, 1920: 464; Lindinger, 1912: 331; Malumphy et al., 2009: 120;
Morrison & Morrison, 1966: 85; Schmutterer, 1952: 416; Tereznikova, 1981: 50;
Williams, 2009: 46; Zahradník, 1972: 402 (Miller et al., 2013).
Coccus ulmi; Linnaeus, 1758: 455. Type status: Unknown. Notes: Type has probably been
lost (Williams, 1985a).
Coccus lanigera; Gmelin, 1790: 2221. Type status: Unknown. Synonymy by Hoy, 1963: 130.
Coccus gramuntii; Planchon, 1864: 24. Type status: Unknown. Type locality: France.
Synonymy by Herbert, 1924: 1.
Nidularia lanigera; Targioni Tozzetti, 1868: 727. Change of combination.
Nidularia gramuntii; Targioni Tozzetti, 1869: 727. Change of combination.
Gossyparia gramuntii; Signoret, 1875: 23. Change of combination.
Gossyparia spuria; Cockerell, 1899b: 268. Change of combination.
Nidularia spuria; Lindinger, 1933a: 108. Change of combination.
Gossyparia spuria; Borchsenius, 1949: 327. Revived combination.
Eriococcus spurius; Ferris, 1955: 164. Change of combination.
Gossyparia spuria; Kosztarab & Kozár, 1978: 76. Revived combination.
Acanthococcus spurius; Danzig, 1980: 207. Change of combination.
Gossyparia spuria; Kosztarab & Kozár, 1988: 289. Revived combination.
Eriococcus spurius; Miller & Gimpel, 2000: 341. Revived combination.
Gossyparia spuria; Kozár, 2009: 102. Revived combination.
Description
Unmounted female
Adult female oval, chestnut colour, covered with white wax powder, 3.5 mm long, 2
mm wide. Ovisac ovoid, compact, panlike, grayish-white, covers only venter and body
margins, exposing dorsum.
Mounted female
Adult female broadly oval, largest strongly nodulose 3 mm long, 1.8 mm wide. Antennae
7 rarely 6 segmented, 221–300 µm long; each segment covered with a few, strong hair-like
setae (except third segment); apical segment with apical seta and 3 sensory falcate setae.
Frontal lobes well developed. Eyes situated on venter near margin. Anal lobes strongly
developed, about twice as long as wide, rounded at apex, sclerotised, strongly nodulose,
with sclerotized teeth, each with 3 enlarged setae on dorsal surface; apical seta 230–247
µm long; ventrally each with two flagellate setae; subapical setae 148 µm long.
Venter: Labium 150 µm long, stylet loop reaches sternite of abdominal segment V, with
normal flagellate setae in median areas, each short and slender but stouter and stiff in
lateral areas; apical labial segment with 5 pairs of hair-like setae, of which the median is
somewhat shorter and thicker. Legs well developed; hind femur 136–140 µm, tibia 105–
116 µm, tarsus 136–140 µm long; claw thick, with small denticle. Meso- and metathoracic
coxae each with spinulae on ventral surface. Tibiae of meso and metathoracic legs each
with 4 setae (median seta absent), tibiae of prothoracic legs each with 5 setae (median
seta present), tarsi each with 6 setae. Setae in median areas flagellate, long, setae in lateral
areas spine like. Multilocular pores each 3–5 µm in diameter with 5 loculi, distributed in
sparse bands on all abdominal segments and scattered on thorax and head, few trilocular
pores in median area of thoracic venter. Cruciform pores on generally on submedian area
in band on thorax and abdominal segments and a few on head; macrotubular ducts form
a wide band on lateral margins; ventral hair-like setae in transverse rows on abdominal
sternites, some scattered on prosoma.
Dorsum: Strongly nodulose; with numerous spinose setae in regular distribution except on
intersegmental areas of thorax where they are absent, but present on mid-line. Setae long
and slender with almost straight sides, with blunt point. Marginal dorsal setae long blunted
each 60 µm forming a marginal row and differentiated from dorsal setae. Macrotubular
ducts, each 7.5–10.0 µm ducts form a wide band on lateral margins. Microtubular ducts
long, each 10 µm long with oval or bifurcated dermal orifice, scattered over dorsum. Anal
ring with partly double rows of pores and 8 setae, each 125–136 µm. Cauda (median
dorsal plate) triangular, lightly sclerotised, strongly nodulose (after Kosztarab & Kozár
(1988); Williams (1985a); with modifications).
Other stages
Males
Two winged forms of adult males described in detail by Afifi (1968).
Nymphs
Immature stages described by Herbert (1924) and Dziedzicka (1961).
Mounted first instar nymph (Fig. 97, top right).

with strong sensory setae as in adult female. Frontal tubercle present. Dorsum with strong
spines with pointed apices of two sizes, forming middorsal longitudinal pair of rows, the
larger ones five on thorax and two on abdominal segments VI-VII, four segments of
abdomen with very short setae, the submedian row contains only one large spines, what
is continued by nine very small setae. Microtubular ducts long, few in two middorsal
longitudinal rows on both side. Venter with transverse rows of six small hair-like setae
on each abdominal segment; median setae longer than others. Cruciform pores present
Acanthococcidae 267
Figure 97. Gossyparia spuria (Modeer, 1758), female, after Williams (1985) with
modifications. First instar at top right, original.
on thoracic and first abdominal segments, with 1 or 2 pair of quinquelocular pores on

each thoracic segment, 1 near each spiracle, and two longitudinal submedian rows on
abdomen. Anal ring with 6 spine-like setae.
Ecology
Host plant: Alnus sp., A. crispa, A. glutinosa, Corylus sp., Fraxinus sp., Salix sp., Ulmus sp.,
U. americana, U. angustifolia, U. campestris var. pyramidalis, U. glabra, U. glabra var. camperdownii,
U. laevis, U. minor, U. procera, U. pumila, U. rubra, U. thomasii, Ulmus x hollandica Dampieri,
Viscum sp.
Distribution: Austria, Bulgaria, Canada, China, Croatia, former Czechoslovakia,
Denmark, Finland, France, France (Corsica), Georgia, Germany, Greece, Hungary,
Iraq, Iran, Italy, Italy (Sardinia), Japan, Latvia, Lithuania, Moldova, Mongolia, Morocco,
Netherlands, Norway, Poland, Portugal, Romania, Russia, Slovenia, Spain, Sweden,
Switzerland, Turkey, Ukraine, United Kingdom, USA, Uzbekistan, former Yugoslavia,
Ukraine. The species is widely distributed in most of the countries in the Palaearctic.
Biology: Lives on trunk, branches and twigs, nymph on leaves, sometimes serious pest
on elms. It has bisexual or rarely parthenogenetic reproduction; one yearly generation;
second instars overwinter; adults appear end of April or first part of May; lay 117-416
eggs per female starting in June; eggs hatched from end of July (Schmutterer, 1952).
Herbert (1924) reported from Western USA that eggs hatched in less than one hour after
laying; first instars migrated to both sides of leaves; often settled near the main veins;
molted in about six weeks into second instars; nymphs moved in the fall to crevices of
the bark for overwintering; male nymphs started forming cocoons from late January,
while female nymphs molted into adults in March and April; egg laying started end of
May and lasted for several months, until about mid- August.
Often becomes a pest on ornamental elms in urban areas. The large quantity of honeydew
they produce coats the leaves, and on the honeydew sooty mold, Coniothecium effusum
(Dziedzicka, 1961) forms and covers the foliage.
Acanthococcidae 269
Genus:
Gossypariella Borchsenius, 1960
Type sp.: Rhizococcus siamensis Takahashi, 1942, by monotypy and original designation.
Lit.: Borchsenius, 1960c: 920; Hoy, 1963: 132; Kozár, 2009: 113; Kozár et al., 2007:
88; Miller & Gimpel, 2000: 384; Morrison & Morrison, 1966: 86; Takahashi, 1942:
6–8; Tang & Hao, 1995: 504, 651; Wang, 2001: 236–237; Yang, 1982: 101 (Miller et
al., 2013).
Description
According to Takahashi (1942), dried specimens blackish brown, body subcircular.
Adult female elongate-oval, narrowed posteriorly, anal lobes conical and normally
heavily sclerotized. Antennae 7 segmented. Frontal lobes, or tubercles present. Labium
3 segmented, with well developed segments and a weakly developed basal segment with
two pairs of hair-like setae. Legs well-developed, midcoxae and hindcoxae often with
spinulae on anterior surfaces, translucent pores sometimes present, inner side of hind
tibia with only four setae, claw usually with a denticle; tarsal and claw digitules longer
than claw clavate. Spiracles often with a few associated disc pores. Discoidal pores usually
quinquelocular (rarely 3 or 4 loculi), cruciform pores on prosomal venter in a marginal
band. Tubular ducts of 2 types: micro- and macrotubular ducts; microtubular ducts
slender, scattered or form transverse rows or bands on dorsum, macrotubular ducts form
longitudinal marginal band or row; or in transverse sparse bands on dorsum. Enlarged
conical setae elongate, with acut apices, on entire dorsum, where they form transverse
bands or rows; hair-like setae on venter only. Anal ring well developed, sclerotized with
partly double row of pores and with 8 anal ring setae, latter often as long as apical seta
on anal lobes. Each anal lobe with a long apical seta and usually with 3 elongate dorsal
conical setae, at least with 2 ventral hair-like setae also present; suranal setae hair-like.
Cauda usually well developed ( after Kozár et al. (2007).
Distribution
Palearctic-Oriental, with 3 species in the Far-Eastern and Mediterranean subregions.
Biology
Key to species
1. – On the abdominal margin with 2 long, setose spines.....................................G. juniperi

– On the abdominal margin 3 long setose spines.............................................................2
2. – Quinquelocular pores on dorsum present.................................................... G. phyllanthi
– Quinquelocular pores on dorsum absent...................................................... G. siamensis
Gossypariella juniperi (Goux, 1936) (Fig. 98) Redescription, Combination

nova
Eriococcus juniperi Goux, 1936: 353.
Holotype: Female. France: Bouches-du-Rhone, Marseille, Maraseilleveyre, on

Juniperus phoenicea, by L. Goux, 25. IV. 1933. (MNHN 14498-01). Paratypes: 7 slides
are with the same data as holotype (MNHN 14498-02-08), on slide MNHN 14499,
collected in the same place in August, marked a male but it was not found, on slide
MNHN 14500, collected in the same place in May a first instar larva was found.
Deposited in MNHN.
Lit.: Balachowsky, 1949: 115; Foldi, 2001: 303; 2002: 246; Köhler, 1998: 400; Kozár,
2009: 106; Lindinger, 1943: 223; Miller & Miller, 1993: 239; Miller, 1984: 323; Miller
& Gimpel, 2000: 475; Miller et al., 2005: 141; Ouvrard & Kozár, 2009: 102-118; Panis,
1981: 6; Zahradník, 1972: 401 (Miller et al., 2013).
Nidularia juniperi; Lindinger, 1943: 223. Change of combination.

Eriococcus juniperi; Hoy, 1963: 97. Revived combination.
Rhizococcus juniperi; Kozár & Walter, 1985: 75. Change of combination.
Eriokermes juniperi; Miller & Miller, 1993: 239. Change of combination, and removed from
Eriococcidae to the Kermesidae.
Rhizococcus juniperi; Köhler, 1998: 400. Revived status and combination.
Description
Unmounted female
Adult female oval, chestnut colour, covered with white wax powder.
Mounted female
173–202 µm long, the size of the segments: I: 30–33, II: 26–29, III: 38–43, IV: 20–28,
V: 15–17, VI: 16–20, and VII: 28–32 µm; each segment covered with a few, strong hair-
like setae (except third segment); apical segment 45–57 µm long with apical seta and
3 sensory falcate setae, the longest 21–28 µm long; falcate sensory setae on preapical
segment 18–24 µm long. Frontal tubercle present, frontal lobes not seen. Eyes situated
Acanthococcidae 271
Figure 98. Gossypariella juniperi (Goux, 1936), female. Original.

on venter near margin. Anal lobes slightly sclerotized, fused at the base, each with two
setose spines along inner margin, and one setose spine on outer margin, similar in size to
those on margin, apical seta 90–95 µm long. The subapical seta hair-like.
Venter: Labium 3 segmented, 49–55 µm long; stylet loop long reaching the posterior
legs. Legs well developed; lengths of segments and digitules of prothoracic legs; coxa:
45–55, trochanter: 35–40, femur: 90–106, tibia: 55–61, tarsus: 85–87, claw 24–30 µm long;
lengths of segments and digitules of mesothoracic legs: coxa: 50–56, trochanter: 35–42,
femur: 100–125, tibia: 60–69, tarsus: 90–99, claw: 24–29 µm long; lengths of segments
and digitules of prothoracic legs: coxa: 55–61, trochanter: 32–49, femur: 100–136, tibia:
60–68, tarsus: 93–108, tarsal digitules knobbed, 38–42, claw 23–29, claw digitules, 30–32
µm long, slightly knobbed. Meso- and metathoracic coxae each with spinulae on ventral
surface. Tibiae of meso and metathoracic legs each with 4 setae (median seta absent),
tibiae of prothoracic legs each with 5 setae (median seta present), tarsi each with 6 setae.
The diameter of anterior spiracles 13–15 µm. Setae in median areas flagellate, long, setae
in lateral areas spine like. Some quinquelocular pores found around spiracles, 4 µm in
diameter. Multilocular pores with 5-12 loculi found in rows on last abdominal segments.
Venter with a small number of scattered, hair-like setae. Microtubular duct present in a
submarginal band. Macrotubular ducts about 3–5 µm wide and 17 µm long, scattered.
Cruciform pores present on submargin.
Dorsum: Dorsal setae few, hair-like, each spine 5–8 µm long. On margin of segments two
spines with acute apices, 18–24 µm long. Macrotubular ducts similar to those on venter,
sparse, present throughout, 5–8 µm wide and 19–20 µm long. Microtubular ducts 4 µm
long, scattered throughout. Disc pores absent. Anal ring 35–56 µm wide, 40–42 µm long,
with pores and with 8 hair-like setae, each 75–90 µm long. Cauda present .
Other stages
First instar described by Goux (1936).
Ecology
Host plant: Cupressus sp., Juniperus phoenicea.
Biology: Goux (1936) only found this species in small numbers and stated that G. juniperi
probably has one generation per year. He observed the formation of the ovisac in the
middle of April.
Comments
Gossypariella juniperi was transferred to the Kermesidae family by Miller & Miller (1993) in
a newly established genus Eriokermes. According to our study G. juniperi is not congeneric
with E. gillettei, because E. gillettei has microtubular ducts, G. juniperi does not have simple
discoidal pores. Here it is resurrected to the Acanthococcidae in a new combination
with Gossypariella genus. The presence of setose spines on margin and on anal lobes
Acanthococcidae 273
needs further special studies. The setae and anal lobes of Eriokermes junipericola (De Lotto,
1954) from Eritrea, collected also on Juniperus, are similar to G. juniperi, and probably
belongs to Gossypariella genus. This character is very sparse among genera and species
of Acanthococcidae. In the collection of MNHN (11827) there is a slide from Spain
(Espagne: Zaragoza, Pine de Ebro), by JBZ 3251, on Juniperus phoenicea, containing a first
instar nymph of an Acanthococcidae. This specimen shows some characters that are
quite different, as the dorsal spines on anal lobes not setose as on the first instar nymph
of G. juniperi, but with blunted enlarged spines, as usual in other Acanthococcidae.
Gossypariella phyllanthi (Ferris, 1957) (Fig. 99)

Eriococcus phyllanthi Ferris, 1957b: 86.
Holotype: Female. India: Kerala, Bangalore, on Phyllanthus emblica, by T.S.

Muthukrishnan. Unknown type status, type designation unknown. Notes: No type
material has been found till now in either the BMNH, MNHN, ZMAS, UCDC or the
USNM collections.
Lit.: Ali, 1970: 77; Ferris, 1957b: 86; Hoy, 1963: 108; Kozár, 2009: 102; Kozár et al.,
2007: 88; Miller & Miller, 1993: 239; Miller & Gimpel, 2000: 384; Tang & Hao, 1995:
504, 651; Williams, 1985a: 349 (Miller et al., 2013).
Gossypariella phyllanthi; Tang & Hao, 1995: 504. Change of combination.
Description
Unmounted female
According to the mounted slide insect partially enclosed within a sac that is entirely open
along the back, with secretion occuring as a wall along the sides of body (Ferris, 1957b).
Mounted female
Body broadly oval 1.0 mm long. Antennae 7 segmented. The segments of the antennae
are covered with few setae. Anal lobes sclerotized, with teeth on margin, slender, with
two setae along inner margin, and one setae on outer margin.
Venter. Labium 3 segmented. Legs small, tarsus twice longer than tibia. Claw with
denticle. Legs with few setae. Quinquelocular pores distributed in sparse rows on all
segments of abdomen and thorax. Venter with a small number of scattered, spinelike
setae. Macro- and microtubular duct not shown.
Dorsum. Dorsal setae short, slender, elongate, apices acute, found in sparse rows on all
segments, and in 2 or 3 on the margin, other dorsal setae hair like, slightly smaller than
marginal enlarged setae. Macrotubular ducts shown only on the margin. Microtubular
ducts not shown. Quinquelocular pores scattered mostly on the last abdominal segments
of dorsum. Anal ring not clearly seen on the drawing, and not mentioned in the
description. Apical setae longer than lobe. Anal plate heavily sclerotized, surface with
reticulation. Cauda present, triangular, with several teeth on posterior margin (Ferris,
1957b).
Ecology
Host plant: Phyllanthus emblica.
Distribution: India (Kerala).
Gossypariella siamensis (Takahashi, 1942) (Fig. 100)

Rhizococcus siamensis Takahashi, 1942: 6
Lectotype: Female. Thailand (Chiengmai), on Ficus sp., 06/04/1940, by R. Takahashi.

Designated by Kozár et al (2007), on the same slide one paralectotype female and one
Coccidae species present. Deposited in the Taichung: Taiwan Agricultural Research
Institute, Entomology Collection, China, Taiwan.
Lit.: Ali, 1970: 78; Borchsenius, 1960c: 920; Hoy, 1963: 132; Hua, 2000: 138; Köhler,
1998: 389; Kozár, 2009: 102; Kozár, et al., 2007: 84; Kozár & Walter, 1985: 74; Miller
& Gimpel, 2000: 385; Takahashi, 1942: 6; Tang & Hao, 1995: 504; Tao, 1999: 34;
Wang, 1982b: 156; 2001: 237; Yang, 1982: 105 (Miller et al., 2013).
Gossypariella siamensis; Borchsenius, 1960c: 920. Change of combination.
Description
Unmounted female
Not seen, according to Takahashi (1942) description blackis brown in dried specimens,
body subcircular, about 1.25 times longer than wide.
Mounted female
Body elongate oval, 1.91 mm long, 1.47 mm wide. Antennae 7 segmented, the size of the
segments; I: 31, –II: 36, III: 34, IV: 34, V: 19, VI: 21, and VII: 31 µm. The third segment
is almost parallel sided. Apical setae of antenna 56 µm long. On apical segment three
sensory falcate setae are found. On the two preapical segments 12–24 µm long, falcate
sensory sete present. The segments of the antenna are covered with few long hair-like
setae. Frontal lobes not seen, some sclerotization seen at the base of the antennae. Eyes
situated on venter near margin. Anal lobes with two hair-like setae along inner margin,
42–68 µm long, and one 55 µm long setae on outer margin. Apical setae 193 µm long.
Anal lobes heavily sclerotized, surface with reticulation.
Venter: Labium 3 segmented, 119 µm long. Stylet loop almost as long as the body. Legs
well developed, small: lengths of segments and digitules of prothoracic legs; coxa: 53,
trochanter: 34, femur: 96, tibia: 70, tarsus: 86, claw: 26 µm long; lengths of segments
Acanthococcidae 275
Figure 99. Gossypariella phyllathi (Ferris, 1957), female. After Ferris (1957) with
modifications.
and digitules of mesothoracic legs; coxa: 53, trochanter: 36, femur: 91, tibia: 70, tarsus:
84, claw: 26 µm long; lengths of segments and digitules of metathoracic legs; coxa: 65,
trochanter: 34, femur: 113, tibia: 74, tarsus: 96, tarsal digitules: 42 µm long, knobbed,
claw: 29 µm, claw digitules: 30 µm long, slightly knobbed. Middle and posterior coxae
with some reticulation, on posterior coxa some pores present, too. Claw with denticle.
Tibiae of meso and metathoracic legs each with 4 setae (median seta absent), tibiae of
prothoracic legs each with 5 setae (median seta present), tarsi each with 6 setae. The
diameter of anterior spiracles 29 µm. Quinquelocular pores distributed in sparse bands
and rows on all segments of abdomen and thorax, 4–5 µm in diameter. Venter with a
small number of scattered, hair-like setae. Microtubular duct absent. Macrotubular ducts
of one size, about 4 µm wide and 12 µm long, present in a submarginal band. Cruciform
pores 4 µm in diameter, scattered on margin and form groups in submarginal band, some
scattered in the venter of middle of thorax. Multilocular pores with 5–7 pores, 4–5 µm
in diameter.
Dorsum: Dorsal setae with acute apices, 36–92 µm long, found in sparse rows on all
segments. On the margin 2-3 setae present, other dorsal setae hair like, slightly smaller
than marginal enlarged setae. Macrotubular ducts absent. Microtubular ducts double
apex, 5 µm long, scattered among dorsal setae. Anal ring not well seen, situated between
venter and dorsum. Anal ring with eight, hair-like setae, about 105 µm long. Anal ring
pores not seen. On penultimate segments, before of anal ring, a sclerotized plate present,
86 µm wide. Cauda 67 µm wide (after Takahashi (1942)).
Ecology
Host plant: Ficus sp.
Distribution: China (Yunnan), Thailand.
Biology: On the twigs.
Comments
In study of specimens collected and described as type species of the genus by Borchsenius
(1960c) from China, were compared with the lectotype and paralectotype of G. siamensis
collected by Takahashi and important differences were not found. This species is very
similar to G. phyllanthi (Ferris, 1957b). According to the drawing of Ferris (1957b), there
are some differences in number of pores on the dorsal margin, in the shape of cauda
and structure of anal lobes, this question needs further study, if the type material of G.
phyllanthi will be found.
Acanthococcidae 277
Figure 100. Gossypariella siamensis (Takahashi, 1942), female. After Kozár et al (2007),
with modifications.
Genus:
Greenisca Borchsenius, 1948
Type sp.: Eriococcus inermis Green, 1915 (misidentified) = Anophococcus gouxi Balachowsky,
1954, by original designation. Synonymy by Williams, 1985a: 356. Notes: According
to Williams (1985a) "there are some nomenclatural problems concerning the type
species." According to Danzig (1962a) the species on which Borchsenius (1948) based
Greenisca was not E. inermis Green, but another species, which Danzig (1980) stated,
was later described as Anophococcus gouxi Balachowsky; the type species, therefore, of
Greenisca should be either A. gouxi Balachowsky, 1954 or = E. inermis Green, 1915.
Article 70 of the International Code on Zoological Nomenclature states that "if a zoologist
considers that a type-species designated for a new genus has been misidentified,
then that person should refer the case to the Commission to designate as the type-
species whichever species will in its judgement best serve stability and uniformity of
nomenclature..." A petition has not been submitted to the commission. Furthermore,
the anal lobes of G. gouxi illustrated by Danzig (1980) have three enlarged setae, but
the same species illustrated by Balachowsky (1954) has the outer setae much reduced
in size. E. inermis has minute outer setae on the anal lobes, frontal tubercules and
microducts without a bifid orifice. This question remains to be resolved.
Lit.: Balachowsky, 1954: 61; Borchsenius, 1948a: 502; 1949:367; 1956; Borchsenius &
Danzig, 1966: 41; Brozek, 2006; Danzig, 1980: 205; 2006: 203; Ferris, 1957b: 86; Hoy,
1963: 132; Gavrilov & Smirnova, 2005; Golan & Drabik, 2004; Kaydan & Kozár,
2008: 6; Kosztarab & Kozár, 1988: 291; Koteja & Zak Okaza, 1981: 508; Kozár &
Konczné Benedicty 2008b: 256; Kozár & Walter, 1985: 74; Marotta, 1993; Matesova,
1960: 205; Morrison & Morrison, 1966: 87; Ouvrard & Kozár, 2009; Pellizari &
Kozár, 2011: 58; Schmutterer, 2008: 84; Tang & Hao 1995: 642; Tereznikova, 1981:
52; Williams, 1985a: 356 (Miller et al., 2013).
Comments
Greenisca laingi; (Bodenheimer, 1953).
Pseudococcus laingi; (Bodenheimer, 1953): 121.
Syntype: Female. Turkey (Kaya-ardi near Nigde), on grass. Original type material lost
(Ben-Dov & Harpaz, 1986). Notes: There are no characters in the description and
drawing that could be regarded as a female acanthococcid. On the drawing, the anal
region (2) shows something like a cerarium with two strong spines and according to
the text, 20 pores on the preceding abdominal segment could be also a mistake.
Lit.: Ben-Dov, 1994: 396; Kaydan, et al., 2007: 90; Kozár & Walter, 1985: 74. Köhler,
1998: 389; Miller & Gimpel, 2000: 253 (Miller et al., 2013).
Greenisca laingi; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus laingi; Miller & Gimpel, 2000: 253. Change of combination.
Greenisca?? laingi; Kozár, 2009: 102. Change of status. Notes: Ben-Dov (1994) treated it as
Acanthococcidae 279
a member Pseudococcus of Pseudococcidae. Kozár & Walter (1985) erroneously transferred

this species from Pseudococcus to Eriococcidae sensu lato in to Greenisca based on strong
marginal spines in its description and figure. This was cited later by Köhler (1998), Miller
& Gimpel (2000), Kaydan et al. (2007). Later Kozár (2009) supposed that it could be
similar to Lecanopsis turcica (Bodenheimer, 1951) in the Coccidae family according to
the drawing of a first instar nymph of Bodenheimer (1953). Because of this, the name
should be excluded from the Acanthococcidae.
Description
Ovisac elongate-oval, felt-like, white, or yellowish. Adult female elongate-oval, almost
parallel-sided with convex dorsum. Antennae 6 or 7 segmented, base some distance from
body margin; frontal tubercle usually present.
Venter: Labium 3 segmented, basal segment with a pair of short setae, total number
of setae on labium 8 pairs; stylet loop slightly longer than labium; legs well developed,
slender, each tibia about as long as tarsus, hind tibia with 5 setae, 3 on inner margin, claws
usually with denticle; mid-coxae and hind-coxae with reticulations on anterior surfaces,
hind-coxae with some translucent pores on posterior surfaces; quinquelocular pores
present on both surfaces in small number; cruciform pores on ventrolateral area only;
macrotubular ducts present, of 2 sizes, main ducts with terminal gland flower-shaped;
microtubular ducts present, with small filamentous ductile.
Dorsum: Enlarged conical setae few, these with truncate or sharp-pointed apex, and
present on anal lobes, rarely on margin of thorax and head; hair-like setae on both
surfaces. With small spine-like, or setose spines in rows on all segments, and anal lobes
well developed, each with three spine-like setae; anal ring oval, with 6 or 8 setae, each
as long as the ring diameter and with few pores present in a single row; cauda present;
quinquelocular pores numerous on all segments; macrotubular ducts and microtubular
ducts present (Pellizari & Kozár, 2011).
Distribution:
This genus contains six species, mostly restricted to the Western part of the Palaearctic
Region, (Kosztarab & Kozár, 1988; Pellizari & Kozár, 2011).
Biology:
All species lives on leaves of Poaceae and Cyperaceae; overwinter as eggs, males unknown.
Key to species
1. – Part of sclerotized dorsal quinquelocular pores in wide bands on dorsum

.......................................................................................................................... G. brachypodii
– Sclerotized dorsal quinquelocular pores not in wide bands on dorsum.....................2
2. – Outer dorsal enlarged setae of anal lobes usually much shorter (less than half) of
other two anal lobe setae; spine-like setae absent on margin of abdomen...............3
– All three dorsal anal lobe spinose setae subequal in length; spine-like setae present
on margin of abdomen......................................................................................................4
3. – Marginal enlarged setae on head present...........................................................G. placida
– Marginal enlarged setae on head absent.............................................................. G. gouxi
4. – 3-5 marginal enlarged setae on each body segments........................................ G. erwini
– Marginal enlarged setae not on each body segment .....................................................5
5. – One or two enlarged setae on all abdominal segments and head...............G. oreophila
– Enlarged setae only on last two abdominal segments......................................G. alpina
Greenisca alpina Pellizzari, 1999 (Fig. 101)

Greenisca alpina Pellizzari & Kozár, 1999: 26.
Holotype: Female. Italy (Croce d'Aune, Belluno), on unidentified Gramineae,

21.viii.1990. Deposited in DAFNAE.
Lit.: Kozár, 2009: 102; Miller & Gimpel, 2000: 122 (Miller et al., 2013).
Eriococcus alpina; Miller & Gimpel, 2000: 122. Change of combination.

Kaweckia alpina; Kozár, 2009: 102. Change of combination.
Greenisca alpina; Pellizzari & Kozár, 2011: 66. Revived combination.
Description
Unmounted female
Ovisac ovoid, white, felted.
Mounted female
Adult female elongate-oval, 3.7 mm long, 1.56 mm wide. Antennae 6 segmented, 230
µm long, third segment longest. Frontal tubercle present. Eyes situated on venter near
margin. Anal lobes well developed with 3 enlarged setae (two inside and one outside)
each with 52 µm long and one hair like apical setae.
Venter: Legs well developed, slender, hind trochanter+femur 200 µm long, hind tibia
135 µm long, hind tarsus 142 µm long, claw slightly curved, slender, 33 µm long, with
a minute denticle at near apex; claw digitules longer than claw; tarsal digitules longer
than claw; hind coxa with a group of translucent pores; tibia with 4 setae. Macrotubular
Acanthococcidae 281
Figure 101. Greenisca alpina Pellizzari, 1999 (in Pellizzari & Kozár, 1999), female. After
Pellizzari & Kozár (1999) with modifications.
ducts in two sizes, scattered all surface. Multilocular pores with 5-9 loculi, numereous
around vulva; 4-7 multilocular pores around the spiracle opening. Cruciform pores not
numerous, on head and thorax; hair-like setae slender of various sizes 38–64 µm long,
longer than those on dorsum; ventral surface of anal lobe with 2 slender setae and slender
suranal seta.
Dorsum: Numerous quinquelocular disc pores each 3.2 µm diameter, rarely trilocular
pores, scattered on dorsum, these form transverse bands or groups. Macrotubular ducts
in two sizes, larger ones 16 µm long, 9.6 µm width, smaller ones 16 µm long, 4 µm width,
scattered on thorax and head. Hair-like setae short, no enlarged setae on dorsum except
on anal lobes and head margin. Anal ring oval, with double incomplete rows of pores and
8 setae; anal lobes slightly sclerotized. Marginal enlarged setae on abdominal segment VI
and VII segment (2–4 setae on the last segments), each 30–45 µm long, minute pointed
setae scattered on the surface (after Pellizzari & Kozár (1999), with modifications).
Ecology
Biology: Found on leaves of grasses.
Greenisca brachypodii Borchsenius & Danzig, 1966 (Fig. 102)

Greenisca brachypodii Borchsenius & Danzig, 1966: 43.
Holotype: Female, Kazakhstan: Eastern Kazakhstan Oblast, Zyryanovski District,

Stolbucha, on Brachypodium pinnatus leaflets, 20.vi.1961, Matesova. Paratypes are from
Russia and Latvia. Deposited in ZMAS.
Common name: Falsebrome felt scale.
Lit.: Borchsenius & Danzig, 1966: 43; Brozek, 2006: 255; Danzig, 1971a: 823;
Dziedzicka & Koteja, 1971: 576; Fetykó, et al, 2010: 296; Komosinska & Podsiadlo,
1967: 684; Kosztarab & Kozár, 1988: 292; Koteja & Zak-Ogaza, 1981: 510; Kozár et
al., 2013: 56; Kozár & Walter, 1985: 74; Köhler, 1998: 389; Lagowska & Koteja, 1996:
31; Matesova, 1968: 117; Miller & Gimpel, 2000: 147; Pellizzari & Kozár, 2011: 66.
Tang & Hao 1995: 506 (Miller et al., 2013).
Acanthococcus brachypodii; Miller & Gimpel, 1996: 599. Change of combination.

Eriococcus brachypodii; Miller & Gimpel, 1999: 213. Change of combination.
Greenisca brachypodii; Kozár, 2009: 102. Revived combination.
Acanthococcidae 283
Figure 102. Greenisca brachypodii (Borchsenius & Danzig, 1966), female. After Borchsenius
& Danzig (1966), with modifications.
Description
Unmounted female
Ovisac ovoid, white or yellowish white, loosely felted, fluffy, 4.5 mm long, 1.8 mm wide.
Mounted female
Adult female elongate-oval, 3.1–3.9 mm long, 1.0–2.2 mm wide; antennae 7 (rarely 6)
segmented, 278 µm long, fourth segment longest; frontal tubercle present; eyes situated
on venter near margin. Anal lobes slightly developed with two enlarged setae and one
hair-like setae.
Venter: Labium 93 µm long, 80 µm wide, with 18 setae; basal segment with two pairs
of setae with different sizes; median setae on apical segment spine-like; stylet loop 142
µm long. Posterior leg 710 µm long, hind coxa with a group of translucent pores, claw
without denticle, its digitules about as long as claw; quinquelocular and trilocular disc
pores on venter with a submarginal band of cruciform pores and quinquelocular pores,
and with some multilocular pores with up to 10-loculi in transverse bands on abdominal
sternites 5-8; macrotubular ducts on submarginal band on abdomen, thorax and head;
hair-like setae on venter longer than those on dorsum.
Dorsum: Numerous quinquelocular and trilocular disc pores on dorsum, these form
transverse bands or groups; macrotubular ducts on transverse rows on the segments;
hair-like setae short, no conical setae on dorsum except on anal lobes; anal ring oval, with
one row of pores and 8 setae, each 173 µm long; anal lobes sclerotized, each lobe with 3
dorsal conical setae, 2 setae 50 µm long, 1 seta only 15 µm long, one subapical seta half as
long as anal ring setae, and an apical seta 309–325 µm long (after Borchsenius & Danzig
(1966), with modifications).
Ecology
Host plant: Brachypodium sp., B. pinnatum, Bromus sp., B. erectus, Calamagrostis sp., Festuca
sp., Lolium sp., L. perenne, Stipa sp.
Distribution: former Czechoslovakia, Hungary, Italy, Kazakhstan, Latvia, Lithuania,
Poland, Romania, Russia.
Biology: A fairly common steppe inhabiting species, on warm rocky hills in Poland, in
Hungary in the forests. Found on leaves on grasses. Adult females collected in July in
Kazakhstan and in July and September in Poland.
Acanthococcidae 285
Greenisca erwini Kozár, 1966 (Fig. 103) Revived combination.

Greenisca erwini Kozár, in Kozár & Hippe 1996: 91.
Holotype: Female. Switzerland (Gersau), on Brachypodium sp., 15.viii.1993. Deposited

in PPI.
Lit.: Danzig, 2006: 203; Kozár, 2009: 102; Kozár & Hippe, 1996; Kozár et al., 2002:
38; Kozár & Nagy, 1998: 55; Miller & Gimpel, 2000: 202; Pellizzari & Kozár, 2011:
Eriococcus erwini; Miller & Gimpel, 2000: 202. Change of combination.

Kaweckia erwini; Kozár, 2009: 102. Change of combination.
Description
Unmounted female
Not seen.
Mounted female
Adult female elongate-oval, 3 mm long, 1 mm wide. Antennae 7 segmented, I: 44.7,
II: 45.4, III: 49.1, IV: 60.9, V: 33.2, VI: 29.0 and VII: 44.3 µm long; frontal tubercle
present; eyes circular situated on venter near margin. Anal lobes well developed with
three enlarged setae (two inside and one outside) each with 60.6 µm long and one hair
like apical setae 301.1 µm long.
Venter: Labium 87.1 µm long; stylet loop twice longer than labium; basal segment with
two pairs of setae with different sizes; median setae on apical segment spine-like. Legs
well developed, strong; lengths of segments and digitules of prothoracic legs; coxa: 70.4
µm, trochanter: 56.9 µm, femur: 166.2 µm, tibia: 137.0 µm, tarsus: 137.3 µm and claw
29.5 µm long; tarsal digitules 56.6 µm long; claw with a denticle; hind coxa with a group
of translucent pores, tibia with 4 setae. Macrotubular ducts in two sizes, scattered over
entire surface. Multilocular pores with 5-7 loculi, each 5.4 µm diameter numerous on
last abdominal segments; quinquelocular pores 4.3 µm diameter on thorax and head;
cruciform pores not numerous, on head and thorax; hair-like setae slender of various
sizes, 29.4–71.6 µm long, longer than those on dorsum. Ventral surface of anal lobe with
2 slender setae and slender suranal seta.
Dorsum: Numerous quinquelocular disc pores, each 6.0 µm diameter, rarely trilocular
pores, scattered on dorsum; macrotubular ducts in two size, larger one 28.4 µm long,
11.5 µm width, smaller ones 22.6 µm long, 5.3 µm width, scattered on thorax and head.
Microtubular ducts each 4.3 µm long, in large numbers. Anal ring oval 71.2 µm long, 46.3
µm width, with double incomplete rows of pores (4.8 µm in diameter) and 8 setae, each
105.9 µm long. Marginal enlarged setae on each segment arranged in groups of 3-5 setae
on each segment, each 52.9–60.6 µm long, minute pointed setae scattered on the surface
7.1 µm long. Cauda present (after Kozár and Hippe (1996), with modifications).
Ecology:
Host plant: Brachypodium sp., Phleum sp.
Distribution: Switzerland.
Biology: Found on leaves on grasses.
Comments
In the revision of the Kozár’s collection (PPI) we found that the G. erwini specimen
cited from Hungary by Kozár, et al. (2002: 38); and Kozár & Nagy (1998: 55) is a
misidentification, and this record belongs to Gregoporia istriensis, so here the distribution
record for that region is deleted.
Greenisca gouxi (Balachowsky, 1954) (Fig. 104)

Anaphococcus gouxi Balachowsky, 1954: 61.
Syntype: Female. France (Croix-du-Grand-Maitre, Fontainebleau), on Monilia caerulea,

08.ix.1953, 13.ix.1953, 04.x.1953, A. Balachowsky. Deposited in MNHN.
Common name: Balachowsky's felt scale.
Lit.: Ali, 1970: 76; Balachowsky, 1954: 61; Danzig, 1980: 228; 1986: 265; Foldi, 2001:
305; Hoy, 1963: 133; Koteja & Zak-Ogaza, 1981: 510; Kozár et al., 2013: 56; Kozár &
Walter, 1985: 74; Köhler, 1998: 389; Ouvrard & Kozár, 2009:103; Pellizzari & Kozár
2011: 60 (Miller et al., 2013).
Greenisca gouxi; Hoy, 1963: 133. Change of combination.

Eriococcus gouxi; Williams, 1985a: 357. Change of combination.
Greenisca gouxi; Kosztarab & Kozár, 1988: 293. Revived combination.
Acanthococcus gouxi; Miller & Gimpel, 1996: 601. Change of combination.
Eriococcus gouxi; Miller & Gimpel, 2000: 217. Revived combination.
Greenisca gouxi; Kozár, 2009: 102. Revived combination.
Description
Unmounted female
Ovisac elongate-oval, white or light cream coloured. Adult female very elongate-oval,
tapering posteriorly, yellow, 2.3–4.0 mm long, 1.0–1.6 mm wide
Mounted female
Adult female elongate-oval, 2.3–4.0 mm long, 1.0–1.6 mm wide; antennae 7 (rarely 6)
segmented, 278–309 µm long, when 6 segmented third segment longer than combination
of first and second segments; frontal tubercle present; eyes situated on venter near
margin. Anal lobes slightly developed with two enlarged setae and one spine like setae.
Acanthococcidae 287
Figure 103. Greenisca erwini Kozár 1996, female. After Kozár & Hippe (1996) with
modifications.
Venter: Labium 93 µm long and wide, with 18 setae; stylet loop 124µm long; legs long
and slender, hind leg 760 µm long, hind coxa and hind femur with a group of translucent
pores, claw without denticle, its digitules about as long as claw; multilocular pores with
3–10 loculi, transverse bands on abdominal sternites 5–8, scattered on thorax and head;
cruciform pores form a band on submarginal area; hair-like setae on venter longer than
those on dorsum.
Dorsum: Slide-mounted adult female with enlarged setae narrow, conical spices slightly
rounded or acute, largest setae on anal lobes, small thin setae scattered over entire dorsum,
7 locular, quinquelocular and trilocular disc pores scattered on dorsum; macrotubular
ducts on transverse rows on the segments; hair-like setae short, no conical setae on
dorsum except on anal lobes; anal ring oval, with one row of pores and 8 setae, each
124 µm long; microtubular ducts short, with 2 sclerotized areas, scattered sometimes
in rows on the segments, anal lobes slightly sclerotized, each lobe with 2 dorsal conical
and 1 spine like setae, 2 50 µm long, 1 only 15 µm long, one preapical seta half as
long as anal ring setae, and apical seta 285–325 µm long (after Balachowsky (1954), with
modifications).
Other stages

with strong sensory setae as in adult female. Dorsum with very small spines of equal
sizes. Usually with six setae on each segment. Tubular ducts present on dorsum. Venter
with transverse rows of four small hair-like median setae on each abdominal segment;
and two rows of small submarginal spines. Two cruciform pores present on margin of
thorax. With some quinquelocular pores on each thoracic segment, 1 near each spiracle,
plus 1 pair on frons. Stylet loop twice longer than labium. Anal ring normal and with 6
hair-like setae, cauda present.
Ecology
Host plant: Carex sp., Brachypodium sp., B. pinnatum, Bromus sp., Calamagrostis sp., Festuca
ovina, Molinia? caerulea, Poa sp.
Distribution: Bulgaria, France, Georgia, Hungary, Italy, Poland, Russia (Primor'ye,
Sakhalin Oblast), Slovakia, Sweden, Switzerland.
Biology: Adult females collected from July to September. Occurs on leaves of grass.
Acanthococcidae 289
Figure 104. Greenisca gouxi (Balachowsky, 1954), female. After Koteja & Zak–Ogaza
(1981), with first instar on top. Original.
Greenisca oreophila Pellizzari & Kozár, 2011 (Fig. 105)

Greenisca oreophila Pellizzari & Kozár, 2011:59.
Holotype: Female. Italy (Lamon, Belluno district), Poaceae, leaves, 26. 08, 1989, by
G. Pellizzari, No. 199 in Pellizzari collection. Paratypes, five adult females, on separate
slides, same data as holotype, five adult females from Agorno (Belluno district),
02.07. 1993, by G. Pellizzari, No. 495 in Pellizzari’s collection, by original designation.
Holotype and paratypes deposited in DAFNAE and one paratype deposited in PPI.
Lit.: Kozár & Hippe, 1996: 92 (Miller et al., 2013).
Description
Unmounted female
Body enclosed in a felted, whitish egg sac.
Mounted female
Body of slide-mounted specimens, elongate-oval, 3.0–4.72 mm long, 1.55–2.28 mm
wide. Antennae usually 7 segmented; segment lengths; I: 36–48, II: 31–41, III: 40–53,
IV: 62–74, V: 19–31, VI: 19–31 and VII: 38–48, all segments with few setae, segment
VII with apical setae 38–48 µm long, and with 3 sensory falcate setae, 31–36 µm long,
segments V and VI each with single falcate seta 17–36 µm long. Frontal tubercles not
seen. Eyes present on ventral margin.
Venter: Labium 3 segmented, 81–91 µm long, basal segment well developed, with two
pairs of different sized setae; stylet loop a little bit longer than labium. Legs well developed,
lengths of segments of prothoracic legs; coxa: 65–75, trochanter: 46–58, femur: 149–154,
tibia: 123–139, tarsus: 127–134, claw: 32–36; lengths of segments of mesothoracic legs;
coxa: 72–84, trochanter: 46–65, femur: 142–170, tibia: 125–144, tarsus: 127–144, claw:
34–36; lengths of segments of metathoracic legs; coxa: 82–102, trochanter: 58–72, femur:
161–171, tibia: 160–180, tarsus: 138–168, tarsal digitules knobbed: 45–57, claw: 32–34,
claw digitules: 36–40, slightly knobbed; meso- and metacoxae with spinulae on anterior
(ventral) surfaces hindcoxae with translucent pores on posterior (dorsal) surface; claws
with denticles; mesothoracic spiracles 31–38 µm in diameter, with some quinquelocular
pores at spiracular opening; quinquelocular pores 6 µm in diameter, scattered in small
numbers on much of surface, some on last abdominal segments 10 loculars; slender setae
present, scattered on abdomen; some longer median flagellate setae, present on head,
thorax and anterior abdominal segments; cruciform pores in a sparse submarginal band;
macrotubular ducts of 2 sizes; the smaller about 3–5 µm wide, 20 µm long, present on
all segments, each duct with sclerotized rim surrounding orifice, particularly discernible
on larger ducts; inner ductule of larger type of duct longer than duct, ending in a flower-
shaped gland, on venter ducts are shorter and narrower; few microtubular ducts present
around margin.
Acanthococcidae 291
Figure 105. Greenisca oreophila Pellizzari & Kozár, 2011, female. After Pellizzari & Kozár
(2011).
Dorsum: Enlarged dorsal setae 1-3 on margin of abdominal segments of 2 sizes, 13

and 20 µm long; dorsal spines 9 µm long, hair-like setae about 14 µm long scattered
all over the body; macrotubular ducts 25 µm long, 7–8 µm wide, in sparse bands on
each segments; microtubular ducts present, each about 4 µm long and 2 µm wide, with
oval orifice, scattered among dorsal setae; sclerotized quinquelocular (sometimes three
locular) pores numerous, 7 µm wide, not form groups; anal ring situated on dorsum, well
developed, 67–74 µm wide, and 78–84 µm long, with a single row of pores and 6-8 setae,
each 137–170 µm long; anal lobes well developed, not sclerotized, with 3 spine-like setae,
each 43 µm long; apical setae 270–326 µm long. Cauda present (Pellizzari & Kozár, 2011).
Ecology
Host plant: Brachypodium sp.
Distribution: Italy, Switzerland.
Biology: Adult females collected from July, August. On leaves of grass.
Comments
Other materials: Two more specimens were collected and identified as G. gouxi by F.
Kozár in Gersau, Switzerland, 08.viii.1993 and 15.viii.1993, on Brachypodium leaves, (PPI:
4221 and PPI: 4236) (Kozár & Hippe, 1996). In these specimens the number of marginal
spines is smaller, because of this they were not included into the paratype series.
Greenisca placida (Green, 1921) (Fig. 106)

Eriococcus placidus Green, 1921: 148.
Lectotype: Female. England (Kent, Thurnham), on Festuca sp., 08/09/1920.

Designated by Williams (1985a). Deposited in BMNH.
Common name: Smooth Felt Scale.
Lit.: Borchsenius & Danzig, 1966: 43; Hoy, 1963: 134; Koteja & Zak-Ogaza, 1981:
510. Williams, 1985a: 376; Kozár & Walter, 1985: 75; Kosztarab & Kozár, 1988: 294;
Pellizzari & Kozár, 2011: 63-66; Williams, 1985a: 376 (Miller et al., 2013).
Nidularia placida; Lindinger, 1933a: 116. Change of combination.

Greenisca placida; Danzig, 1959: 446. Change of combination.
Acanthococcus placidus; Miller & Gimpel, 1996: 603. Change of combination.
Eriococcus placidus; Miller & Gimpel, 2000: 302. Revived combination.
Greenisca placida; Kozár, 2009: 102. Revived combination.
Acanthococcidae 293
Figure 106. Greenisca placida (Green, 1921), female. After Williams (1985) with
modifications.
Description
Unmounted female
Ovisac strongly convex, grayish-yellow, 3.5–4.5 mm long, 2.0–2.2 mm wide.
Mounted female
Adult female elongate-oval, according to Green (1921), 2.5–3 mm long, 1.0–1.2 mm wide;
antennae 7 segmented, 280–300 µm long; frontal tubercle present, minute, just anterior
the each basal antennal segment; eyes situated on venter near margin. Anal lobes slightly
developed with two enlarged setae and one small spine-like setae (Williams, 1985a).
2 pairs of different sized setae; legs well developed, slender, hind trochanter+femur 230–
240 µm long, hind tibia 140–150 µm long, hind tarsus 160–170 µm long, claw slightly
curved, slender, 40 µm long, with a minute denticle near apex; hind coxa and hind femur
with a group of translucent pores. Macrotubular ducts in two sizes; a larger type same
as on dorsum around margin, narrower type present in median to submarginal areas;
multilocular pores with 5-7 loculi, septalocular pores generally in transverse bands on
abdominal sternites VI–VIII, others scattered on thorax and head; cruciform pores not
numerous, in narrow submarginal band from about abdominal segment V forward to
head; hair-like setae slender of various sizes longer than those on dorsum; ventral surface
of anal lobe with 2 slender setae and slender suranal seta (Williams, 1985a).
Dorsum: Quinquelocular disc pores present in single to double rows mainly at anterior
edges of segments, each pore with wide sclerotised rim and about half width of a
diameter of cup of macroduct. Macrotubular ducts fairly evently distributed, each about
25 µm long, the cup about 2-3 times as wide as diameter of setal base of small setae.
Microducts in a regular arrangement, each about 6 µm long, with ampulla, tube with inner
end swollen and inner collar. Setae in two types, apart from enlarged setae on anal lobes
there are other present about 30 µm long, on head margin in variying numbers but there
are usually one or two present; elsewhere minute pointed setae scarcely more than 20 µm
long, but often shorter, hair-like; anal ring oval, with one row of pores and 8 setae, each
145 µm long. Anal lobes conical, pointed, about twice as long as wide, slightly sclerotised,
each lobe with apical setae 220–300 µm long, with 2 dorsal conical setae 36–50 µm long,
posterior usually shorter, 32–40 µm long and outer minute seta situated, in most species
towards centre. Cauda weakly developed (after Williams (1985a), with modifications).
Ecology
Host plant: Agropyron repens, Avena flavescens, Brachypodium pinnatum, B. sylvaticum, Festuca
sp.
Distribution: former Czechoslovakia, Germany, Hungary (?), Italy, Russia (St. Petersburg
Oblast), United Kingdom.
Biology: Found on the upper surface of the leaves of grasses.
Acanthococcidae 295
Genus:
Greenoripersia Bodenheimer, 1929
Type sp.: Greenoripersia kaiseri Bodenheimer, 1929.
Comments
Based on the description and drawing of Bodenheimer (1929), this genus is treated here
as a synonym of Neoacanthococcus Borchsenius, 1948.
Genus:
Gregoporia Danzig, 1979
Type sp.: Gregoporia distincta Danzig, 1979 (= Gregoporia rosaceae Balachowsky, 1932), by
monotypy and original designation.
Lit.: Danzig, 1979: 46; 1983: 521; 1985: 110; Foldi, 2001: 305; Goux 1948: 69; Hoy,
1963: 113; Kosztarab, et al., 1986: 86; Kozár, 1983: 142; 2009: 102; Kozár et al., 1984:
5; Kozár & Walter, 1985: 75; Köhler, 1998: 390; Lindinger, 1933a: 116; Miller &
Gimpel, 1996: 603; 1999: 214; 2000: 318; Pellizzari & Kozár, 2011: 66; Tang & Hao,
1995: 508 (Miller et al., 2013).
Description
Adult female elongate-oval yellowish. Antennae 7 segmented; frontal tubercle usually
present. Labium 3 segmented, basal segment with a pair of setae; stylet loop slightly
longer than labium; legs well developed, slender, hind tibia with 5 setae, claws usually
with denticle; hind-coxae with reticulations on anterior surfaces, hindcoxae with some
translucent pores on posterior surfaces; multilocular pores with 5-10 loculi, present on
only venter; cruciform pores on ventrolateral area; macrotubular ducts present, of 2
sizes, enlarged conical setae numerous, these with truncate or sharp-pointed apex, and
present on anal lobes, and on margin hair-like setae long, longest in the median part
of body Marginal enlarged setae on each segment 5-10 in number; small, spine-like, or
setose spines in rows on all segments, and anal lobes well developed, each with 4 or 5
enlarged setae; anal ring oval, with 8 or 10 setae, each as long as the ring diameter and
with few pores present in two incomplete rows; cauda present; dorsal pores with 3-5
loculi, strongly sclerotised with special shape and in groups of 2-9 pores, situated as
logiditual bands; macrotubular ducts and microtubular ducts present.
Distribution
The Gregoporia genus contains only two species and restricted to the western part of the
Palaearctic Region.
Biology
All species are known from leaves of plants; eggs yellow. Lives on leaves of Poaceae.
Comments
The genus was synonymized by Miller & Gimpel (1996: 605), with notes that the
diagnostic characters of Gregoporia of 8 segmented antennae and dorsal quinquelocular
pores are within the range of variation expected within Eriococcus sensu lato (Miller &
Gimpel, 1996). However here we accept reestablishment of this genus by Kozár (2009)
as Gregoporia.
Key to species
1. – Number of marginal enlarged setae on each segment 5-6; quinquelocular pores in

row among marginal setae.................................................................................G. istriensis
– Number of marginal enlarged setae on each segment 7-10; quinquelocular pores
not in row among marginal setae..................................................................... G. rosaceus
Gregoporia istriensis Kozár, 1983 (Fig. 107)

Gregoporia istriensis Kozár, 1983: 142
Holotype: Female. Croatia (former Yugoslavia), Lipica, on unidentified Poaceae,

27.vi.1981, by F. Kozár. By original designation. Deposited in PPI.
Lit.: Kozár, 2009: 102; Miller & Gimpel, 2000: 601; Kozár, et al., 2002: 38; Kozár &
Nagy, 1998: 55; Tang & Hao 1995: 508, 651 (Miller et al., 2013).
Acanthococcus istriensis; Miller & Gimpel, 1996: 601. Change of combination.

Eriococcus istriensis; Miller & Gimpel, 1999: 214. Change of combination.
Gregoporia istriensis; Kozár, 2009: 102. Revived combination.
Description
Unmounted female
Adult female is elongate.
Acanthococcidae 297
Figure 107. Gregoporia istriensis Kozár, 1983, female. After Kozár (1983) with modifications.
Mounted female
Adult female elongate-oval, 3 mm long, 1 mm wide. Antennae 7 segmented, 250–260 µm
long, fourth segment longest; frontal tubercle not seen; eyes circular, situated on venter
near margin. Anal lobes well developed with three (outer) and two (inner) enlarged setae;
and one hair like apical setae.
Venter: Labium 80 µm long, stylet loop somewhat longer than labium. Legs well
developed, strong, hind legs 640 µm long, claw with denticle at near apex. Macrotubular
ducts in two sizes, scattered all surface. Multilocular pores with 5-10 loculi, numereous
around vulva, 2-5 multilocular pores around the spiracle opening. Cruciform pores not
numerous, on head and thorax. Hair like setae long. Ventral surface of anal lobe with 2
slender setae and slender suranal seta.
Dorsum: With groups of especially heavily sclerotised tri- and quinqueloculars pores in
large numbers, one group with 2-9 pores, these pores in a row among marginal enlarged
setae; dorsal multilocular pores in mediolateral and marginal areas. Macrotubular ducts
scattered on body surface. Setae short, spine-like. Microtubular ducts with 2 sclerotized
areas, scattered on body surface. Anal ring oval, with double incomplete rows of pores
and ten setae each 120 µm long. Marginal enlarged setae cylindrical, apices rounded or
truncate, marginal setae conspicuously larger than other setae on dorsum, 5 or 6 lateral
setae on margin or each abdominal segment; setae on anal lobe with truncated end.
Cauda not seen (after Kozár (1983), with modifications based on type material).
Ecology
Distribution: Croatia (former Yugoslavia), Hungary.
Biology: On the leaves. Eggs are yellow.
Comments
In the revision of the Kozár’s collection we found that the G. erwini specimen cited from
Hungary by Kozár, et al. (2002); and Kozár & Nagy (1998) was a misidentification and
this record belongs to the Gregoporia istriensis, what is a new country record.
Acanthococcidae 299
Gregoporia rosaceus (Balachowsky, 1932) (Fig. 108)

Eriococcus rosaceus Balachowsky, 1932: 233.
Syntype: Female. France (between La Turbie and Peille, Alpes-Maritimes), on Bromus

erectus, 03.09.1932, by A. Balachowsky. Notes: There are six slides containing ten
syntype specimens (Matile-Ferrero, personal communication, November 20, 1996).
Deposited in MNHN.
Lit.: Danzig, 1979: 46; 1983: 521; 1985: 110; Foldi, 2001: 305; Goux 1948: 69; Hoy,
1963: 113; Kozár, 1983: 142; 2009: 102; Kozár et al., 1984: 5; Kozár & Walter, 1985:
75; Köhler, 1998: 390; Lindinger, 1933a: 116; Miller & Gimpel, 1996: 603; 1999: 214;
2000: 318; Pellizzari & Kozár, 2011: 66; Tang & Hao, 1995: 508 (Miller et al., 2013).
Nidularia rosaceus; Lindinger, 1933a: 116. Change of combination.

Gregoporia distincta Danzig, 1979: 46.
Syntype: Female. Russia, 22/07/1976, by E. Danzig. Deposited in ZMAS.
Synonymized by Danzig, 1983: 521.
Gregoporia rosacea; Danzig, 1983: 521. Change of combination.
Acanthococcus rosaceus; Miller & Gimpel, 1996: 603. Change of combination.
Eriococcus rosacea; Miller & Gimpel, 1999: 214. Revived combination.
Gregoporia rosaceus; Kozár, 2009: 102; Ouvrard & Kozár, 2009: 102. Revived combination.
Description
Unmounted female
Adult female is elongate, pale yellow in color, 3–4 mm long, 1.5 mm wide. Ovisac is
cylindrical, color varies from white to cream (Balachowsky, 1932).
Mounted female
Adult female elongate-oval. Antennae 7 segmented, fourth segment longest; frontal
tubercle present; eyes circular, situated on venter near margin. Anal lobes very well
developed with four enlarged setae.
Venter: Labium 3 segmented, basal segment with two pairs of setae, one of them long,
hair-like, other short, spinelike, with six pair of setae on apical segment, the median
ones short, spine-like. Stylet loop somewhat longer than labium. Legs well developed,
strong, posterior coxae with pores, claw with denticle at near apex; macrotubular ducts
in two sizes, scattered all surface; multilocular pores with 5-9 loculi numereous around
vulva; about 4-8 multilocular pores around the spiracle opening; cruciform pores not
numerous, on head and thorax; hair like setae long; ventral surface of anal lobe with 2
slender setae and slender suranal seta.
Dorsum: With groups of especially heavily sclerotised tri- and quinquelocular pores in
large numbers, one group with 2-9 pores, these pores in a row among marginal enlarged
setae. Enlarged setae thin, sides concave, apices acute, marginal setae conspicuously larger
than all other setae on dorsum, very abundant on lateral margin, 8-9 on each abdominal
segment. Macrotubular ducts scattered on body surface. Microtubular ducts scattered
on body surface. Anal ring oval, with double incomplete rows of pores and ten setae.
Marginal enlarged setae on body segment in number 8-9. Setae on anal lobe truncated
end. Cauda present (after Balachowsky (1932), and Danzig (1979), with modifications
Ecology
Host plant: Brachypodium pinnatum, Bromus erectus, Cynodon sp.
Distribution: France, Italy, Russia.
Biology: On the upper surface of the leaf.
Genus:
Hispaniococcus Kozár, 2008
Hispaniococcus Kozár & Konczné Benedicty, 2008b: 253.
Type sp.: Ovaticoccus agenjoi Gómez-Menor Ortega, 1954.

Lit.: Kozár, 2009: 102 (Miller et al., 2013).
Description
Antennae 6 segmented; frontal tubercle present. Legs short, with tibia shorter than tarsus;
claw with denticle, claw digitules spinose, shorter than claw; all coxae with spinulae;
posterior coxae also with large pores; multilocular pores only on venter, each with 5
loculi; macrotubular ducts over both surfaces; microtubular ducts present; cruciform
pores absent; ventral setae short and hair-like. Anal lobes not well developed; dorsal
surface of each lobe with three dome shaped setae, ventral surface of each lobe with a
long apical seta and two shorter subapical seta, dorsal enlarged setae dome-shaped; anal
ring sclerotized, but not well developed with few anal ring pores and six strong setae, all
shorter than diameter of ring; cauda absent, macrotubular ducts heavily sclerotized, each
with inner ductules short, terminal gland not observed; microtubular ducts short.
Distribution
The genus contains only two species and restricted to the southern part of the Palaearctic
Region.
Acanthococcidae 301
Figure 108. Gregoporia rosaceus (Balachowsky, 1932), female. After Danzig (1979) with
modifications.
Biology
Lives on Artemisia sp.
Key to the species
1. – Enlarged setae on dorsum dome shaped; anal ring setae shorter than
anal ring width....................................................................................................... H. agenjoi
– Enlarged setae on dorsum elongate; anal ring setae longer than anal
ring width.......................................................................................................H. oligacanthus
Hispaniococcus agenjoi (Gómez-Menor Ortega, 1954) (Fig. 109)

Gymnococcus agenjoi Gómez-Menor Ortega, 1954: 142.
Lectotype: Female. Spain (Alcalá de Henares, Madrid), on Artemisia sp., iii.1952, by

R. Agenjo. Lectotype and two paralectotype slides (with the same data as lectotype,
but the year of collection is different, deposited in MNCN.
Lit.: Boratinsky, 1958: 174; Hoy, 1963: 183; Kozár, 2009: 102; Kozár & Konczné
Benedicty, 2008b: 254; Kozár & Walter, 1985: 75; Köhler, 1998: 393-394; Martin-
Mateo, 1985: 94; Miller & Gimpel, 2000: 430 (Miller et al., 2013).
Ovaticoccus agenjoi; Boratynski, 1958: 174. Change of combination.

Hispaniococcus agenjoi; Kozár, 2009: 102. Change of combination.
Description
Unmounted female
Body of adult female oval, cuticle membranous, whitish in color.
Mounted female
Body of slide-mounted specimens oval, 2.02–2.28 mm long, 1.81–1.92 mm wide.
Antennae 6 segmented, with some sign of constriction on the third segments; I: 25–28,
II: 18–20, III: 30–33, IV: 16–17, V: 13–10, VI: 27–22 µm, all segments with few setae,
segment VI with apical setae 44 µm long, and with 3 sensory falcate setae, 18–20 µm
long, segments IV and V each with single falcate seta 11–13 µm long. Apical segment
with two 4 µm long coeloconic sensilla. Frontal tubercles present, 3 µm in diameter. Eyes
present on venter. Anal lobes not well developed, not sclerotized, with 3 spine-like dome
shaped setae, as long as marginal ones; apical setae each 100–110 µm long.
Venter: Labium 3 segmented, 80–84 µm long, basal segment not well developed, with
two pairs of setae, total number of setae 9 pairs, apical median setae half long of the
longest lateral ones; stylet loop not seen. Legs well developed, lengths of segments of
prothoracic legs: coxa: 40–45, trochanter: 30–33, femur: 77–78, tibia: 58–60, tarsus: 65,
Acanthococcidae 303
Figure 109. Hispaniococcus agenjoi (Gómez–Menor Ortega, 1954), female. After Kozár &
Konczné Benedicty (2008) with modifications.
claw: 26–27 µm long; lengths of segments of mesothoracic legs; coxa: 43–46, trochanter:
31–32, femur: 60–61, tibia: 54, tarsus: 68–73, claw: 25–27 µm long; lengths of segments
and digitules of mesothoracic legs; coxa: 60–62, trochanter: 37, femur: 71–73, tibia:
57–60, tarsus: 71–81, tarsal digitules knobbed, 32–40, claw 26–27, claw digitules 10–
13 µm long, spinose, shorter than claw; all coxae with spinulae on anterior (ventral)
surfaces, hindcoxae with translucent pores and spinulae on posterior (dorsal) surface;
claws with denticles; all legs with few flagellate setae and with sensory pore at base of
each tarsus. Mesothoracic spiracles 23–25 µm in diameter, with several quinquelocular
pores at spiracular opening. Quinquelocular pores 6 µm in diameter, scattered in medium
numbers on much of surface; setae slender short, scattered on abdomen, some longer
flagellate setae, present on head and thorax. Macrotubular ducts about 4 µm wide, 26
µm long, present on all segments, each duct with sclerotized rim surrounding orifice,
particularly discernible on larger ducts; inner ductule shorter than duct; few microducts
present around body margin.
Dorsum: Setae on last abdominal segments robust, dome shaped spine-like, of one sizes,
9–13 µm long, present mostly in single rows across segments, 3-4 present on margin
of each abdominal segment in loose groups. Macrotubular ducts about 10-20 on each
segments, 50 µm long and 6 µm wide; microtubularducts short, sclerotized, each about 4
µm long with normal orifice, scattered among dorsal setae. Quinquelocular pores similar
as those on venter, scattered on margin. Anal ring situated on dorsum, well developed,
sclerotized 38–40 µm wide, 35 –38 µm long, with single row of pores and 6 strong setae
each 36–30 µm long (Gómez-Menor Ortega (1954).
Ecology
Host plant: On Artemisia sp., Kalidium sp.
Distribution: Spain.
Biology: This species occurs under the bark of the branches.
Hispanicoccus oligacanthus (Danzig, 1972) (Fig. 110) Redescription,

Combination nova
Rhizococcus oligacanthus Danzig, 1972b: 341.
Holotype: Female. Mongolia (South Gobi Aymag, near Dund-Gol), on Kalidium

gracile on stem near roots, 20.viii.1969, by I. Kerzhner. By original designation.
Deposited in ZMAS.
Lit.: Danzig, 1974: 70; Kozár, 2009: 94; Kozár & Walter, 1985: 75; Miller & Gimpel,
Acanthococcus oligacanthus; Miller & Gimpel, 1996: 602. Change of combination.

Eriococcus oligacanthus; Miller & Gimpel, 1999: 214. Change of combination.
Acanthococcidae 305
Figure 110. Hispaniococcus oligacanthus (Danzig, 1972), female. After Danzig (1972b)
with modifications.
Rhizococcus oligacanthus; Köhler, 1998: 393. Revived combination.

Eriococcus oligacanthus; Miller & Gimpel, 2000: 430. Revived combination.
Anophococcus oligacanthus; Kozár, 2009: 94. Change of combination.
Description
Unmounted female
Wide oval, convex, 3 mm long, 2.5 wide.
Mounted female
Adult female oval. Antennae 7 segmented. Frontal tubercle present. Anal lobes weakly
sclerotized.
Venter: Labium 3 segmented, with 6 pairs of setae on apical segment. Legs short,
strong, with wide posterior coxae, coxae and femur with pores, tarsal digitules longer
than claw, slightly capitated, claw digitules short, hair-like. Cruciform pores numerous
in submarginal and submedial area on each thoracic segments, and a few on submedial
part of head. Quinquelocular pores numerous on venter, forming large groups around
spiracles.
Dorsum: Anal ring developed. Enlarged setae short, blunted, forming rows and bands
on dorsum, anal lobe with three spines. Marginal row of spines absent. Macrotubular
ducts and microtubular ducts few, scattered on dorsum (after Danzig (1972b) with
modifications).
Ecology
Host plant: Kalidium gracile.
Biology: On stem near roots.
Acanthococcidae 307
Genus:
Hujinlinococcus Kozár & Wu Genus nova
Type sp.: Eriococcus nematosphaerus Hu, Xie & Yan, 1981: 75.
Lit.: Kozár, 2009: 102; Tang & Hao, 1995: 453 (Miller et al., 2013).
Description
According to Hu et al. (1981), body crooked in shape, about 2.4 mm in length. Antennae
7 segmented; labium 3 segmented, with well developed segments and a weakly developed
basal segment with two pairs of hair-like setae; legs well-developed, claw with a denticle
near the apex; tarsal and claw digitules longer than claw clavate, spiracles often with a
few associated disc pores; discoidal pores usually quinquelocular, cruciform pores on
prosomal venter in a marginal band; tubular ducts of 2 types: micro- and macrotubular
ducts; microtubular ducts slender, scattered or form transverse rows or bands on dorsum,
macrotubular ducts form row; or in transverse sparse bands on dorsum; enlarged setae
of 2 distinct shapes, one nipple shaped, sides convex, apices rounded, base broad,
second type cylindrical, sides slightly concave, apices truncate, enlarged setae abundant
over dorsal surface; anal lobes sclerotized, with teeth on inner margin, apparently with 2
enlarged setae; hair-like setae on venter only; anal ring well developed, sclerotised with
partly double row of pores and 8 anal ring setae, each anal lobe with a long apical seta
and usually with 3 elongate dorsal enlarged setae (after Hu et al., 1981).
Distribution
Palearctic-Oriental, with 1 species.
Biology
Feed on grasses.
Comments
Because the great variation of several characters in E. nematosphaerus, a new genus is
established in hope that more new species would be discovered in the future.
Hujinlinococcus nematosphaerus (Hu, Xie & Yan, 1981) (Fig. 111)

Combination nova
Eriococcus nematosphaerus Hu et al., 1981: 75.
syntype: Female. China, on Phyllostachys sp. Syntypes, both sexes. Deposited in SIEC.
Notes: The original publication states that "the holotype specimens of female and
male are preserved in Shanghai." Because more than one holotype is mentioned, the
series must be considered syntypic.
Lit.: Fang et al., 2001:104; Hua, 2000: 137; Kozár, 2009: 93; Miller & Miller, 1996: 602;
Miller & Gimpel, 2000: 279; Tang & Hao, 1995: 453; Tao, 1999: 32; Wang, 2001: 208
Acanthococcus nematosphaerus; Miller & Gimpel, 1996: 602. Change of combination.
Description
Unmounted female
Adult female body crooked in shape, male body slender.
Mounted female
Body elongate oval. 2.4 mm long. Antennae 7 segmented; anal lobes strongly developed,
about twice as long as wide, rounded at apex, each with 3 enlarged setae on dorsal surface
and five sclerotised tubercles.
Venter: Legs well developed; tarsus longer than the tibia, with a pair of setae inflated
at the apex; claw with denticle close to apex; claw digitules longer than claw slightly
knobbed. Setae flagellate and long. Quinquelocular pores found in rows on last abdominal
segments, some around spiracles. Cruciform pores present on submargin. Macrotubular
ducts scattered on body. Microtubular duct present in a submarginal band.
Dorsum: Enlarged setae of 2 distinct shapes, one nipple shaped, sides convex, apices
rounded, base broad, second type cylindrical, sides slightly concave, apices truncate,
enlarged setae abundant over dorsal surface. Macrotubular ducts in 2 sizes, long. Pores
on the abdominal segments, scattered on thorax and head. Microtubular ducts, scattered
throughout. Anal ring well developed, with 8 setae (after Hu et al. (1981) and Tang & Hao
(1995) with modifications).
Other stages

Body of slide-mounted specimens, oval, 0. 50 mm long, 0.24 mm wide. Antennae six
segmented, apical three segments with strong sensory setae as in adult female. Dorsum
with large spines in median longitudinal rows, the median andominal spines larger the
others of equal sizes. Dorsum with six rows of setae. Macrotubular ducts absent. Venter
Acanthococcidae 309
Figure 111. Hujinlinococcus nematosphaerus (Hu, Xie & Yan, 1981), female, after Tang &
Hua (1995). First instar on top right, after Hu et al. (1981) with modifications.
with transverse rows of four small hair-like median setae on each abdominal segment;
and two rows of submarginal setae. Cruciform pores not shown, or not mentioned.
With some quinquelocular pores on each thoracic and abdominal segment, 1 near each
spiracle, plus 1 pair on frons. Stylet loop reaching the posterior coxae. Anal lobe with four
conical setae and with long apical setae. Anal ring normal and with 6 hair-like setae (after
Hu et al. (1981), with some modifications).
Ecology
Host plant: Phyllostachys sp., P. glauca, P. nigra, P. nuda, P. praecox, P. propinqua, P. pubescens,
P. rubella and P. viridis.
Biology: This species has two generations a year. It overwinter in the adult female stage.
Each female lays 9-587 eggs. First instars of the first generation occur in late April to
late May; the second generation occur in late July to early August. Males of the first
generation occur in June and males of the second generation occur in late September to
early October.
Genus:
Kaweckia Koteja & Zak-Ogaza, 1981
Type sp.: Eriococcus glyceriae Green, 1921: 146.
Lit.: Danzig, 1980: 228; Kawecki, 1985: 32; Kosztarab et al., 1986: 9; Kosztarab
& Kozár, 1988: 294; Koteja, 2000a: 242; Kozár, 2009: 583; Kozár & Walter, 1985:
75; Köhler, 1998: 391; Miller & Gimpel, 1996: 601; Miller & Gimpel, 2000: 214;
Schmutterer, 2008: 84; Tang & Hao, 1995: 510, 652; Tereznikova, 1981: 52 (Miller et
al., 2013).
Description
Living female red. Adult female elongate oval, almost parallel-sided. Antennae 6 or 7
segmented, base at body margin; frontal tubercles present; labium 3 segmented generally
wide; stylet loop extends to line between mid-coxae. Legs well developed, hind tibia with
4 setae, 2 on inner side, coxae generally with translucent pores; claw usually with a denticle
at apex; tarsal and claw digitules longer than claw clavate. Spiracles often surrounded by
sclerotised area, with a few associated disc pores. Discoidal pores generally 3-9 loculi,
on both surfaces. Cruciform pores on dorsum and venter; micro- and macrotubular
ducts on both surfaces. Enlarged setae often with truncate apex on abdominal segments
and on anal lobes; dorsal setae small needle-like, ventral setae in medial area enlarged,
Acanthococcidae 311
hair-like. Anal ring circular, broad, twice smaller than in Greenisca. Anal lobes prominent
(Kosztarab & Kozár, 1988).
Distribution
Transpalearctic genus, with 5 species.
Biology
On stem, root collar, and in leaf sheaths of various grasses, often below soil surface; eggs
overwinter, males unknown.
Key to species
1. – Anal lobe with less than three enlarged setae.................................................................2

– Anal lobe with three enlarged setae..................................................................................3
2. – Anal lobe with one enlarged setae.................................................................. K. matesovae
– Anal lobe with two enlarged setae.....................................................................K. hellenica
3. – Cruciform pores on dorsum scattered on thorax; all dorsal setae about the same
shape and size..................................................................................................... K. orientalis
– Cruciform pores on dorsum form bands on thorax; dorsal setae various shapes
and sizes...............................................................................................................................4
4. – Marginal enlarged setae on abdominal segment VII, wide and short (about
three times longer than width of base) and not reaching posterior segment
of thorax; high number of multilocular pores around spherical atrium (about
7-14)....................................................................................................................... K. glyceriae
– Marginal enlarged setae on abdominal segment VII, narrow and long (about
four-five times longer than width of base) and reaching posterior segment
of thorax; small number of multilocular pores around spherical atrium
(about 2-4).............................................................................................................K. vanensis
Kaweckia glyceriae (Green, 1921) (Fig. 112, 113)

Eriococcus glyceriae Green, 1921:146.
Holotype: Female. England (Blakeney Point), on Glyceria maritima (=Puccinellia

maritima), ?.vii.1920, by E.E. Green. Lectotype female, by subsequent designation
Williams, 1985a:367-368. Deposited in BMNH. Notes: In addition to the lectotype
there are two adult female paralectotypes on the same slide in the BMNH (D. J.
Williams, personal communication, June 19, 1996). Type material also in UCRC (Gill
& Fromer, 1979).
Common Name: Grass felt scale.
Lit.: Danzig, 1980: 228; 1986: 265; Fetykó et al., 2010: 296; Foldi, 2001: 305; Goux,
1937: 93; Hoy, 1963: 133; Kawecki, 1985: 32; Ben-Dov & Kosztarab, 1996: 9; Koteja,
1974a: 297; 2000a: 242; Kozár, 2009: 583; Kozár et al., 2013: 56; Kozár & Walter,
1985: 75; Kwon & Han 2003: 156; Pellizzari & Kozár, 2011: 66; Tereznikova, 1981:
52; Tang & Hao, 1995: 510, 652; Zahradník, 1977: 121 (Miller et al., 2013).
Nidularia glyceriae; Lindinger, 1933a: 116. Change of combination.

Greensica glyceriae; Borchsenius, 1949: 368. Change of combination.
Eriococcus glyceriae; Hoy, 1963: 133. Revived combination.
Greenisca baltica; Rasina, 1966: 28.
Holotype: Female. Latvia (Rigas Jurmala), on Festuca rubra, 04.viii.1948, by A. Rasina,
by original designation. Deposited in Riga: Museum of the Plant Protection Institute,
Latvia. Synonymy by Danzig, 1980: 230.
Kaweckia baltica; Koteja & Zak-Ogaza, 1981: 506. Change of combination.
Kaweckia glyceriae; Koteja & Zak-Ogaza, 1981: 506. Change of combination.
Eriococcus glyceriae; Williams, 1985a: 367. Revived combination.
Kaweckia glyceriae; Kosztarab & Kozár, 1988: 294. Revived combination.
Acanthococcus glyceriae; Miller & Gimpel, 1996: 601. Change of combination.
Kaweckia glyceriae; Köhler, 1998: 391. Revived combination.
Eriococcus glyceriae; Miller & Gimpel, 2000: 214. Revived combination.
Kaweckia glyceriae; Kozár, 2009: 583. Revived combination.
Description
Unmounted female
Ovisac compact, white to yellowish, up to 4 mm long, 1.6 mm wide; covers female
completely.
Mounted female
Adult female elongate, twice as long as wide, 3 mm long, 1.0–1.7 mm wide, bright pink
(eggs are also bright pink), covered with powdery wax. Antennae 7 segmented, 217–248
µm long, each segment covered with a few, hair-like setae; apical segment with apical seta
and 3 sensory falcate setae. Frontal tubercle present. Eyes situated on venter near margin.
Anal lobes broad, subapical setae 88–99 µm long, each with 3 enlarged setae on dorsal
surface.
Venter: Labium 3 segmented, median setae on apex spine-like; stylet loop reaches line
between mid-coxae. Legs well developed; hind legs 440 µm long; hind coxa with a group
of translucent pores; hind femur 112–140, tibia 87–110, tarsus 110–117 µm long; claw
with a small denticle, tarsal digitules knobbed, claw digitules slightly knobbed. Tibiae
of meso and metathoracic legs each with 4 setae (median seta absent). Oval disc pores
with 5, 7 and 9 loculi, scattered, found in rows on last abdominal segments. Setae in
median areas flagellate, long, setae in lateral areas spine like, some quinquelocular pores
found around spiracles. Microtubular duct present in a submarginal band. Macrotubular
Acanthococcidae 313
Figure 112. Kaweckia glyceriae (Green, 1921), female, after Koteja & Zak-Ogaza (1981)
with modifications. First instar on top right, original.
ducts of two sizes, scattered on body surface and forming rows and bands on abdominal
segments. Cruciform pores present on submargin.
Dorsum: Dorsal setae few, hair-like, 8-11 enlarged cylindrical setae with truncate tip and
broadened at base form a row on each dorsal margin of abdominal segments IV to VII.
and 19–20 µm long. Microtubular ducts 4 µm long, scattered throughout. Oval disc pores
with 5, 7 and 9 loculi, found in rows on the edge of the segments. Anal ring small, partly
with double rows of pores and with 8, rarely 6 setae, each 65–95 µm long. Anal lobes
broad, subapical setae 88–99 µm long, apical setae 195–235 µm long, 3 enlarged setae
with truncate tip, each 20–29 µm long, on dorsum of each lobe. Cauda not seen (After
Koteja & Zak-Ogaza (1981)).
Other stages

with strong sensory setae as in adult female. Stylet loop long, reaches posterior coxae.
Dorsum with hair-like setae in two median longitudinal rows. The marginal row of setae
consist on last abdominal segments one strong sharp-pointed spines, the others are hair-
like. Microtubular ducts in a marginal row. Venter with transverse rows of four small
hair-like median setae on each abdominal segment; and two rows of small submarginal
setae. Some cruciform pores present on thorax margin of venter. With one multilocular
pore, on each thoracic and abdominal segment, one near each spiracle, plus some on
frons, one in middle-line of thorax and one on each segment of abdomen, forming two
longitudinal rows. Anal lobe with three conical setae and with long apical setae. Anal ring
normal and with 6 hair-like setae. This stage was described by Schmutterer (1952) and
Tereznikova (1981).
Mounted second instar female (Fig. 113)

with strong sensory setae as in adult female. Frontal tubercle present near base of
antennae. Stylet loop long, reaching posterior coxae. Dorsum with hair-like setae in six
median longitudinal rows. The marginal row of setae consist on last abdominal segments
strong spines 2 truncated spines on last two segments and 1on two other segments, the
others are hair-like. Microtubular ducts in a marginal row and scattered on dorsum. Venter
with transverse rows of hair-like setae on each abdominal segment. Some cruciform
pores present on thorax margin of venter, and about 30 scattered on midthorax of
dorsum. Multilocular pores with 6-9 loculi, scattered on venter and dorsum too. Stylet
loop reaching the posterior coxae. Anal lobe with three conical setae and with long apical
setae. Anal ring normal and with 6 hair-like setae (after Williams (1985a)).
Acanthococcidae 315
Figure 113. Kaweckia glyceriae (Green, 1921), second instar female. Original.
Ecology
Host plant: Agropyron repens, A. tsukushiense var. transiens, Agrostis sp., A. canina, A.
capillaris, A. gigantea, Anthoxanthum sp., Arrhenatherum elatius, Brachypodium sp., Calamagrostis
sp., Carex sp., Corynephorus canescens, Elymus arenarius, Elytrigia sp., Festuca ovina, Festuca sp.,
F. rubra, F. sulcata, F. supina, Glyceria sp., G. maritima, Hierochloa odorata, Koeleria sp., Lolium
perenne, Phleum pratense, Piptatherum songaricum, Poa angustifolia, P. compressa, P. pratensis,
Puccinellia sp., Secale cereale, Sedum sp., Triticum aestivum, T. vulgare.
Distribution: Austria, China (Inner Mongolia), former Czechoslovakia, France,
Germany, Hungary, Italy, Kazakhstan, Latvia, Poland, Romania, Russia (Karelia AR,
Primor'ye, St. Petersburg (= Leningrad) Oblast), South Korea, Ukraine (Krym (=Crimea)
Oblast, Odessa Oblast), United Kingdom, former Yugoslavia.
Biology: Lives on leaves, under leaf sheaths and on root crown of grasses under soil
surface. This species has one generation per year and overwinters in egg stage (Kosztarab
& Kozár, 1988), and each female lays 15–127 eggs. First instars hatch in spring, females
become adult in July and August. According to Tereznikova (1975) first instars hatch in
summer, this is not consistent with the data of others (Koteja & Zak-Ogaza, 1981). This
species is considered a pest in Kazakhstan (Kosztarab & Kozár, 1988), and become very
common in Hungary on highways (Kozár, 2009).
Kaweckia hellenica (Kozár) 1999 (Fig. 114)

Greenisca hellenica Kozár in Pellizzari & Kozár, 1999: 28.
Holotype: Female. Greece (Ialissos), unknown host, 20.x.1996, by B. Nagy. Holotype

female, by original designation. Deposited in PPI.
Lit.: Kozár, 2009: 102; Miller & Gimpel, 2000: 227; Milonas et al., 2008: 143; Pellizzari
& Kozár, 1999: 28 (Miller et al., 2013).
Eriococcus hellenica; Miller & Gimpel, 2000: 227. Change of combination.

Kaweckia hellenica; Kozár, 2009: 102. Revived combination.
Description
Unmounted female
Unknown.
Mounted female
Adult female elongate, 3.2 mm long, 1.5 mm wide. Antennae 7 segmented, 281 µm long,
the length of the segments, I: 60, II: 38, III: 58, IV: 46, V: 17, VI: 26 and VII: 36 µm long,
each segment covered with a few, hair-like setae. Frontal tubercle present. Eyes circular.
Anal lobes well developed, each with 5 enlarged setae on dorsal surface.
Acanthococcidae 317
Figure 114. Kaweckia hellenica Kozár, 1999, female. After Pellizzari & Kozár (1999) with
modifications.
Venter: Labium 3 segmented, 62 µm long, stylet loop three times longer than labium.
Legs well developed; hind coxa with a group of translucent pores; hind coxa 84, hind
femur 135, hind tibia 144, hind tarsus 147 µm long; claw with a small denticle at apex,
tarsal digitules knobbed, 43 µm long, claw digitules slightly knobbed. Tibiae of meso and
metathoracic legs each with 4 setae (median seta absent). Diameter of anterior spiracles
43 µm long. Oval disc pores with 5, 7 and 10 loculi, each 7–8 µm in diameter, scattered
found in rows on last abdominal segments, 10 locular discodial pores numerous around
the genital opening. Cruciform pores sparse on body margin. Setae in median areas
flagellate, long, setae in lateral areas spine like. Some five-locular pores found around
spiracles. Macrotubular ducts two sizes, scattered on body surface and form rows and
bands on abdominal segments. Microtubular duct present in a submarginal band.
Dorsum: With 1-6 marginal enlarged setae with truncate apex, each 18–32 µm long,
on each abdominal segments; small setae 14 µm long, scattered on body. Multilocular
pores numereous, scattered on all body, not in group; cruciform pores in high number
on thorax. Macrotubular ducts two sizes 14–18 µm long, scattered on body surface and
form rows and bands on abdominal segments. Microtubular ducts 5 µm long scattered
throughout body. Anal ring small 41 µm diameter, partly with double row of pores and
with 6 setae, each 85.5 µm long. Anal lobes well developed, apical setae 173 µm long.
Cauda not seen (after Pellizzari & Kozár, 1999).
Ecology
Host plant: Unknown.
Distribution: Greece.
Biology: Unknown.
Kaweckia matesovae (Danzig, 2006) (Fig. 115) Combination nova

Greenisca matesovae Danzig, 2006: 203.
Holotype: Female. Russia (Saratov Prov. railway station Ozinki), the steppe, from
stems of Agropyron fragile, 7.ix.1969, by G. Matesova, type no. 3461. Paratypes 3
females in separate slides with labels identical to those of the holotype. Deposited
in ZMAS.
Lit.: Danzig, 2006: 203 (Miller et al., 2013).
Description
Unmounted female
Adult females in the leaf sheaths on the root crown.
Acanthococcidae 319
Figure 115. Kaweckia matesovae (Danzig), female. After Danzig (2006) with modifications.
Mounted female
Body elongate oval, 3 mm long. Antennae 7 segmented. Eyes situated on venter near
margin. Anal lobes wide developed, each with 1 short, truncated setae and two long
slender setae.
Venter: Legs small, well developed. Metathoracic coxae with pores. Setae flagellate, long.
Multilocular pores each 7 µm in diameter and generally with 7 loculi, distributed in sparse
bands on all abdominal segments and scattered on thorax and few on head. Macrotubular
ducts similar size, scattered on all body. Microtubular ducts few. Cruciform pores very
few on thorax, and situated on body margin.
Dorsum: Marginal dorsal setae on preanal segments absent. Dorsal setae slender, often
curved but stiff, arranged in transverse rows across each body segment, rows irregular
on head. Macrotubular ducts similar size, scattered throughout dorsum, generally in
segmental bands. Microtubular scattered over dorsum. Cruciform pores numerous
situated on median part of thorax and first abdominal segments (after Danzig (2006).
Ecology
Host plant: Agropyron fragile.
Biology: Unknown.
Kaweckia orientalis (Borchsenius, 1956) (Fig. 116) Redescription

Greenisca orientalis Borchsenius, 1956: 676.
Syntype: Female. North Korea: Sariwon, on undetermined Poaceae, 19.vii.1950, by

N. Borchsenius. Deposited in ZMAS.
Lit.: Borchsenius & Danzig, 1966: 42; Danzig, 1980: 228; 2006: 203; Kozár, 2009:
102; Kozár & Walter, 1985: 75; Köhler, 1998: 391; Kwon & Han, 2003: 156; Miller
& Gimpel, 2000: 227; Milonas et al., 2008: 143; Paik, 1978:422; Rasina, 1966: 23;
Pellizzari & Kozár, 1999: 28; Tang & Hao, 1995: 510, 652; Tao, 1999: 35; Wang, 2001:
225 (Miller et al., 2013).
Kaweckia orientalis; Koteja & Zak-Ogaza, 1981: 508. Change of combination.

Acanthococcus orientalis; Miller & Gimpel, 1996: 602. Change of combination.
Eriococcus orientalis; Miller & Gimpel, 1999: 215. Change of combination.
Kaweckia orientalis; Kozár, 2009; 102. Revived combination.
Description
Unmounted female
Acanthococcidae 321
Figure 116. Kaweckia orientalis (Borchsenius, 1956), female. Original.

Mounted female
Body elongate oval, 3.1–3.2 mm long, 1.50 mm wide. Antennae 7 segmented, 250–254
µm, length of segments: I: 34–41, II: 41–43, III: 45–48, IV: 45–48, V: 19–22, VI: 19–24,
and VII: 36–39 µm long, each segment covered with a few, strong hair-like setae (except
third segment); apical segment with apical seta 46–50 µm long; apical segment also with
3 sensory falcate setae, each 27–31 µm long; segment VI with 1 sensory falcate seta,
27–29 µm long, segment V with 1 sensory falcate seta, 19–24 µm long. Frontal tubercle
present. Eyes situated on venter near margin. Anal lobes slightly developed, each with 3
enlarged setae (inside two shorter 17–18 µm, outside one larger 41 µm long) plus a few
microtubular ducts on dorsal surface; apical seta 308–312 µm.
short. Stylet loop medium size. Legs well developed; lengths of segments of prothoracic
legs; coxa: 53–55, trochanter: 41–44, femur: 139–143, tibia: 115–120, tarsus: 118 µm
long; lengths of segments of mesothoracic legs; coxa: 63–67, trochanter: 45–48, femur:
139–144, tibia: 115–120, tarsus: 120; lengths of segments of metathoracic legs; coxa:
74–77, trochanter: 45–48, femur: 144–147, tibia: 132–134, tarsus: 125–140 µm long.
Claw length 32–35 µm long, claw digutules 41–45, tarsal digitules 56–62 µm long. Meso-
and metathoracic coxae each with spinulae on ventral surface and metathrocic coxae
with pores. Tibia of meso and metathoracic legs each with 4 setae, tarsi each with 6
setae. Diameter of spiracles 41 µm; posterior spiracles slightly larger than anterior. Setae
flagellate, long. Multilocular pores each 7 µm in diameter and with 7-9 loculi, distributed
in sparse bands on all abdominal segments and scattered on thorax and few on head.
Macrotubular ducts each 3–6 µm wide and 15–18 µm long, scattered on all body.
Microtubular ducts few, each 6 µm long and. Cruciform pores very few on thorax, each
3.0–5.0 µm in diameter and situated on body margin.
Dorsum: Marginal dorsal setae long, truncated each 28–45 µm forming a marginal
row on last 5 abdominal segments. Dorsal setae short, flagellate each seta 10–17 µm
long; arranged in transverse rows across each body segment, rows irregular on head.
Macrotubular ducts, each 3–5 µm wide and 15–18 µm long, scattered throughout dorsum,
generally in segmental bands. Microtubular ducts short, each 6 µm long, scattered over
dorsum. Cruciform pores each 5 µm in diameter, situated median part of thorax and
first three abdominal segments. Anal ring strongly sclerotized, 46 µm in diameter, with
8 setae, each 60–65 µm long; anal ring situated on margin of dorsum. Cauda not seen
(Redescription based on type material).
Ecology
Distribution: North Korea.
Biology: Unknown.
Acanthococcidae 323
Kaweckia vanensis Kaydan Species Nova (Fig. 117 )
Holotype: Female. Turkey (Van-Başkale road, N: 38°20’727’’, E: 043° 45’230”,

1854 m altitude), on plant from Poaceae, M. B. Kaydan, 09.vi.2005, (KPCT: 1709).
Paratypes: 3 adult females, on same slide, with same data as holotype (KPCT: 1709); 3
adult female on one slide, Turkey, Van-Başkale road, N: 38°20’727’’, E: 043° 45’230”,
1854 m altitude, on plant from Poaceae, M. B. Kaydan, 09.vi.2005, (KPCT: 1708);
5 adult females (on one slides), Turkey, Van-Başkale road, N: 38°20’727’’, E: 043°
45’230”, 1854 m altitude, on plant from Poaceae, M. B. Kaydan, 09.vi.2005, (KPCT:
1705); 2 adult females, Turkey, Van-Başkale road, N: 38°18’395’’, E: 043° 48’663”,
1989 m altitude, on plant from Poaceae, M. B. Kaydan, 14.vi.2006, (KPCT: 2975).
Description
Unmounted female
Mounted female
270–280 µm, length of segments: I: 45.0–55.0, II: 37.5–45.0, III: 55–65, IV: 47.5–55.0, V:
22.5–25.0, VI: 22.5–25.0, and VII: 35.0–37.5 µm long, each segment covered with a few,
strong hair-like setae (except third segment); apical segment with apical seta 40–45 µm
VI with 1 sensory falcate seta 27.5–32.5 µm long, segment V with 1 sensory falcate
seta 15.0–17.5 µm long. Frontal tubercle present. Eyes situated on venter near margin.
Anal lobes slightly developed, each with 3 enlarged setae, each 22.5–32.5 µm plus 4-6
microtubular ducts on dorsal surface; apical seta 190–220 µm; ventral hair-like subapical
seta 75–85 µm long.
Venter: Labium 107.5–110.0 µm long, 95–105 µm wide, median setae on apex of labium
short, 7.5–10 µm long. Stylet loop medium size. Legs well developed; lengths of segments
and digitules of prothoracic legs: coxa: 60–70, trochanter: 45–55, femur: 135–140, tibia:
102.5–115, tarsus: 120–135 and claw: 30.0–32.5, trochanther + femur: 177.5–195, tibia
+ tarsus: 220–235, tarsal digitules: 45–50, claw digitules 30 µm long; lengths of segments
and digitules of mesothoracic legs; coxa: 60.0–67.5, trochanter: 55, femur: 135–145,
tibia: 110–135, tarsus: 115–125 and claw: 32.5–35.0, trochanther + femur: 185–195, tibia
+ tarsus 220–245, tarsal digitules 47.5–52.5, claw digitules 32.5–37.5 µm long; lengths
of segments and digitules of metathoracic legs; coxa: 75–85, trochanter: 55–60, femur:
140–155, tibia: 125–135, tarsus: 130–135 and claw: 35, trochanther + femur: 197.5–
210.0, tibia + tarsus: 260–285, tarsal digitules 47.5–50.0, claw digitules 35.0 µm long.
Meso- and metathoracic coxae each with spinulae on ventral surface and metathrocic
coxae with pores. Tibia of meso and metathoracic legs each with 4 setae, tarsi each with
6 setae. Length of spiracles 55–70 µm; diameter of spiracular peritreme 25–32.5 µm,
posterior spiracles slightly larger than anterior. Setae flagellate, long, each 15–90 µm long.
Multilocular pores each 5–7 µm in diameter and with 3, 5, 7 and 9 loculi, distributed
in sparse bands on all abdominal segments and scattered on thorax and few on head.
Macrotubular ducts each 3–6 µm wide and 15.0–17.5 µm long, scattered on all body.
Microtubular ducts, each 3–6 µm long and situated on body margin. Cruciform pores
very few on thorax, each 3.0–5.0 µm in diameter.
Dorsum: Marginal setae long, truncated, each 25–45 µm forming a marginal row on
abdominal segments. Dorsal setae short, flagellate each seta 15.0–50.0 µm long; arranged
in transverse rows across each body segment, rows irregular on head. Macrotubular ducts,
each 6.0–7.5 µm wide and 15.0–17.5 µm long, scattered throughout dorsum, generally in
segmental bands. Microtubular ducts short, each 3.0–4.0 µm long, scattered over dorsum.
Cruciform pores each 3.0–5.0 µm in diameter, situated median part at thorax and first
two abdominal segments in groups. Anal ring strongly sclerotized, with 16–19 pores
on each side, 42.5–45.0 µm in diameter, with 6 setae, each 62.5–80 µm long; anal ring
situated on margin of dorsum. Cauda not seen.
Etymology
The new species is named after the locality of collection the Van Province (Turkey).
Ecology
Biology: Unknown.
Genus:
Kotejacoccus Kaydan & Kozár, 2008
Type sp.: Kotejacoccus turcicus Kaydan & Kozár, 2008: 21.
Lit.: Hodgson & Trencheva 2008: 36; Kozár, 2009: 103 (Miller et al., 2013).
Description
Antennae 6 or 7 segmented; frontal lobe or tubercle present. Venter often with groups
of bilocular micropores or tubular ducts. Legs normal; tibia longer than tarsus. Claw
with a denticle. Cruciform pores absent. Multilocular pores, each with 5-11 loculi, sparse
throughout most of venter, but numerous around spiracles. Setae all hair-like, except
some setae spinose on margin. Dorsum, with strong, capitate, (drum stick-shaped, sensu
Henderson (2007a,b) setae, mostly as long as those near margin. Anal lobes not well
Acanthococcidae 325
Figure 117. Kaweckia vanensis Kaydan sp. n., female.

developed, dorsal surface of each lobe with three-four capitate dorsal setae. Anal ring
sclerotized, not well developed, with six-eight short, strong setae and a few pores. Cauda
usually absent. Macrotubular ducts short, broad. Microtubular ducts narrow, long, few
(Kaydan & Kozár, 2008)
Distribution:
Palearctic, with 3 species known in Euro-Siberian and Irano-Turanian subregions.
Biology
Key to species
1. – Anal ring reduced, venter with bilocular pores in groups...............................K. turcicus

– Anal ring well developed, venter without bilocular pores............................................2
2. – Ventral setae slightly blunted, frontal lobe absent.......................................K. korotyaevi
– Ventral setae hair like, frontal lobe present.....................................................K. orbiculus
Kotejacoccus korotyaevi (Danzig, 1982) (Fig. 118) Combination nova

Acanthococcus korotyaevi Danzig, 1982: 145.
Holotype: Female. Mongolia (Shin-Djinsta, Bayan-Chongorskii), on Reaumuria

soongarica, 21.viii.1981, by Korotyaev. Deposited in ZMAS.
Lit.: Miller & Gimpel, 2000: 248 (Miller et al., 2013).
Eriococcus korotyaevi; Miller & Gimpel, 1999: 214. Change of combination.

Acanthococcus korotyaevi; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Unknown.
Mounted female
Body elongate oval 4.0 mm long. Frontal tubercle present. Anal lobe as long as wide, with
three capitated spines, and short apical setae.
Venter: Labium 3 segmented, basal segment well developed with two steae of different
sizes; apical segment with 5 pairs of setae, median is about the same size. Antennae 7
segmented, antennal segments with short hair-like setae. Legs narow; posterior coxae
with spinulae on anterior surface, and with pores; claw with a denticle, claw digitules
Acanthococcidae 327
Figure 118. Kotejacoccus korotyaevi (Danzig, 1982), female. After Danzig (1982a) with
modifications.
shorter than claw. Legs with a few hair-like setae, tibia each with 4 setae, tarsus with 5
setae. Multilocular pores with 5-10 loculi, distributed in sparse bands and rows on last
abdominal segments, plus a small groups around each spiracle, each group with 4 pores.
Hair-like setae numerous on venter, long strong, some capitated. Macrotubular ducts of
one size, scattered, microtubular ducts absent. Bilocular-micropores and cruciform pores
absent. Suranal setae hair-like.
Dorsum: Dorsal setae all stout and capitate, those on the dorsum being shorter than
those on anal lobe. Marginal row of setae absent. Macrotubular and microtubular ducts
short, wide, scattered and sparse throughout; terminal gland of each duct not sclerotized,
tube-like. Anal ring oval, with 6 fairly short setae. Cauda absent (after Danzig (1982), with
modifications based on holotype).
Ecology
Host plant: Reaumuria soongarica.
Biology: Unknown.
Kotejacoccus orbiculus (Matesova, 1960) (Fig. 119) .

Acanthococcus orbiculus Matesova, 1960: 205.
Holotype: Female. Kazahstan (Ili River, Chulaktau, 160 km. from Iliysk), on Tamarix
ramosissima, 05.ix.1953, by Mitjaev, by original designation. Deposited in AAKA.
Notes: 6 female paratypes, 3 immatures on 5 slides in ZMAS.
Common name: Gall forming felt scale.
Lit.: Bazarov, 1968a: 73; Beardsley, 1984: 86; Borchsenius, 1963: 208; Hoy, 1963: 105;
Köhler, 1998: 380; Kozár, 2009: 93; Kozár & Walter, 1985: 74; Matesova, 1960: 205;
Tang & Hao, 1995: 451 (Miller et al., 2013).
Eriococcus orbiculus; Hoy, 1963: 105. Change of combination.

Acanthococcus orbiculus; Danzig, 1980: 62. Revived combination.
Eriococcus orbiculus; Miller & Gimpel, 2000: 288. Revived combination.
Acanthococcus orbiculus; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult females green coloured.
Mounted female
Body elongate oval 2.3 mm long, 1.2 mm wide. Frontal lobe present. Eyes situated on
venter near margin.
Acanthococcidae 329
Figure 119. Kotejacoccus orbiculus (Matesova, 1960), female. After Matesova (1960) with
modifications.
Venter: Labium well developed, three segmented, basal segment with two pair of
setae one of them about three times longer, apical with five pairs of setae. Antennae 7
segmented, length of each segment: I: 33.6, II: 46.2, III: 63, IV: 65, V: 32.5, VI: 33.6; and
VII: 38 µm long; segment III almost parallel sided; apical segment with 3 falcate sensory
setae, two penultimate segments with falcate sensory setae, each 16 µm long; all antennal
segments with sparse rings of hair-like setae. Legs large; coxae wide, posterior coxae with
spinulae on anterior surface, with numerous small pores; metathoracic legs: femur 243
µm, tibia 210 µm, tarsus with claw 227 µm, tarsal digitules longer than claw, each slightly
knobbed; claw digitules shorter than claw, hair-like, all claw with a denticle. Legs with
a few long hair-like setae, tibia each with 4 setae, tarsus with 5 setae. Quinquelocular
pores distributed in bands on all abdominal segments, and with groups around anterior
spiracles, each group with more than 10 pores. Hair-like setae numerous, in midventer,
and forming a wide submarginal band, about 37–49 µm long. Macrotubular ducts of two
sizes, scattered on venter, 28 µm long and 7 µm wide, microtubular ducts scattered on
margin. Cruciform pores absent. Suranal setae hair-like.
Dorsum: Dorsal setae of two kinds, strong capitate, 18–32 µm long, and longer blunted
hair-like 14–49 µm long. Marginal row of spines absent. Macrotubular ducts short, wide,
each 25 µm long, 9 µm wide, form bands on all segments. Microtubular ducts scattered
over dorsum. Anal ring oval, with 6–8 setae, 88 µm long; anal ring with about 26 pores
in one row. Anal lobes well developed, each with three-four capitate setae 14–22 µm
long, each apical seta 155 and subapical setae 92 µm long. Cauda present (after Matesova
(1960), with modifications).
ECOLOGY
Host plant: Tamarix gracilis, T. leptostachys, T. ramosissima.
Distribution: Kazakhstan, Mongolia.
Biology: The first instars of the scale appear in the middle of summer. They settle
on internodal, terminal, or green twigs of Tamarix. The feeding first instar is gradually
overgrown by fleshy green folds from which the shoot continues to grow. The folds
close to form a gall. The cavity of the gall is smooth and irregular. In autumn the folds
of the gall open and the female leaves. Galls abandoned by the females turn yellow, dry
up and fall off (Matesova, 1960) This species infests Tamarix species so heavily that ‘the
reduction of the assimilating surface in individual plants reaches 80 percent. The growth
of such plants is conspicuously depressed’ (Matesova, 1960).
Acanthococcidae 331
Kotejacoccus turcicus Kaydan & Kozár, 2008 (Fig. 120)

Kotejacoccus turcicus Kaydan & Kozár, 2008: 21.
Holotype: Female. Turkey: Van-Catak, N: 37°55'005", E: 042°57'791" 1438 in altitude

on Quercus sp., 16.v.2006, by M.B. Kaydan. By monotypy. Paratypes: one female (with
two well-developed 1st-instar nymphs inside), Hakkari-Cukurca Road, N: 37º30’986"
E: 043º43’741", 956 m, on Quercus sp., M. B. Kaydan, 15.ix.2005, (KPCT: 2343); also
a separate slide with 2, 1st-instar nymphs, 2, 2nd- and 1, 3rd-instar female nymphs.
Deposited in KPCT, and two females with the same label in PPI.
Common name: Turkish felt scale.
Lit.: Erkılıç el al., 2011: 16; Hodgson & Trencheva 2008: 36; Kozár, 2009:103; Kozár
& Konczné Benedicty, 2008b: 256 (Miller et al., 2013).
Description
Unmounted female
Adult females lilac; found inside small creamy-coloured ovisacs at the bifurcation of
young branches, in a similar manner to Acanthococcus aceris Signoret.
Mounted female
Body elongate oval, 1.19–2.0 mm long, 0.6–1.34 mm wide. Frontal tubercle present. Eyes
situated on venter near margin.
Venter: Labium 3 segmented, 100–110 µm long; basal segment not well developed, but
with two equal long setae on each side; stylet loop much longer than body. Antennae 6
segmented, length of segments: I: 30–35, II: 30, III: 58–62, IV: 20, V: 20, and VI: 38–40
µm long; segment III almost parallel sided; apical seta 70–74 µm long; apical segment
with three falcate sensory setae, longest 32–38 µm long; two penultimate segments with
falcate sensory setae, each 30–32 µm long; all antennal segments with sparse rings of
hair-like setae. Legs normal; lengths of segments and digitules of prothoracic legs; coxa:
40–40, trochanter: 32–40, femur: 78–80, tibia: 65–72; tarsus: 52–55, claw: 21–22, claw
digitules 20–22 µm long; lengths of segments and digitules of mesothoracic legs; coxa:
40, trochanter: 35, femur: 90, tibia: 68, tarsus: 62, claw: 21–22, claw digitules 22 µm long;
lengths of segments and digitules of metathoracic legs; coxa: 40–42, trochanter: 27–30,
femur: 97, tibia: 70–75, tarsus: 55–60; tarsal digitules knobbed, claw 22–25 µm long, claw
digitules each slightly knobbed; all coxae with spinulae on anterior surface, but without
pores; claw with a denticle. Legs with a few hair-like setae, plus a sensory pore on tarsus;
tibia each with 3 setae, tarsus with 4 setae. Quinquelocular pores, each 4.1–5.0 µm in
diameter, distributed in sparse rows on all abdominal and thoracic segments, plus groups
around each spiracle, each group with more than 20 pores. Diameter of anterior spiracles
48–42 µm. Hair-like setae few, scattered on venter, longest setae near vulva where each
about 25–30 µm long. Macrotubular and microtubular ducts absent. Bilocular-micropores
in more than 15 large groups on venter plus a group between antennae. Cruciform pores
absent. Suranal setae setose.
Dorsum: Dorsal setae all stout and capitate, those on the dorsum being 10–18 µm long
and those on the margin 12–26 µm long. Macrotubular ducts short, wide, each 16.4–17.2
µm long, 8.2 µm wide, scattered and sparse throughout; terminal gland of each duct
heavily sclerotized and globose. Microtubular ducts each 4.0–4.1 µm wide and 7.4–9.4
µm long, few, scattered over dorsum. Anal ring oval, 38–42 µm in diameter, with 6 fairly
short, strong setae, each 34–42 µm long; anal ring with 14 pores in one row. Anal lobes
weakly developed, each with four capitate setae, longest 22–29 µm long; each apical seta
strong but short, each 82–120 µm long. Cauda absent.
Ecology
Host plant: Quercus sp.
Acanthococcidae 333
Figure 120. Kotejacoccus turcicus Kaydan & Kozár, 2008, female. After Kaydan & Kozár
(2008).
Genus:
Neoacanthococcus Borchsenius, 1948
Type sp.: Neoacanthococcus tamaricicola Borchsenius, 1948.
Lit.: Bazarov, 1962: 53; Borchsenius, 1949: 364; Ferris, 1957b: 87; Hoy, 1962: 201;
Hoy, 1963: 170; Köhler, 1998: 392; Kaydan & Kozár, 2010: 165; Koteja & Zak-Ogaza,
1981: 502; Kozár, 2009: 104; Kozár & Walter, 1985: 75; Miller & Gimpel, 2000: 402;
Morrison & Morrison, 1966: 129 (Miller et al., 2013).
Greenoripersia Bodenheimer, 1929. Synonym nova

Type sp.: Greenoripersia kaiseri Bodenheimer, 1929: 112. Notes: Hardy et al. (2008: 62)
transferred this genus and species from the Pseudococcidae to the Eriococcidae.
Lit.: Ben-Dov, 1994: 175; Ben-Dov & Harpaz, 1986: 32; Hardy et al., 2008: 62; Kozár,
2009: 102 (Miller et al., 2013).
Description
Body elongate oval. Antennae 6 or 7 segmented. Frontal lobes present. Anal lobes
medium long, hardly sclerotized. Labium 3 segmented; basal segment well developed
with 2 pairs of equal long setae; apical segment with 6 pairs of equally long, strong setae.
Stylet loop long, reaching to level with area between metathoracic legs. Legs long, with
tibia longer than tarsus; tibia with four setae (median seta absent), meso- and metathoracic
coxae with spinulae, and metathoracic coxa also with small pores on femur; claw of all
legs with denticle. Claw digitules shorter than claw. Claw and tarsal digitules hair-like.
Body setae short and hair-like; also with enlarged setae submarginally in one species.
Multilocular pores each with 3 to 5 loculi, present throughout the body. Macrotubular
ducts of one size, heavily sclerotized, some with inner gland ductule ending in a simple
sclerotized hole; terminal gland simple more abundant on dorsum than venter of where
most abundant submarginally but also scattered throughout and scattered over derm.
Microtubular ducts with oval sclerotized orifice, few on margin on venter but numerous
on dorsum. Cruciform pores absent. Anal lobes well developed dorsal surface of each
lobe with 3 spine-like setae, ventral surface of each lobe with a long apical seta and
one shorter subapical seta. Anal ring sclerotised well developed, with anal ring pores in
one row and with 8-20 setae, seta usually shorter than diameter of ring. Enlarged setae,
nearly hair-like, blunt, or sharp pointed at apex and curved slightly mainly distributed
in rows across segments and form a marginal band. Quinquelocular pores scattered on
abdominal segments (generally in rows), thorax and head on both sides. Cauda present.
Distribution:
Palearctic, with 6 species, known in Mediterranean and Irano-Turanian subregions.
Acanthococcidae 335
Biology:
Feed on Tamarix spp. and Centaurea sp.
Key to species
Notes: The species N. kaiseri is not included into the key, because of the inadequate
description and drawing.
1. – Quinquelocular pores on dorsum present.......................................................................2

– Quinquelocular pores on dorsum absent........................................................................4
2. – Marginal and dorsal enlarged setae short and conical................................. N. centaurea
– Marginal and dorsal enlarged setae long, elongated, not conical.................................3
3. – Enlarged setae on dorsum of abdominal segment VII numbering
more than 25; total number of quinquelocular pores on segments
VII-VIII fewer than 15................................................................................ N. tamaricicola
– Enlarged setae on dorsum of abdominal segment VII fewer than 20;
total number of quinquelocular pores on segments VII-VIII numbering
more than 30........................................................................................................N. atlihani
4. – Anal ring with 5 or more pairs of setae.....................................................N. gracilispinus
– Anal ring fewer than 5 pairs of setae.............................................................N. pamiricus
Neoacanthococcus atlihani Kaydan & Kozár, (Fig. 121)

Neoacanthococcus atlihani Kaydan & Kozár, 2010: 171.
Holotype: Female. Turkey, Iğdir - Tuzluca-Digor Road, N: 40°07’ 326”, E: 043°37’720”,

940 m altitude, on Tamarix sp., M.B. Kaydan, 31.viii.2005 (KPCT: 2237). Paratypes:
13 adult females, same data as holotype (KPCT: 2237), 2 adult females, Turkey, Iğdir-
Tuzluca-Aralık Road, N: 39°55’ 24”, E: 044°24’272”, 830 m altitude, on Tamarix sp., M.B.
Kaydan, 01.ix.2005 (KPCT: 2234).
Lit.: Kaydan & Kozár, 2010: 171 (Miller et al., 2013).
Description
Unmounted female
Adult females yellowish red; found at bifurcation of young branches, felt-like test not
seen at collection date.
Mounted female
280–295 µm, length of segments: I: 70–80, II: 40–45, III: 45–55, IV: 55, V: 22.5–35, VI:
20–30, and VII: 37–45 µm long; each segment covered with a few, strong hair-like setae;
apical segment with apical seta 45.0–47.5 µm long; apical segment also with 3 sensory
falcate setae, each 25–35 µm long; segment VI with 1 sensory falcate seta 25.0–27.5 µm
long, segment V with 1 sensory falcate seta 20 µm long. Frontal lobes present. Eyes
situated on venter near margin. Anal lobes developed, each with 3 enlarged setae, each
22.5–40.0 µm plus 2 or 3 microtubular ducts on dorsal surface; apical seta 160–212.5 µm;
Venter: Labium 195–220 µm long, 110–140 µm wide, basal segment well developed,
with two pairs of equal long setae, a pair of median setae on apex of labium 25–35 µm,
about equal long. Legs well developed; lengths of segments and digitules of prothoracic
legs; coxa: 110–125, trochanter: 70–80, femur: 170–195, tibia: 130–160, tarsus: 140–155
and claw: 40.0–47.5, trochanther + femur: 250–270, tibia + tarsus: 270–300, tarsal
digitules: 52.5–62.5, claw digitules 17.5–25.0 µm long; lengths of segments and digitules
of mesothoracic legs; coxa: 120–130, trochanter: 75–90, femur: 180–190, tibia: 160–190,
tarsus: 140–170 and claw: 45–50, trochanther: + femur: 260–280, tibia + tarsus: 310–350,
tarsal digitules 50–60, claw digitules 17.5–25.0 µm long; lengths of segments and digitules
of metathoracic legs; coxa: 140–170, trochanter: 70–100, femur: 180–205, tibia: 180–190,
tarsus: 160–185 and claw: 45–50, trochanther + femur: 270–290, tibia + tarsus: 350–390,
tarsal digitules 55–60, claw digitules 17.5–22.5 µm long. Meso- and metathoracic coxae
each with spinulae on ventral surface and translucent pores on metathoracic coxa and
femur. Tibiae each with 4 setae (median seta absent), tarsi each with 5 setae. Length
of spiracles 80–100 µm; diameter of spiracular peritreme 40–45 µm, posterior spiracles
slightly larger than anterior. Derm with a sparse covering of scattered flagellate hair-like
Acanthococcidae 337
Figure 121. Neoacanthococcus atlihani Kaydan & Kozár, 2010, female. After Kaydan &
Kozár (2010).
setae, each 17.5–137.5 µm long. Enlarged setae, situated on submargin in 2 or 3 rows,

each 17.5–25.0 µm long. Multilocular pores each 6.0–7.5 µm in diameter and with 3-5
loculi, distributed in sparse bands on all abdominal and thoracic segments and head.
Macrotubular ducts of one size, each 6–9 µm wide and 12.5–20.0 µm long, present on
submargin of posterior abdominal segments, and scattered elsewhere on abdomen,
thorax and head. Microtubular ducts sclerotized, with oval orifice, present on margin but
scarce, 6.0–7.0 µm long; cruciform pores absent.
Dorsum: Marginal setae spine-like, blunted, each 27.5–37.5 µm long, forming a marginal
band and not strongly differentiated from dorsal setae; other dorsal setae conical, spine-
like, each setae 17.5–27.7 µm long, those on thorax, head and anterior abdominal segments
slightly larger than setae on posterior 2 or 3 abdominal segments; arranged in transverse
rows across each body segment, rows irregular on head. Multilocular pores each 7.0–
7.5 µm in diameter and with 5 loculi, distributed in sparse bands on all abdominal and
thoracic segments and on head. Macrotubular ducts, each 8–10 µm wide and 20–25 µm
long, scattered throughout dorsum, generally in segmental bands. Microtubular ducts,
6.0–7.5 µm long with oval dermal orifice, scattered over dorsum. Anal ring strongly
sclerotized, with 17-27 pores on each inner side, 90 µm in diameter, with 8 setae, each
85–125 µm long; anal ring situated on margin of dorsum. Cauda present, not well seen
(after Kaydan & Kozár (2010)).
Ecology
Host plant: Tamarix sp.
Neoacanthococcus centaurea (Savescu, 1985) (Fig. 122) Combination

nova
Acanthococcus centaureae Savescu, 1985: 122.
Syntypes: Female. Romania (District de Constantza, Basarabi), on Centaurea solstitialis,

1955. Deposited in Academie des Sciences Agricoles et Forestieres, Bucarest,
Romania. Notes: The slide was searched kindly by Dr. Sonica Drosu and Dr. Liliana
Vasiliu-Oromulu in the Plant Protection Institute and in Biological Institute. We
thanks their great efforts. It was not found, only some other species were present.
The basic part of the collection probably is present in the hands of the wife of
Professor A. Savescu. We hope it will appear sometime in some national collection,
as well as we can hope that his manuscript of a book on Coccoidea fauna of Romania
will be published.
Lit.: Fetykó, et al., 2010: 295; Miller & Gimpel, 1999: 213; 2000: 159-160 (Miller et
al., 2013).
Acanthococcidae 339
Figure 122. Neoacanthococcus centaurea (Savescu, 1985), female. After Savescu (1985)
with modifications.
Eriococcus centaureae; Miller & Gimpel, 1999: 213. Change of combination.

Acanthococcus? centaureae; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female oval, 2.4–2.8 mm long, 1.1–1.3 mm wide and yellowish pink in color. Ovisac
is white and eggs are oval and yellowish orange.
Mounted female
Antennae short wide, 7 segmented, 390–410 µm long, I: 33, II: 63, III: 90, IV: 84, V: 33,
VI: 33 and VII: 56 µm long.
Venter: Legs well developed; lengths of segments of prothoracic legs: femur 257, tibia
230, tarsus 208 µm long, coxae with small pores. Claw with denticle, claw and tarsal
digitules clavate. Quinqelocular pores scattered.
Dorsum: Anal lobes, each with four dorsal setae, anal lobe setae 252 µm long, anal ring
with partly two rows of circular pores, anal ring setae 230 µm long. Quinquelocular
pores scattered on whole dorsum. Macrotubular ducts scattered. All dorsum covered
with enlarged conical setae, forming a submarginal band (after Savescu (1985).
Ecology
Host plant: Centaurea solstitialis.
Distribution: Romania.
Biology: This species reproduces parthenogenetically.
Comments:
The original description is fragmentary, the drawing was printed dark, and authors of this
work could not study the slides of the type material. If it would be possible to study on
the type material it can validate our decision.
Acanthococcidae 341
Neoacanthococcus gracilispinus (Borchsenius & Matesova, 1955) (Fig.

123) Redescription, Combination nova
Acanthococcus gracilispinus Borchsenius & Matesova, 1955: 227.
Syntype: Female. Kazakhstan (Near the river Ili), on Tamarix sp., 23.viii.1951, by G.
Matesova. Deposited in ZMAS.
Lit.: Bazarov, 1968a: 73; Borchsenius, 1963: 211; Borchsenius & Matesova, 1955: 227;
Danzig, 1982: 147; Hoy, 1963: 92; Köhler, 1998: 377; Matesova, 1955: 202; 1960: 209;
1971: 30; Ossianilson, 1955: 4-5; Tang & Hao, 1995: 448, 644 (Miller et al., 2013).
Eriococcus gracilispinus; Hoy, 1963: 92. Change of combination.

Acanthococcus gracilispinus; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus gracilispinus; Miller & Gimpel, 2000: 219; Revived combination.
Acanthococcus gracilispinus; Kozár, 2009: 92. Revived combination.
Proteriococcus gracilispinus; Kaydan & Kozár 2010: 174. Change of combination.
Description
Unmounted female
Adult female is oval, dark claret. At the time of oviposition the female is enclosed in a
tough white sac, 5 mm long, 2.5 mm wide.
Mounted female
320–400 µm, length of segments: I: 70–105, II: 55–80, III: 70.0–82.5, IV: 50–65, V:
27.5–37.5, VI: 25–35, and VII: 35–45 µm long; each segment covered with a few, strong
hair-like setae; apical segment with apical seta 40.0–47.5 µm long; apical segment also
with 3 sensory falcate setae, each 22.5–37.5 µm long; segment VI with 1 sensory falcate
seta 30 µm long, segment V with 1 sensory falcate seta 20 µm long. Frontal lobe present.
Eyes situated on venter near margin. Anal lobes strongly developed, each with 3 enlarged
setae, each 32.5–85.0 µm plus 2 or 3 microtubular ducts on dorsal surface; apical seta
170.0–172.5 µm; ventral hair-like subapical seta 85 µm long.
34–40 µm long. Legs well developed; lengths of segments and digitules of prothoracic
legs; coxa: 135–150, trochanter: 97.5–115.0, femur: 240–245, tibia: 185.0–202.5, tarsus:
145–170 and claw: 55.0–57.5, trochanther + femur: 340–390, tibia + tarsus: 360–390,
tarsal digitules 77.5–80.0, claw digitules 27.5–30 µm long; lengths of segments and
digitules of mesothoracic legs; 127.5–170.0, trochanter: 115–120, femur: 242.5–265.0,
tibia: 210–220, tarsus: 185–195 and claw: 62.5–75.0, trochanther + femur: 350–410, tibia
+ tarsus: 400–460, tarsal digitules 70–85, claw digitules 30 µm long; lengths of segments
and digitules of metathoracic legs; coxa: 140–190, trochanter: 125.0–137.5, femur: 240–
275, tibia: 230–245, tarsus: 205–210 and claw: 55–65, trochanther + femur: 370–440,
tibia + tarsus: 430–470, tarsal digitules 75.0–87.5, claw digitules 30–35 µm long. Meso-
and metathoracic coxae each with spinulae on ventral surface and translucent pores on
metathoracic coxa. Tibiae each with 4 setae (median seta absent), tarsi each with 5 setae.
Length of spiracles 95.0–112.5 µm; diameter of spiracular peritreme 50–60 µm, posterior
spiracles slightly larger than anterior. Derm with a sparse covering of scattered flagellate
hair-like setae, each 47.5–110.0 µm long. Multilocular pores each 5.0–7.5 µm in diameter
and with 3-5 loculi, distributed in sparse bands on all abdominal and thoracic segments
and head. Macrotubular ducts of two sizes, larger one each 7.5–10.0 µm wide and 25–35
µm long, present on submargin of posterior abdominal segments and scattered elsewhere
on abdomen, thorax and head; smaller one each 5–6 µm wide and 17.5–25 µm long,
present on submargin of posterior abdominal segments. Microtubular ducts sclerotized,
with oval orifice, present on margin but scarce, 6.0–7.5 µm long; cruciform pores absent.
Dorsum: Dorsal marginal enlarged setae spine-like, long, each 27.5–42.5 µm, not forming
a marginal row and not strongly differentiated from dorsal setae; other dorsal setae conical,
spine-like, each setae 25–35 µm long, those on thorax, head and anterior abdominal
segments slightly larger than setae on posterior 2 or 3 abdominal segments; arranged in
transverse rows across each body segment, rows irregular on head. Multilocular pores
absent. Macrotubular ducts, each 7.5 µm wide and 25–30 µm long, scattered throughout
dorsum, generally in segmental bands. Microtubular ducts, each 6.0–7.5 µm long with
oval dermal orifice, scattered over dorsum. Anal ring strongly sclerotized, 90.0–92.5 µm
in diameter, with 6-10 setae, each 120–155 µm long; anal ring situated on margin of
dorsum. Cauda present. (Redescription based on type material).
Ecology
Biology: The egg sac is evidently secreted shortly before oviposition, before which the
females go from the branches of the host to the root crown where they are found under
the shelter of a thin layer of earth (Borchsenius & Matesova, 1955; Miller et al., 2013).
Comments
The new drawing and redescription is based on syntype material of collection Matesova
(AAKA), one of them with three females on one slide marked as “paratype” (what was
not mentioned in the original description), two other slides contain two females.
Acanthococcidae 343
Figure 123. Neoacanthococcus gracilispinus (Borchsenius & Matesova, 1955), female.

Original.
Neoacanthococcus kaiseri (Bodenheimer, 1929) Combination nova

Greenoripersia kaiseri Bodenheimer, 1929: 112.
Syntype: Female. Egypt (Sinai Peninsula), on Tamarix mannifera. Probably lost; Ben-
Dov & Harpaz (1986).
Lit.: Ben-Dov, 1994: 175; Ben-Dov & Harpaz, 1986: 32; Hardy et al., 2008: 62; Kozár,
2009: 102 (Miller et al., 2013).
Greenoripersia kaiseri; Bodenheimer, 1929: 112. Status nova. Notes: Kozár & Walter (1985:
69) mentioned that this genus and species may belong to Eriococcidae sensu lato. Hardy
et al., (2008) and Kozár (2009) independently from each other formally transferred this
species from the Pseudococcidae to the Eriococcidae. Kozár (2009) also proposed a
possible relation with Neoacanthococcus.
Description
Unmounted female
Adult female large 4–5 mm long, 2.5 mm wide elongate oval, yellow-brownish, with
white eggsacs.
Mounted female
Antennae short wide, 6 segmented, I: 36, II: 30–36, III: 42–57, IV: 18–21, V: 21, VI: 39
µm long.
Venter: Stylet loop long reaching the third coxae. Legs well developed; lengths of
segments of prothoracic legs; femur 280, tibia 160, tarsus 135; median legs: femur 280,
tibia 165, tarsus 260, lengths of segments of metathoracic legs: femur 300, tibia 180,
tarsus 160 µm long. Posterior spiracles bigger than anteriors.
Dorsum: Anal lobes, each with three dorsal setae, anal ring with one row of circular
pores, anal ring setae 60–100 µm long. Macrotubular ducts with sclerotic vestibules. Hair-
like setae on head shorter, on abdomen 180–200 µm long. All dorsum covered with
enlarged setae with hair-like apices (after Bodenheimer (1929).
Ecology
Host plant: Tamarix mannifera.
Distribution: Egypt.
Biology: On the branches.
Comments
New collection is needed to clarify the position of this species.
Acanthococcidae 345
Neoacanthococcus pamiricus (Bazarov, 1968) (Fig. 124) Combination

nova
Acanthococcus pamiricus Bazarov, 1968a: 73.
Holotype: Female. Tadjikistan (Vanch Valley, 20 km. from Vanch), on Tamarix sp.,
05.vii.1965, by B. Bazarov, by original designation. Deposited in ZMAS. Notes: 1
paratype in TASZ.
Lit.: Köhler, 1998: 382; Kozár & Walter, 1985: 74; Miller & Gimpel, 2000: 29 (Miller
et al., 2013).
Eriococcus pamiricus; Tang & Hao, 1995: 485. Change of combination.

Acanthococcus pamiricus; Kozár, 2009: 93. Revived combination.
Proteriococcus pamiricus; Kaydan & Kozár, 2010: 165. Change of combination.
Description
Unmounted female
Adult females light-green; covered by wax layer, egglaying female enclosen in white, oval
eggsacs, 3.8 mm long, 1.9 mm wide.
Mounted female
Body elongate oval, 2.4 mm long, 1.2 mm wide. Antennae 7 segmented, 280–295 µm,
length of segments: I: 55, II: 24, III: 45, IV: 34, V: 22, VI: 22, and VII: 42 µm long; each
segment covered with a few, hair-like setae; apical segment also with 3 sensory falcate
setae, segment V and VI with 1 sensory falcate seta. Frontal lobe present. Eyes situated
on venter near margin. Anal lobes developed, each with 3 enlarged setae, each 24 µm plus
2 or 3 microtubular ducts on dorsal surface; apical seta 90 µm; ventral hair-like subapical
seta 24 µm long.
Venter: Labium 3 segmented, large, basal segment with two pairs of different size of
setae, median setae on apex of labium strong, short. Legs normal; lengths of segments
of metathoracic legs: femur 198, tibia 175, tarsus and claw 198 µm long, tarsal and claw
digitules pointed, shorter than claw. Claw with denticle. Meso- and metathoracic coxae
each with spinulae on ventral surface and the posterior with translucent pores on coxa.
Tibiae each with 4 setae (median seta absent), tarsi each with 5 setae. Derm with a sparse
covering of scattered flagellate hair-like setae, each 17.5–137.5 µm long. Enlarged setae,
situated on submargin in 2 or 3 rows, each 17.5–25.0 µm long. Multilocular pores each
6.0–7.5 µm in diameter and with 3–5 loculi, distributed in sparse bands on all abdominal
and thoracic segments and head. Macrotubular ducts of two sizes, each 17 µm wide
and 28 µm long, present on submargin of posterior abdominal segments, and scattered
elsewhere on abdomen, thorax and head. Microtubular ducts sclerotized, short, with oval
orifice, and with 2 sclerotized areas, present on margin but scarce, 2 µm wide and 16 µm
long. Cruciform pores absent.
Dorsum: Marginal dorsal setae hair-like or slightly enlarged, apices acute or slightly
rounded, long, each 18–35 µm, forming a marginal band and not strongly differentiated
from dorsal setae; other dorsal setae conical, spine-like, could be 45 µm long, those on
thorax, head and anterior abdominal segments slightly larger than setae on posterior 2
or 3 abdominal segments; arranged in transverse rows across each body segment, rows
irregular on head. Multilocular pores absent. Macrotubular ducts, scattered throughout
dorsum, generally in a marginal and segmental bands. Microtubular ducts scattered over
dorsum. Anal ring oval, medium sized, with partially two rows of pores, with 6 setae,
each 88 µm long; anal ring situated on margin of dorsum. Cauda present, not well seen
(after Bazarov (1968a), with modifications based on the type material).
Ecology
Distribution: Iran, Tajikistan.
Acanthococcidae 347
Figure 124. Neoacanthococcus pamiricus (Bazarov, 1968), female. Original.

Neoacanthococcus tamaricicola Borchsenius, 1948 (Fig. 125)

Neoacanthococcus tamaricicola Borchsenius, 1948: 502.
Lectotype: Female. Turkmenistan (Kara-Kala, Turkmenian Quarantine Inspection),

on Tamarix sp., 08.x.1936, by subsequent designation by Danzig (1996: 522).
Deposited in ZMAS.
Lit.: Borchsenius, 1949: 364; 1963: 211; Bustshik, 1960: 169; Danzig, 1982: 147; 1996:
522; Ferris, 1957b: 87; Hoy, 1962: 23; 1963: 170; Kaydan & Kozár, 2008: 35; Kozár,
2009: 104; Kozár & Walter, 1985: 75; Köhler, 1998: 392; Lashin, 1956: 115; Miller &
Gimpel, 2000: 402; Morrison & Morrison, 1966: 129; Myartseva, 1978: 114; Tang &
Hao, 1995: 512, 513; Ter-Grigorian, 1962: 132, 152, 155; 1969a: 87; 1983: 881 (Miller
et al., 2013).
Description
Unmounted female
Adult female body oval, 2.5 mm long.
Mounted female
Body elongate oval, 2.6 mm long, 1.4 mm wide. Antennae 7 segmented, length of
segments: I: 42, II: 39, III: 46, IV: 43, V: 20, VI: 20, and VII: 38 µm long; each segment
with a few, strong hair-like setae; apical segment with apical seta 40 µm long and with 3
sensory falcate setae, each about 26 µm long; segment VI with 1 sensory falcate seta 28
µm long; segment V with 1 sensory falcate seta 18 µm long. Frontal lobes and sclerotized
frontal tubercles present. Eyes situated on venter near margin. Anal lobes strongly
developed, each with 3 enlarged setae plus with 2 microtubular ducts on dorsal surface;
apical seta 150 µm; ventral subapical hair-like seta 60 µm long.
Venter: Labium 161 µm long, basal segment with two pairs of almost equal long setae,
apical segments with 6 pairs of setae, median one the same size as others. Legs well
developed: lengths of segments and digitules of prothoracic legs; coxa: 91, trochanter:
82, femur: 173, tibia: 149, tarsus: 134 and claw: 46; tarsal digitules: 50, claw digitules:
20 µm long; lengths of segments and digitules of mesothoracic legs; coxa: 103, with
spinulae, trochanter: 77, femur: 185, tibia: 173, tarsus: 144 and claw: 48, tarsal digitules:
50, claw digitules: 20 µm long; lengths of segments and digitules of metathoracic legs;
coxa: 108, trochanter: 70–100, femur: 180, tibia: 175, tarsus: 151 and claw: 48, tarsal
digitules: 53, claw: digitules 22 µm long. All coxae with spinulae and metathoracic coxae
and femur with translucent pores on ventral surface. Tibia each with 4 setae, tarsi each
with 5 setae. Multilocular pores each 5–6 µm in diameter and with 3–5 loculi, distributed in
sparse rows on all abdominal and thoracic segments and also head. Diameter of anterior
spiracular peritreme 43 µm. Derm with a sparse covering of scattered flagellate hair-like
setae. Enlarged setae situated on submargin in 2 or 3 rows. Macrotubular ducts of one
size, each 5–7 µm wide and 20–22 µm long, distributed on submargin of abdominal
Acanthococcidae 349
Figure 125. Neoacanthococcus tamaricicola Borchsenius, 1948, female. After Kaydan &
Kozár (2010).
segments, and scattered on thorax and head. Microtubular ducts present marginally but
few, 5–6 µm long. Cruciform pores absent.
Dorsum: Dorsal enlarged setae nearly hair-like, long, each 18–31 µm; setae on thorax,
head and anterior abdominal segments slightly larger than setae on posterior 2 or 3
abdominal segments; arranged in transverse bands across each body segment, rows
irregular on head. Multilocular pores each 5–6 µm in diameter and with 3–5 loculi,
distributed sparsely on all abdominal and thoracic segments and on head. Macrotubular
ducts each 5–7 µm wide and 20–22 µm long, scattered throughout dorsum, generally in
segmental bands. Microtubular ducts, each 6 µm long with oval dermal orifice, scattered
over dorsum. Anal ring strongly sclerotized, 105 µm long, and 75 wide, with 8 setae
according to Borchsenius (1949), but studied paratype with 4 on one side and 5 on other
side, each 62–100 µm long, and with about 40 pores on inner side of ring in total; anal
ring situated on margin of dorsum. Cauda present, not well seen (Redescription is based
on lectotype and paralectotype).
Ecology
Host plant: Polygonum sp., Tamarix sp.
Distribution: Turkmenistan.
Acanthococcidae 351
Genus:
Neokaweckia Tang & Hao, 1995
Type sp.: Greenisca rubra Matesova, 1960: 205; by monotypy and original designation.
Lit.: Danzig, 2006: 203; Kozár, 2009: 104; Miller & Gimpel, 1999: 215; 2000: 320;
Description
Adult female elongate oval, almost parallel-sided; antennae 6 or 7 segmented, base at
body margin; frontal tubercles present; labium 3 segmented generally wide; stylet loop
extends to line between mid-coxae. Legs well developed, hind tibia with 4 setae, 2 on
inner side, coxa generally with translucent pores; claw usually with a denticle at apex; tarsal
and claw digitules longer than claw clavate. Spiracles often surrounded by sclerotised
area, with a few associated disc pores. Discoidal pores generally 3-9 loculars, on both
surfaces. Cruciform pores on dorsum and ventrum; micro- and macrotubular ducts on
both surfaces. Enlarged setae cone-shaped, apices rounded, restricted to posterior 2 or
3 abdominal segments and on anal lobes. Anal lobes with 4 enlarged setae; dorsal setae
small needle-like, ventral setae in medial area enlarged, hair-like; anal ring circular small,
broad.
Distribution
Palearctic, with 2 species known in Euro-Siberian and Irano-Turanian subregions.
Biology
These are rare steppe-inhabiting xerophilous species.
Comments
Miller & Gimpel (1999: 215) synonymyzed Neokaweckia with Eriococcus genus. However
Kaydan & Kozár (2008: 23), Kozár & Konczné Benedicty (2008b: 256), Kozár (2009:
104) accepted the name of Neokaweckia as a valid name. For further studies are needed
to clarify status of this genus.
Key to species
1. – Enlarged marginal setae present on abdominal segments IV-VIII+IX; cruciform

on dorsum in big group on mid thorax..............................................................N. rubra
– Enlarged marginal setae present only on abdominal segments VII-VIII+IX;
cruciform on dorsum in only small numbers on mid thorax................... N. laeticornis
Neokaweckia laeticoris (Tereznikova, 1965) (Fig. 126)

Greenisca laeticoris Tereznikova, 1965: 975.
Holotype: Female. Ukraine (Donezk Oblast, Krasnolimansky District, Zakotnoe),

on Festuca sulcata leaves, 05.vii.1962, by E. Tereznikova, by original designation.
Deposited in ZMAS, paratypes in Institute of Zoology, Ukraine Academy of Science,
Kiev.
Common name: Tereznikova's Felt Scale.
Lit.: Danzig, 1980: 231; 2006: 203; Kosztarab & Kozár, 1988: 295; Koteja, 1974b:
77; 1986: 218-219; Kozár et al., 2013: 56; Kozár & Walter, 1985: 75; Miller & Gimpel,
1996: 601; 1999: 214; 2000: 249; Tang & Hao, 1995: 514; Tao, 1999: 34 (Miller et al.,
2013).
Kaweckia laeticoris; Koteja & Zak-Ogaza, 1981: 507. Change of combination.

Neokaweckia laeticoris; Tang & Hao, 1995: 514. Change of combination.
Acanthococcus laeticoris; Miller & Gimpel, 1996: 601. Change of combination.
Kaweckia laeticoris; Köhler, 1998: 391. Revived combination.
Eriococcus laeticoris; Miller & Gimpel, 1999: 214. Change of combination.
Neokaweckia laeticoris; Kozár, 2009: 104. Revived combination.
Description
Unmounted female
Female elongate oval, reddish, 2.5–4 mm long, 1.0–1.5 mm wide. Ovisacs in leaf sheaths,
cocoon-like, whitish, flattened, thin.
Mounted female
Antennae 6 or 7 segmented, each segment covered with a few. Frontal tubercle present.
Eyes situated on venter near margin; anal lobes short and broad with long apical setae,
each with 4 cone-shaped, apices rounded enlarged setae on dorsal surface.
Venter: Labium 3 segmented, basal segment with two pairs of sort setae, apical segment
with 6 pairs of setae, the median are short, spine-like, stylet loop reaches line between
middle legs. Legs small, tibia with 4 setae, hind coxae with a large group of translucent
pores, claws with small denticle, tarsal and claw digitules capitated, longer than claw.
Spiracles small, with sclerotized oval rim; a large group of disc pores (quinque- or
septaloculars) anterior of each spiracle. Cruciform pores scattered on entire dorsum;
3-5-7-locular pores frequent on body surface. Macrotubular ducts of one size numerous
on body surfaces, form a submarginal band; microtubular ducts scattered along margin
of abdomen. Hair-like setae narrow, short.
Dorsum: Dorsal setae few, hair-like; enlarged setae cone-shaped, apices rounded, restricted
to posterior 2 abdominal segments. Cruciform pores scattered on entire dorsum;
Multilocular pores with 3, 5 and 7 loculi, frequent on body surface. Macrotubular ducts
Acanthococcidae 353
Figure 126. Neokaweckia laeticoris (Tereznikova, 1965), female. After Tereznikova (1965)
with modifications.
of one size numerous on body surfaces. Microtubular ducts scattered along margin of
abdomen. Anal ring circular, with partly double row of pores, and 8 setae. Cauda present
(after Tereznikova (1981), with modifications based on paratype).
Ecology
Host plant: Agrostis sp., A. capillaris, Elymus sp., Festuca sp., F. sulcata, Stipa sp.
Distribution: China (Inner Mongolia), former Czechoslovakia, Hungary, Poland,
Ukraine.
Biology: This is a rare steppe-inhabiting xerophilous species. Often found with Kaweckia
glyceriae. Adult females collected in July and August (Kosztarab & Kozár, 1988).
Neokaweckia rubra (Matesova, 1960) (Fig. 127) Redescription

Greenisca rubra Matesova, 1960: 205.
Syntype: Female. Kazakhstan (Karagandin region, North Valley), on Elymus sp.

Deposited in ZMAS.
Common name: Matesova's felt scale.
Lit.: Borchsenius & Danzig, 1966: 42; Danzig, 1980: 228; 1986: 265, 269; Dziedzicka,
1977: 5; Hoy, 1963: 134. Kosztarab & Kozár, 1988: 297; Koteja, 1974a: 77; 1986:
219; Koteja & Zak-Ogaza, 1981: 508; Kozár, 2009: 104; Kozár & Walter, 1985: 75;
Köhler, 1998: 391; Matesova, 1971: 26; Miller & Gimpel, 1996: 603; 1999: 215; 2000:
321; Tang & Hao, 1995: 514; Tereznikova, 1965: 958; 1975: 8, 20, 51, 64, 74; Williams,
1985a: 356 (Miller et al., 2013).
Kaweckia rubra; Koteja & Zak-Ogaza, 1981: 508. Change of combination.

Neokaweckia rubra; Tang & Hao, 1995: 515. Change of combination.
Acanthococcus rubra; Miller & Gimpel, 1996: 603. Change of combination.
Kaweckia rubra; Köhler, 1998: 392. Revived combination.
Eriococcus rubrus; Miller & Gimpel, 1999: 215. Change of combination.
Neokaweckia rubra; Kozár, 2009: 104. Revived combination.
Description
Unmounted female
Adult female body is elongate-oval, flat, clear rose in color. Ovisac cocoon-like, with pink
eggs. Second instar is flat, elongate-oval and clear rose also.
Mounted female
Body elongate oval, 3.30 mm long, 1.46 mm wide. Antennae 7 segmented, 220 µm, length
of segments; I: 45, II: 35, III: 40, IV: 35, V: 20, VI: 20, and VII: 35 µm long; each segment
covered with a few (except third segment), strong hair-like setae; apical segment with
Acanthococcidae 355
Figure 127. Neokaweckia rubra (Matesova, 1960), female. Original.

apical seta 50 µm long; apical segment also with 3 sensory falcate setae, each 22.5–30.0
µm long; segment VI with 1 sensory falcate seta 30 µm long, segment V with 1 sensory
falcate seta 15 µm long. Frontal tubercle present. Eyes situated on venter near margin.
Anal lobes strongly developed, each with 4 enlarged setae, short, truncated, each 17.5–
22.5 µm plus on dorsal surface; apical seta 280 µm; ventral hair-like subapical seta 100 µm
long, in addition on the ventral side of the anal lobe with one hair-like setae 45 µm long.
Venter: Labium 110 µm long, 105 µm wide, median setae on apex of labium short,
spine-like, 7.5–10.0 µm long. Legs well developed; lengths of segments and digitules of
prothoracic legs; coxa: 60, trochanter: 45, femur: 125–135, tibia: 102.5–105.0, tarsus: 115–
117.5 and claw: 30–35, trochanther + femur: 175.0–177.5, tibia + tarsus: 215.0–217.5,
tarsal digitules 50–55, claw digitules 37.5 µm long; lengths of segments and digitules of
mesothoracic legs; coxa: 60, trochanter: 45, femur: 122.5, tibia: 110.0–112.5, tarsus: 122.5
and claw: 32.5–35.0, trochanther + femur: 170–175, tibia + tarsus: 237.5, tarsal digitules
52.5–57.5, claw digitules 35 µm long; lengths of segments and digitules of metathoracic
legs; coxa: 75–80, trochanter: 50–55, femur: 135, tibia: 112.5–120, tarsus: 127.5–132.5 and
claw: 35, trochanther + femur: 185–190, tibia + tarsus: 245, tarsal digitules 57.5–60, claw
digitules 40.0–42.5 µm long. Metathoracic coxae with translucent pores on both surface
of coxa. Tibiae each with 4 setae (median seta absent), tarsi each with 5 setae. Length
of spiracles 60–70 µm; diameter of spiracular peritreme 35–45 µm, posterior spiracles
setae, each 12.5–80.0 µm long. Multilocular pores each 5–7 µm in diameter and with 5-7
loculi, distributed in sparse bands on all abdominal segments and bands on spiracles on
thoracic segments and head. Macrotubular ducts of one size, each 3–6 µm wide and
15–22 µm long, present on scattered on abdomen, thorax and head. Microtubular ducts,
each 3–5 µm long scattered over submargin of thorax and head. Cruciform pores absent.
Dorsum: Enlarged setae cylindrical, sides straight, apices truncate, enlarged setae restricted
to posterior 2 or 3 abdominal segments, other dorsal setae hair like; each 15.0–22.5 µm.
dorsal multilocular pores present, predominantly with 7 loculi. Macrotubular ducts of
one size, each 3–6 µm wide and 15–22 µm long, present on scattered on abdomen,
thorax and head. Microtubular ducts, each 3–5 µm long scattered over submargin of
thorax and head. Anal ring sclerotized, with one row of pores, 40 µm in diameter, with
8 setae, each 65–70 µm long; anal ring situated on margin of dorsum Cruciform pores
scattered on thorax and first three abdominal segments, each 3–4 µm. Small cauda present
(Redescription is made based on type material).
Other stages
Last nymphal instar described by Matesova (1960).
Ecology
Host plant: Agrostis sp., Elymus sp., Festuca sp., Stipa sp., S. baicalensis.
Distribution: Kazakhstan, Kyrgyzstan, Mongolia, Russia (Primor'ye).
Acanthococcidae 357
Biology: Live in axils of leaves of grasses, infested stems are discolored and die. Nymph
were found at the end of June, but adult females were not yet present.
Comments
Tang & Hao (1995: 731) present a drawing (Fig. 61), as N. rubra, however in almost all
detailes of it identical with N. laetocoris.
Genus:
Neotrichococcus Miller & Gimpel, 1999
Type sp.: Trichococcus filifer Borchsenius, 1948: 503.
Lit.: Bazarov, 1962: 53; Borchsenius, 1949: 322; Ferris, 1957b: 89; Hoy, 1962: 13, 23,
201; 1963: 195; Koteja & Zak-Ogaza, 1981: 502; Kozár, 2009: 104; Kozár & Walter,
1985: 75; Köhler, 1998: 402; Miller & Gimpel, 2000: 404; Morrison & Morrison,
1966: 197; Tang & Hao: 1995: 516 (Miller et al., 2013).
Trichococcus Borchsenius, 1948: 503. by monotypy and original designation. Homonym of

Trichococcus Kanda, 1941; discovered by Miller & Gimpel, 1999, which is now considered
a junior synonym of Beesonia Green.
Nidularia; Lindinger, 1957: 552. Incorrect synonymy; discovered by Hoy, 1963: 195.
Neotrichococcus Miller & Gimpel, 1999: 215. Replacement name for Trichococcus Borchsenius,
1948.
Description
Adult female short, oval, convex, dark crimson in color. Egg sac is white, spherical, and
flocculent and looks like a ball of cotton (Borchsenius, 1949). Antennae 7 segmented.
Frontal lobes or tubercle not found. Anal lobes medium long, hardly sclerotized. Labium
3 segmented; basal segment well developed with 2 pairs of setae; apical segment with 5
pairs of equally long setae. Legs long, tibia with 4 or 5 setae, meso- and metathoracic
coxae with spinulae, and metathoracic coxa also with small pores on femur; claw of all
legs with denticle. Claw digitules shorter or longer than claw. Body setae short and hair-
like; also with enlarged setae submarginally in one species. Enlarged setae cylindrical,
sides concave basally, apices truncate, bases with 3 rings, 2 or 3 sizes of setae, mainly
distributed in double rows across segments and form a marginal band. Multilocular pores
each with 3 to 5 loculi, scattered on abdominal segments (generally in rows), thorax and
head on both sides. Macrotubular ducts of two sizes, heavily sclerotized, some with inner
gland ductule ending in a simple sclerotized hole; more abundant on dorsum than venter
scattered throughout. Microtubular ducts with oval sclerotized orifice, few on margin on
venter but numerous on dorsum. Cruciform pores absent. Anal lobes well developed
dorsal surface of each lobe with 3 spine-like setae, ventral surface of each lobe with a
long apical seta and two shorter subapical seta. Anal ring sclerotised well developed, with
anal ring pores in one row and with 6-8 setae, seta usually shorter than diameter of ring.
Cauda present.
Distribution
Palearctic, with 2 species known in Irano-Turanian subregion.
Biology
Feed on Acantholimon sp. and Salsola spp. On the leaf and shoot.
Key to species
1. – Enlarged setae on posterior segment of abdomen on dorsum in bands, dorsum

without trilocular pores...........................................................................................N. filifer
– Enlarged setae on posterior segment of abdomen on dorsum in rows, dorsum
with trilocular pores...........................................................................................N. notabilis
Neotrichococcus filifer (Borchsenius, 1948) (Fig. 128) Redescription

Trichococcus filifer Borchsenius, 1948: 503.
Lectotype: Female. Tajikistan (Isfara District, Voruch), on Acantholimon sp. stems,

10/09/1944, by N. Borchsenius. By subsequent designation by Danzig, 1996: 521.
Deposited in ZMAS. Notes: 4 female paralectotypes on same slide as lectotype
(Danzig, 1996).
Lit.: Bazarov, 1962: 53; Borchsenius, 1949: 326; 1963: 32; Danzig, 1996: 574; Ferris,
1957b: 89; Hoy, 1962: 13, 23, 201; 1963: 195; Koteja & Zak-Ogaza, 1981: 502; Kozár,
2009: 104; Köhler, 1998: 402; Miller & Gimpel, 2000: 404; Morrison & Morrison,
1966: 197; Tang & Hao, 1995: 516, 517 (Miller et al., 2013).
Nidularia filifer; Lindinger, 1957: 552. Change of combination.

Trichococcus filifer; Kozár & Walter, 1985: 75. Revived combination.
Neotrichococcus filifer; Miller & Gimpel, 1999: 215. Change of combination.
Description
Unmounted female
Adult female short, oval, convex, dark crimson in colour. Egg sac is white, spherical,
Acanthococcidae 359
Figure 128. Neotrichococcus filifer (Borchsenius, 1948), female. Original.

flocculent and looks like a ball of cotton, up to 5 mm in diameter, some needle-like wax
is up to 10 mm (Borchsenius, 1948).
Mounted female
Body elongate oval, 3.2 mm long, 2.0 mm wide. Frontal tubercle not seen. Eyes situated
on venter near margin. Antennae 7 segmented, length of each segment: I: 20–24 µm,
II: 20–22, III: 28–34, IV: 22–28, V: 17–20, VI: 15–19 and VII: 28–31 µm long; segment
III almost parallel sided; apical seta 29–36 µm long; apical segment with three falcate
sensory setae, 22–26 µm long; two penultimate segments with falcate sensory setae, each
11–24 µm long; all antennal segments with sparse rings of hair-like setae. Anal lobes well
developed, each with three strong truncated setae, apical seta 99–101 µm long.
Venter: Labium 3 segmented, 96–101 µm long, basal segment well developed with two
equal long setae, and with equal long five setae on apical segment. Legs well developed;
lengths of segments and digitules of prothoracic legs; coxa: 44–50 µm, trochanter: 35–
38 µm, femur: 96–101 µm, tibia: 60–66 µm; tarsus: 75–81 µm, claw: 22–24 µm, claw
digitules 24 µm; lengths of segments and digitules of mesothoracic legs; coxa: 45–48
µm, trochanter: 30–36 µm, femur: 80–84 µm, tibia: 61–64 µm, tarsus: 80–86 µm; tarsal
digitules knobbed, claw 26–36 µm; lengths of segments and digitules of metathoracic
legs; coxa: 57–60 µm, trochanter: 33–38 µm, femur: 83–91 µm, tibia: 70–75 µm, tarsus:
91–94 µm; tarsal digitules knobbed, claw 26 µm, claw digitules slightly knobbed; all coxae
with spinulae on anterior surface, posterior coxae with numerous small pores; claw with
a denticle. Legs with a few hair-like setae, tibia each with 4 setae, tarsus with 4 setae.
Quinquelocular pores, each 6 µm in diameter, distributed in rows on all abdominal and
thoracic segments. Diameter of anterior spiracles 23–28 µm. Hair-like setae few, scattered
on venter, longest setae near vulva. Macrotubular ducts short, wide, each 24 µm long, 4
µm wide, scattered and sparse throughout; terminal gland of each duct heavily sclerotized
and globose. Microtubular ducts each 5 µm long, few. Cruciform pores absent. Suranal
setae hair-like.
Dorsum: Enlarged setae conical, sides slightly concave, apices slightly rounded, setae
of 2 or 3 sizes, forming rows and bands over surface, 19–34 µm long on the margin.
Macrotubular ducts short, wide, each 26 µm long, 7 µm wide, scattered and sparse
throughout; terminal gland of each duct heavily sclerotized and globose. Microtubular
ducts each 5 µm long, few, scattered over dorsum. Anal ring oval, 74 µm in diameter, with
8 setae, each 108 µm long. Cauda well developed (Redescription is based on the lectotype
and two paralectotype).
Other stages
First instar described by Borchsenius (1949).
Ecology
Host plant: Acantholimon sp.
Acanthococcidae 361
Biology: The egg sac is evidently secreted shortly before oviposition, before which the
females go from the branches of the host to the root crown where they are found under
the shelter of a thin layer of earth (Borchsenius and Matesova (1955); Miller et al., 2013).
Neotrichococcus notabilis (Borchsenius, 1949) (Fig. 129) Redescription,

Combination nova
Acanthococcus notabilis Borchsenius, 1949: 344.
Lectotype: Female. Tajikistan near Parkhar, on Salsola sp. stems, 11/10/1944, N.

Borchsenius. Subsequent designation by Danzig, 1996: 521. Notes: There are 2 more
female paralectotypes on two slides. Deposited in ZMAS.
Lit.: Danzig1975: 69; 1996: 574; Hoy, 1963: 104; Köhler, 1998: 381; Miller & Gimpel,
2000: 281 (Miller et al., 2013).
Nidularia notabilis; Lindinger, 1957: 543. Change of combination.

Eriococcus notabilis; Hoy, 1963: 104. Change of combination.
Acanthococcus notabilis; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus notabilis; Tang & Hao, 1995: 452, 649; Revived combination.
Acanthococcus notabilis; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female short, oval, convex, dark crimson in colour. Egg sac is white, oval and
compact, entirely encloses the body (Borchsenius, 1949).
Mounted female
Body elongate oval, 2.0 mm long, 1.0 mm wide. Frontal tubercle not seen. Eyes situated
on venter near margin. Antennae 7 segmented, length of each segment; I: 46, II: 36, III:
48, IV: 36, V: 24, and VI: 19; VII: 38 µm long; segment III almost parallel sided; apical
seta 48 µm long; apical segment with three falcate sensory setae, longest 29 µm long;
two penultimate segments with falcate sensory setae, each 14–26 µm long; all antennal
segments with sparse rings of hair-like setae. Anal lobes well developed, each with three
strong truncated setae, apical seta broken.
Venter: Labium 3 segmented, basal segment well developed with two equal sized setae,
apical segment with six setae, the median is strong, spine-like. Legs well developed;
lengths of segments and digitules of prothoracic legs; coxa: 77, trochanter: 48, femur:
168, tibia: 130; tarsus: 137, claw: 46, claw digitules: 17 µm long; lengths of segments of
mesothoracic legs: coxa: 67 µm long (the rest of legs were lost), lengths of segments and
digitules of metathoracic legs; coxa: 84, trochanter: 55, femur: 168, tibia: 144, tarsus: 154;
tarsal digitules knobbed, claw: 55 µm long, claw digitules each slightly knobbed; all coxae
with spinulae on anterior surface, but without pores; claw with a denticle. Legs with a few
hair-like setae, plus a sensory pore on tarsus; tibia each with 3 setae, tarsus with 4 setae.
Quinquelocular pores, each 5–6 µm in diameter, distributed in rows on all abdominal and
thoracic segments. Diameter of anterior spiracles 31 µm. Hair-like setae few, scattered on
venter, longest setae near vulva. Macrotubular ducts short, wide, each 20 µm long, 4 µm
wide, scattered and sparse throughout; terminal gland of each duct heavily sclerotized
and globose. Microtubular ducts each 5 µm long, few. Cruciform pores absent. Suranal
setae hair-like.
Dorsum: Enlarged setae conical, sides slightly concave, apices slightly rounded, setae
of 2 or 3 sizes, forming rows and bans over surface, 36 µm long and those on the
margin and 24 µm in other places. Trilocular pores scattered on dorsum 4 µm in diameter.
Macrotubular ducts short, wide, each 26 µm long, 7 µm wide, scattered and sparse
throughout; terminal gland of each duct heavily sclerotized and globose. Microtubular
ducts each 5 µm long, few, scattered over dorsum. Anal ring oval, 74 µm in diameter, with
8 setae, each 108 µm long. Cauda well developed (Redescription is based on the lectotype
and two paralectotype).
Ecology
Host plant: Salsola sp.
Biology: Lives on the stems and leaves of its host plant.
Acanthococcidae 363
Figure 129. Neotrichococcus notabilis (Borchsenius, 1949), female. Original.

Genus:
Noteococcus Hoy, 1962
Type sp.: Eriococcus hoheriae Maskell, 1962, by monotypy and original designation.
Lit.: Henderson, 2007b: 3; Hoy, 1962: 164; Köhler, 1998: 392; Kozár, 2009: 113;
Kozár & Walter, 1985: 75; Miller & Gimpel, 2000: 405; Morrison & Morrison, 1966:
136 (Miller et al., 2013).
Description
Antennae short and relatively stout, 6 segmented. Legs much reduced, each consisting
of coxa, fused femur-trochanter, and fused tibia and tarsus, claw without denticle. Derm
membranous except for markedly nodulose posterior abdominal segments. A lunate-
shapped rugose plate anterior to lobes on dorsum. Anal lobes strongly sclerotised, in
addition to two large setae on margins, bearing about 12 small enlarged setae on dorsum;
irregular series of teeth on venter of lobes. Anal ring with 6 setae. Tubular ducts elongate,
numerous on dorsum. Quinquelocular pores numerous on venter, few on dorsal margin
of thoracic region (Hoy, 1962).
Distribution
Australasian and Palaearctic. Only one species is known from this genus, which was
probably introduced to Europe.
Biology
Occurring in crevices in bark of main trunk of host plant. Female not forming definite
sac but surrounded by white cottony wax.
Comments
Hoy (1962) notes that this genus is closely related to Phloeococcus, but can be distinguished
by its reduced legs and the presence of long tubular ducts.
Acanthococcidae 365
Noteococcus hoheriae (Maskell, 1880) (Fig. 130)

Eriococcus hoheriae Maskell, 1880: 298.
Holotype: Female. New Zealand: South Island, on the hills above Lyttelton, on
Hoheria sp., by W.M. Maskell. Lectotype female, by subsequent designation Deitz &
Tocker, 1980: 47. Type depository: Auckland: New Zealand Arthropod Collection,
Landcare Research, New Zealand. Described: female. Illust. Notes: Type material
also in BMNH, ANIC, USNM.
Lit.: Boratynski & Williams, 1964: 91; Hoy, 1962: 164; 1963: 173; Köhler, 1998: 392;
Kozár, 2009: 113; Kozár & Walter, 1985: 75; Miller & Gimpel, 2000: 405; Morrison
& Morrison, 1966: 136; Williams, 1985a: 384 (Miller et al., 2013).
Nidularia hoheriae; Lindinger, 1933a: 116. Change of combination.

Noteococcus hoheriae; Hoy, 1962: 164. Change of combination.
Description
Unmounted female
Adult female without definite sac surrounded by white cottony wax.
Mounted female
Body pyriform, segmentation conspicuous particularly on abdominal segment. 1.22 long.
Antennae 6 segmented, short. Anal lobes heavily sclerotised, complex, triangulate, very
rugose; caudal setae much shorter than lobes; on masal margin of lobes two large, curved,
apically acute setae, on other margin a series of five short conical, apically acute setae; on
dorsal surface a further series of five similar setae; a pair of sharp teeth on apical masal
angle; on ventral surface of lobes a prominent angled series of teeth arranged in two
groups, in apical group teeth being much larger in size than in basal group; at base of
teeth a sharp tipped setae of medium length; towards outer ventral margin about midway
between base and apex a shourt conical apically acute setae.
Venter: Derm membranous with some fine spicules towards mid-line. Setae on last
abdominal segments of similar length on those on dorsum but more slender, remaining
ventral setae slightly larger than small dorsal setae. Legs very much reduced in size,
apparently consisting of three joints composed of a coxa, fused trochanter and femur
and fused tibia and tarsus, claw very small, without denticle. Quinquelocular pores, small,
with thickened lightly sclerotised rim; these pores numerous in two irregular bands from
lobes to posterior spiracles thence to ventral margin and on cephalic region.
Dorsum: Derm membranous, except on last two abdominal segments, these being
conspicuously nodulose: a rugose, lunate sclerotised plate in front of anal lobes. Dorsal
setae few small, sharp tipped and inconspicuous on antepenultimate abdominal segment
a large setae towards each margin, these setae slightly curved and tapering to sharp
apex, also transverse row of six similar slightly smaller setae; on penultimate abdominal
segment a transverse row of six setae similar to those on antepenultimate segment. Anal
ring apparently with 6 setae. Tubular duct with shallow asymmetrical cup, ducts relatively
long, numerous with some marginal concentration. Microtubular ducts not observed.
Quinquelocular pores a few on dorsal margin (after Hoy (1962).
Ecology
Host plant: Hoheria sp., H. populnea.
Distribution: New Zealand, United Kingdom (Scilly Isles).
Biology: Sac is white and cottony, but is often covered with black fungus causing it to
look like a small gall or an excrescence on the bark. The eggs are very minute, oval and
red. Adult female is red.
Acanthococcidae 367
Figure 130. Noteococcus hoheriae (Maskell, 1880), female. After Hoy (1962) with
modifications.
Genus:
Orontesicoccus Kaydan Genus nova
Type sp.: Proteriococcus lauri Erkılıç, 2011 (in Erkılıç et al., 2011: 17). By monotypy.
Description
Unmounted female
Dark red.
Mounted female
Venter: Antennae 7 segmented; frontal lobe present. Basal segment of labium with two
pairs of setae, on apical segment with five pairs of setae. Multilocular pores, each with
5 loculi. Macrotubular ducts of two sizes, scattered on venter especially on margin. Legs
normal, tibia equal size with tarsus. Claw with denticle. All coxae with spinulae; posterior
coxae also with small pores. Microtubular ducts absent. Cruciform pores absent. Ventral
setae in small number, hair-like.
Dorsum: Marginal dorsal setae, long intermediate type (spine-like with pointed and
curved apex), well separated marginal row of spines, 4 or 5 on each segment. Anal lobes
well developed; dorsal surface of each lobe with three setose spines, spinulae present
on anal lobe, on cauda and on derm, ventral surface of each lobe with a long apical seta
and shorter subapical seta. Anal ring sclerotized, well developed with one row of anal
ring pores and with 6 long setae, all longer than diameter of ring. Cauda present. Large
macrotubular ducts present in high number all over the dorsum, heavily sclerotized, each
with inner ductules ending in a simple sclerotized pore; terminal gland not observed.
Microtubular ducts of two kinds, one is narrow, long spread all over dorsum among
spines. Others are smaller, fused with dorsal spines in number of 1 or 2.
Etymology
The new genus is named after the ancient name of the river Asi “Orontes”.
Distribution
Palaearctic, with species in Mediterranean subregion.
Biology
Occurring in crevices in bark and on both side of leaves of host plants. Female forming
definite sac by white cottony wax.
Acanthococcidae 369
Comments
The genus Orontesicoccus is unique in having the following combination of characters:
(i) on dorsum with setose spines with 1 or 2 associated microtubular ducts, what is a
rare character in acanthococcids of the Palaearctic Region, (ii) apical segment of labium
with 5 pairs of setae, (iii) anal lobes with spinulae and setose spines, (iv) with serrated
surface on cauda and frontal lobes, (v) microtubular ducts long, with elongated structure.
The new genus shows some similarities with Proteriococcus Borchsenius, 1960c, having
setose spines on margin and anal lobes, with Sangicoccus Reyne, and some other genera
and Eriococcus sensu lato species from New Zealand, Australia and Neotropical regions,
by having fused microtubular ducts associated with spines. Orontesicoccus differs from
Proteriococcus in having fused microtubular ducts with base of spines, by the absence of
cruciform pores, and from Sangicoccus by the absence of truncated spines. The monotypic
genus Orontesicoccus is known from Turkey in the Palaearctic Region, where it lives, on
Laurus nobilis leaves, in a place known for many endemic plant species.
Orontesicoccus lauri (Erkılıç, 2011) (Fig. 131) Combination nova

Proteriococcus lauri Erkiliç in Erkiliç et al., 2011: 17.
Holotype: Female. Turkey (Hatay-Harbiye, N:36°07' E:36°08', 239 m altitude), on

leaves of "Laurus nobilis, " 26/v/2008, by L. Erkiliç. By original designation. Notes:
Paratypes: Five adult females and 9 first-instar nymphs with same data as holotype.
Deposited in KPCT, and one female in PPI.
Lit.: (Miller et al., 2013).
Proteriococcus lauri; Erkiliç, 2009 in Erkılıç et al. (2011), published by Kozár (2009: 105)
appears to be a nomen nudum.
Description
Unmounted female
Adult female dark red; in the white felt-like eggsacc. The felt sac was full with eggs with
same colour with female.
Mounted female
Body elongate oval, 2.30–2.38 mm long, 1.28–1.36 mm wide. Frontal tubercle present.
Eyes situated on venter near margin.
Venter: Labium 3 segmented, 90–85 µm long, 70–95 µm wide; basal segment not well
developed, but with two equal long setae present on each side; median setae on apex of
labium 8–10 µm long. Stylet loop long, reaching to level with between mesothrax legs
and posterior legs. Antennae 7 segmented, 260–290 m, length of segments: I: 50–52.5,
II: 35–45, III: 37.5–45, IV: 45–55, V: 30, VI: 25–32.5, and VII: 35–45 µm long; segment
II with 1 sensory pore; segment III with almost parallel sides; each segment covered
with a few, strong hair-like setae; apical segment with apical seta 62.5–72.5 µm long;
apical segment also with 3 falcate sensory setae, each 35–40 µm long; two preapical
segments each also with 1 sensory falcate seta 35–40 µm long. Legs well developed,
comparatively small: lengths of segments and digitules of prothoracic legs; coxa: 115–
125 µm, trochanter: 60–75 µm, femur: 130–155 µm, tibia: 100–125 µm; tarsus: 90–105
m and claw: 22.5–30 µm, trochanther+femur: 170–215 µm, tibia+tarsus: 180–225 µm,
tarsal digitules: 40–50 µm, claw digitules: 27.5–35 µm; lengths of segments and digitules
of mesothoracic legs; coxa: 120–135 µm, trochanter: 60–70 µm, femur: 135–140 µm,
tibia: 110–130 µm; tarsus: 100–120 µm and claw 25–30 µm, trochanther+femur: 160–190
µm, tibia+tarsus: 200–240 µm, tarsal digitules: 40–50 µm, claw digitules: 27.5–35 µm;
lengths of segments and digitules of metathoracic legs; coxa: 120–130 µm, trochanter:
60–70 µm, femur: 130–160 µm, tibia: 110–135 m; tarsus: 105–120 µm and claw: 27.5–30
µm, trochanther+femur: 180–210 µm, tibia+tarsus: 210–240 µm, tarsal digitules: 42.5–45
µm, claw digitules: 27.5–30 µm, each slightly knobbed. Mesothoracic and metathoracic
coxae with spinulae on anterior surface. Each trochanter with two pores on each side.
Claw with a small denticle. Legs with a few hair-like setae, and with one sensory pore
on tarsus. Tibia each with 4-5 setae, tarsus with 5 setae. Multilocular pores each 3–5 µm
in diameter and with 5 loculi, distributed in sparse rows on all abdominal and thoracic
segments. Diameter of anterior spiracles 40–50 µm; spiracular perithreme 20–25 µm.
Derm with a sparse covering of scattered flagellate hair-like setae, each about 12.5–110
µm long. Enlarged setae each 17.5–30 µm, without microtubular duct at base, situated
on submargine in 2 or 3 rows. Macrotubular ducts two sizes; larger one each 3.5–5.0 µm
wide and 17.5–20.0 µm long; placed on submargin, smaller one each 2.5 µm wide and
12.5–15.0 µm long; scattered on abdominal segments. Microtubular ducts and cruciform
pores absent. Suranal setae setose.
Dorsum: Marginal enlarged setae long, setose on apex, each 25–60 µm. Dorsal enlarged
setae conical, each seta 15–40 µm long, setae on thorax, head and anterior abdominal
segments slightly larger than setae on last 2 or 3 abdominal segments, arranged in
transverse rows across each body segment, rows irregular on head. Macrotubular ducts
wide, each 7.5–10 µm wide and 17.5–22.5 µm long, scattered throughout dorsum,
generally in the row. Microtubular ducts long, each 1.0–1.5 µm wide and 10.0–12.54 µm
long, scattered over dorsum, with 0–3 (generally 1 or 2) opennig at base of each dorsal
setae and marginal setae. Anal ring strongly sclerotized, with 16-20 pores on inner side
and large teeth on outer side of anal ring, 50–55 µm in diameter, with 6 setae, each 105–
135 µm long; anal ring situated on margin of dorsum. Anal lobes strongly developed and
sclerotized, each with three enlarged setae each 35–50 µm and 2-4 microtubular ducts on
dorsal surface; apical seta 190.0–262.5 µm; subapical setae 60–80 µm long. Cauda present
with large spinules as on dorsal surface.
Acanthococcidae 371
Figure 131. Orontesicoccus lauri (Erkılıç, 2011), female, first instar on top right. After Erkılıç
et al. (2011) with modifications.
Other stages

with strong sensory setae as in adult female. Stylet loop long, reaches behind posterior
coxae. Dorsum with six strong spines on head and thorax and six small setae forming two
median longitudinal rows. The marginal row of setae consist on last abdominal segments
1 strong sharp-pointed spines, the others are hair-like. Long microtubular ducts in a sparse
submarginal and a marginal row on dorsum. Venter with transverse rows of four small
hair-like median setae on each abdominal segment; and two rows of small submarginal
setae. Cruciform pores not seen. With one pair multilocular pores, on each thoracic and
abdominal segment, 1 near each spiracle, plus some on frons, 1 in middle-line of thorax
and 1 on each segment of abdomen. Anal lobe with 3 conical setae and with long apical
setae. Anal ring normal and with 6 hair-like setae (after Erkılıç et al. (2011)).
Ecology
Host plant: Laurus nobilis.
Biology: Found on the both side of leaves in high density.
Acanthococcidae 373
Genus:
Ovaticoccus Kloet, 1944
Type sp.: Coccus agavium Douglas, 1888: 150, by monotypy. Homonym of Gymnococcus
Zopf, 1887 Metazoa; discovered by Kloet, 1944: 86. Notes: Douglas (1888) stated,
in his description of Coccus agavium: "At one time I thought it might constitute the
type of a new genus, under the name of Gymnococcus, but in consideration of the
important and leading characters of the antennae, I have concluded (for the present,
at least) that it is better to regard all the others as specific, and to refer the species
to Signoret's genus Coccus." Cockerell (1894b) designated Gymnococcus as a distinct
genus and gave the credit to Douglas. Since Douglas did not actually use the name
Gymnococcus, and since Cockerell was the first person to do so, Cockerell is here
given credit for describing this genus (see also Hoy, 1963). Parrott (1900) considered
Gymnococcus valid but credited it to Newstead (1897). However, Gymnococcus Douglas
was considered valid until Kloet (1944) discovered that the name Gymnococcus was
preoccupied by a genus of Mycetozoa (Gymnococcus Zopf, 1887) and introduced the
new name Ovaticoccus. This name was disregarded until Ferris (1957b) confirmed
Kloet's designation.
Gymnococcus Douglas, 1888; Fernald, 1903: 79.

Gymnococcus Cockerell, 1894b; 1053; Hoy, 1963: 181.
Gymnococcus (Parrotia) Gómez-Menor Ortega, 1954: 142. Unavailable name; discovered
by Morrison & Morrison, 1966: 149. Notes: According to Morrison & Morrison (1966),
Gómez-Menor Ortega proposed Parrotia as a subgenus of Gymnococcus for two described
American species whose names he did not include. Since no type species was given, it is
unavailable according to Article 13 (b) of the 1961 code. Further, it is preoccupied by
Parrotia Kieffer (1924) in the Diptera.
Ovaticoccus Kloet, 1944: 86. Replacement name for Gymnococcus Cockerell 1894b.
Lit.: Balachowsky, 1942: 4; Boratynski, 1958: 173; Borchsenius, 1949: 322, 369;
Danzig, 1971a: 821; Gill, 1993: 173; Gómez-Menor Ortega, 1954: 141; Ferris, 1955:
179; Hodgson & Miller, 2010: 69; Hoy, 1962: 201; 1963: 181; Kloet, 1944: 86; Koteja,
1974b: 77; Koteja & Zak-Ogaza, 1981: 501; Kozár, 2009: 111; Kozár & Konczné
Benedicty, 2008a: 148; Kozár & Walter, 1985: 393; Miller 2005: 491; Miller & Gimpel,
2000: 426; Miller & McKenzie, 1967: 506; Miller & Miller, 1993: 84; Morrison &
Morrison, 1966: 141; Pellizzari & Kozár, 2011: 61; Tereznikova, 1981: 57; Tranfaglia,
1976: 129; Williams, 1985a: 384 (Miller et al., 2013).
Description
Ovisac elongate-oval, felt-like, white. Adult female oval. Antennae 6 or 7 segmented.
Frontal tubercle present. Labium 3 segmented, setae hair-like, basal segment with two
pair of short setae, apical segment with six setae; stylet loop slightly longer than labium.
Multilocular pores on venter, each with 5, or more loculi. Legs short, with tibia shorter
than tarsus, tibia with four setae. Claw with denticle, claw digitules knobbed, longer than
claw. All coxae with spinulae; posterior coxae also with large pores. Microtubular ducts
present. Cruciform pores present. Ventral setae short and hair-like. Some of dorsal spines
dome-shaped. Anal lobes not developed; ventral surface of each lobe with a long apical
seta and two shorter subapical seta. Quinquelocular pores numerous on all segments
of dorsum. Macrotubular ducts if present, heavily sclerotized, each with inner ductules
short, terminal gland not observed. Microtubular ducts short. Anal ring sclerotized, but
not well developed, with 2-8 setae, without anal ring pores, all shorter than diameter of
ring. Cauda absent.
Distribution
The species of Ovaticoccus genus known from Nearctic Region, especially in south-western
part of USA (Miller & McKenzie, 1967), except O. amplicoxae Williams & Martin, 2003,
which was described from Neotropic Region, and may not belong to Ovaticoccus genus.
The O. agavium as introduced species was found also in the Palaearctic and Ethiopian
regions. Two new species were described recently by Pellizzari & Kozár (2011). One
similar species Hispaniococcus agenjoi (Gómez-Menor Ortega, 1954), which was removed
from Ovaticoccus by Kozár & Konczné Benedicty (2008a), is known only from the
Palaearctic (Canary Island). Most of these species are found on grasses and herbaceous
plants belonging to Agavaceae, Liliaceae, Asteraceae and Poaceae.
Biology
Feed on leaves and leaf sheath of herbaceous plants.
Comments
O. agenjoi Gómez-Menor Ortega (1954: 142) (Gymnococcus) was transferred from Ovaticoccus
genus to the currently established Hispaniococcus Kozár (2008: 148) genus.
Key to species
1. – Ventral cruciform pores in transverse bands and groups across most of

abdominal segments........................................................................................... O. agavium
– Ventral cruciform pores rare in rows across abdominal segments.............................2
2. – Macrotubular ducts absent.................................................................................O. exoticus
– Macrotubular ducts present ....................................................................... O. agavecearum
Acanthococcidae 375
Ovaticoccus agavacearum Pellizzari & Kozár, 2011 (Fig. 132)

Ovaticoccus agavacearum Pellizzari & Kozár, 2011: 63.
Holotype: Female. Italy (Valenzano, Bari district), on potted Yucca sp., 19.v.2008, by G.
Pellizzari. type no. 1498/1. Notes: Paratypes: 23 adult females on 9 slides n.1498/2–
11; 1st instar nymphs, 2nd instar nymphs male and female, on slides1498/12-19,
same data as holotype. Deposited in DAFNAE, and one paratype deposited in PPI.
Lit.: Pellizzari & Kozár, 2011: 63 Miller et al. (2013).
Description
Unmounted female
Body of adult female oval, cuticle membranous.
Mounted female
Body of slide-mounted specimens, oval, 1.76–2.12 mm long, 1.42–1.63 mm wide.
Antennae 7 segmented, segment lengths; I: 26–36, II: 26–33, III: 26–30, IV: 23–40, V:
21–26, VI: 20–23 and VII: 29–32 µm long; all segments with few, setae about 17 µm
long, apical setae on segment VII 45 µm long, and with 3 sensory falcate setae, 18 µm
long, segments V and VI with single falcate seta 11 and 18 µm long, with two 4 µm long
coeloconic sensilla on apical segment. Frontal tubercles present. Eyes present on margin.
Venter: Labium 3 segmented, 67–72 µm long, basal segment not well developed,
with two pairs of setae, total number of setae 9 pairs, apical median setae half long
of the longest lateral ones. Stylet loop reaching the line behind the middle legs. Legs
well developed, lengths of segments of prothoracic legs; coxa 26–30, trochanter 22–32,
femur 65–68, tibia 48–56, tarsus 55–61, claw 21; lengths of segments of mesothoracic
legs; coxa: 26–35, trochanter: 26–30, femur: 62–72, tibia: 49–56, tarsus: 52–58, claw: 21;
femur: 70–75, tibia: 52–60, tarsus: 59–65, tarsal digitules knobbed, 33–36, claw 23, claw
digitules 24–26, knobbed. Coxae with spinulae on anterior (ventral) surfaces, hindcoxae
with translucent pores and spinulae on posterior (dorsal) surface. Each trochanter with 2
sensory pores on each surface. Claws with denticles. All legs with few flagellate setae and
with sensory pore at base of each tarsus. Mesothoracic spiracles 22 µm long in diameter,
with a group of quinquelocular pores at spiracular opening situated in a sclerotized
depression. Quinquelocular pores, sometimes with 3-7 loculi, 6 µm in diameter, scattered
in medium numbers on much of surface. Slender short setae 9–16 µm long, present,
scattered on abdomen. Some longer flagellate setae, present on head, mid abdomen and
thorax. Cruciform pores 5 µm long, present in rows on last abdominal segments, absent
in midthorax and four some scattered on submarginal of last abdominal segments.
Macrotubular ducts absent. Microtubular ducts present on margin. Oviduct or internal
genital organ barely discernible.
Dorsum: Setae on last abdominal segments robust, dome shaped spine-like, of one sizes,
6–7 µm long, some present on margin of last abdominal segments and on head and
thorax, usually 2-9 in number in one specimen it was 14. Small hair-like setae 5–8 µm
long, scattered on abdominal segments. Macrotubular ducts scattered all over dorsum,
12 µm long and 3 µm wide. Microtubular ducts short, sclerotized, each about 4 µm
long, with oval orifice, scattered around margin. Quinquelocular pores same as those on
venter, few, scattered on all segments. Anal ring situated on venter, weakly developed,
sclerotised 33–36 µm wide, 20–24 µm long, without pores, with 6 setae each 16 µm long,
two of them stronger. Anal lobes not developed, not sclerotized, with 2 spine-like setae.
Apical setae each 86 µm long. Cauda absent (after Pellizzari & Kozár (2011).
Other stages
Mounted first instar nymph. Antennae 6 segmented last three segments with strong sensory
setae as in female. Nymph with relatively large size of dome-shaped spines of three size
category, the larger are on last andominal segments. The number of spines usually six on
each segment. Tubular ducts absent. Some cruciform pores present on the submargin of
last abdominal segments, usually 1 on last 3-4 segments, quinquelocular pores scattered
on anterior abdominal segments and on venter of thorax. Anal ring with 3 pairs of spine-
like setae, not well seen on studied specimen, sclerotized plates and with sometimes 2
cruciform pores on the last 3 or 4 segments of abdomen.
Ecology
Host plant: Yucca sp.
Biology: On leaves.
Ovaticoccus agavium (Douglas, 1888) (Fig. 133)

Coccus agavium Douglas, 1888: 150.
lectotype: Female. United Kingdom (England, London, Kew Gardens, imported

from USA), on Agave sp., ?.ii.1885, D. Morris. By subsequent designation Boratynski,
1958: 175. Deposited in BMNH.
Lit.: Balachowsky, 1942: 4; Boratynski, 1958: 173; Borchsenius, 1949: 322, 369;
Danzig, 1971a: 821; Gill, 1993: 173; Gómez-Menor Ortega, 1954: 141; González,
2009: Ferris, 1955: 179; Hodgson & Miller, 2010: 69; Hoy, 1962: 201; 1963: 181;
Kloet, 1944: 86; Koteja, 1974b: 77; Koteja & Zak-Ogaza, 1981: 501; Kozár, 2009:
111; Kozár & Konczné Benedicty, 2008a: 148; Kozár et al., 2013: 56; Kozár & Walter,
1985: 393; Miller 2005: 491; Miller & Gimpel, 2000: 426; Miller & McKenzie, 1967:
506; Miller & Miller, 1993: 84; Morrison & Morrison, 1966: 141; Pellizzari & Kozár,
2011: 61; Tereznikova, 1981: 57; Tranfaglia, 1976: 129; Williams, 1985a: 384 (Miller
et al., 2013).
Acanthococcidae 377
Figure 132. Ovaticoccus agavacearum Pellizzari & Kozár, 2011, female and first instar on
the top right. After Pellizzari & Kozár (2011) with modifications.
Gymnococcus agavium; Cockerell, 1894b: 1053. Change of combination.

Ripersia agavium; Newstead, 1897: 12. Change of combination. Notes: Newstead made no
formal descriptions, but amended structural details of specific features.
Pseudantonina agaves; Chiaromonte, 1929: 61. Synonymy by Boratynski, 1958: 175.
Ovaticoccus agavium; Kloet, 1944: 86. Change of combination.
Description
Unmounted female
Adult female elongate-oval, rotund. Body pinkish-yellow; with clear brown areas on
dorsum between segments, arranged in two longitudinal lines. Segmentation clearly
visible both dorsally and ventrally (Parrott, 1900). Both surfaces lightly dusted with white
mealy secretion. Filamentous ovisac secretion often coalescing with ovisacs of other
females (Boratynski, 1958; Miller & McKenzie, 1967).
Mounted female
Body elongate oval no trace of anal lobes. 2.10–4.00 mm long, 1.10–2.70 mm wide.
Antennae 7 segmented; seventh segment with three sensory setae; sixth segment with
only one noticeably longer and narrower than that on segment five. Frontal tubercle
present. Eyes normal.
Venter: Labium 3 segmented, from apically acute to blunt, with hair-like setae. Legs
small; hind coxae not enlarged, usually with a few pores on either surface; inconspicuous
denticle on each claw. Multilocular pores of one kind, quinquelocular pores primarily
restricted to margin and submargin of body, in small numbers around spiracles.
Cruciform pores numerous; in lateral clusters on abdominal segments VIII through IV,
in medial clusters from segment VIII through mesothorax. Macrotubular ducts sparse,
varying from five to nine on entire surface. Microtubular ducts absent. Body setae long,
lanceolate arranged in clusters forming three longitudinal lines on abdominal segments
VIII through III; posterior anal lobe setae elongate.
Dorsum: Dorsal setae lanceolate infrequent, noticeably smaller than those on venter.
Enlarged setae dome-shaped, with truncated apexes; of several sizes; numerous over
entire surface. Multilocular pores of one kind; quinquelocular pores, common on entire
surface. Macrotubular ducts uncommon; present on abdominal segments IV-VIII.
Microtubular ducts uncommon; present on abdominal segments IV-VIII; rarely found
on margin of thorax (Miller & McKenzie (1967), with modifications).
Ecology
Host plant: Agave sp., A. americana, A. decipiens, A. beaueriana, A. lecheguilla, A. lophantha
var. poselgeri, A. parryi, A. sisalana, A. triangularis, A. utahensis var. nevadensis, Aloe sp., Dracaena
sp., Furcraea selloa, Yucca sp.
Acanthococcidae 379
Figure 133. Ovaticoccus agavium (Douglas, 1888), female. After Miller & McKenzie (1967)
with modifications.
Distribution: Eritrea, Ethiopia, France, Hungary, Italy, Russia, Ukraine, United Kingdom
(England), USA.
Ovaticoccus exoticus Pellizzari & Kozár, 2011 (Fig. 134)

Ovaticoccus exoticus Pellizzari & Kozár, 2011: 61.
Holotype: Female. Italy (Mozia, TP), Agave sp., on leaves, 26. viii, 1989, collected
by G. Pellizzari, No. 1546 in Pellizzari collection. By original designation. Paratypes,
16 adult females, on separate slides, same data as holotype. Holotype and paratypes
deposited in DAFNAE and one paratype deposited in PPI.
Lit.: Pellizzari & Kozár, 2011: 61 Miller et al. (2013).
Description
Unmounted female
Body of adult female oval, pinkish, cuticle membranous, covered with powdery wax and
wax filaments.
Mounted female
Body of slide-mounted specimens, oval, 1.55–2.67 mm long, 0.96–1.74 mm wide.
Antennae 7 segmented, segment lengths I: 28–31, II: 26–30, III: 28–30, IV: 26–28, V:
20–28, VI: 22–24, VII: 28–32 µm long; all segments with few about 22 µm long setae,
segment VII with apical setae 52 µm long, and with 3 sensory falcate setae, 18–20 µm
long, segments V and VI with single falcate seta 10 and 23 µm long, with two 4 µm long.
Frontal tubercles present. Eyes present on margin.
Venter: Labium 3 segmented, 60–79 µm long, basal segment not well developed, with two
pairs of setae, total number of setae 9 pairs, apical median setae half long of the longest
lateral ones; stylet loop reaches the line behind the middle legs. Legs well developed;
lengths of segments and digitules of prothoracic legs; coxa: 27–33, trochanter: 22–28,
femur: 66–73, tibia: 45–55, tarsus: 50–57, claw: 20–22 µm long; lengths of segments of
mesothoracic legs; coxa: 28–35, trochanter: 20–28, femur: 61–70, tibia: 50–53, tarsus:
51–54, claw: 20–22; lengths of segments and digitules of metathoracic legs; coxa: 34–40,
trochanter: 25–30, femur: 68–73, tibia: 45–53, tarsus: 50–58, tarsal digitules knobbed, 29–
35, claw 20–22, claw digitules 24, knobbed. All coxae with spinulae on anterior (ventral)
surfaces, hind coxae with translucent pores and spinulae on posterior (dorsal) surface.
Each trochanter with 2 sensory pores on each surface. Claws with denticles. All legs with
few flagellate setae and with sensory pore at base of each tarsus. Mesothoracic spiracles
21 µm in diameter, with several quinquelocular pores at spiracular opening. Multilocular
pores 5 µm in diameter, with 5 loculi, sometimes with 3 or 7 loculi, scattered in medium
numbers on much of surface. Slender short setae 9–16 µm long, present, scattered on
abdomen. Some longer flagellate setae, present on head and thorax. Cruciform pores 5
Acanthococcidae 381
Figure 134. Ovaticoccus exoticus Pellizzari & Kozár, 2011, female. After Pellizzari & Kozár
µm long, present in rows on last abdominal segments scattered in midthorax and four
small submarginal groups (2-5) on last abdominal segments. Macrotubular ducts absent.
Microtubular ducts absent.
Dorsum: Setae on last abdominal segments robust, dome shaped spine-like, of one
sizes, 7–8 µm long, present mostly in single rows across segments, about 6 present on
each abdominal segments. Small hair-like setae 7–9 µm long, scattered on abdominal
segments. Macrotubular ducts absent. Microtubular ducts short, sclerotized, each about 6
µm long, with oval orifice, scattered around margin. Quinquelocular pores same as those
on venter, scattered on all segments. Anal ring situated on venter, weakly developed,
sclerotized 36–41 µm wide, 22–26 µm long, without pores, with 6 strong setae each 20
µm long. Anal lobes not developed, not sclerotized, with 2 spine-like setae. Apical setae
each 113 µm long. Cauda absent (Pellizzari & Kozár (2011).
Ecology
Host plant: Agave americana.
Distribution: Italy (Sicily).
Biology: On leaves.
Acanthococcidae 383
Genus:
Proteriococcus Borchsenius, 1960
Type sp.: Proteriococcus acutispinatus Borchsenius, 1960c: 916.
Lit.: Ali, 1970: 80; Borchsenius, 1960c: 916; Erkılıç et al., 2011: 17; Hoy, 1963: 186;
Hua, 2000: 138; Kozár, 2009: 105; Kozár & Konczné Benedicty 2008b: 256; Köhler,
1998: 394; Miller & Gimpel, 2000: 441; Morrison & Moorison, 1966: 162; Tang &
Hao, 1995: 516, 597; Tao, 1999: 34; Wang, 2001: 235; Yang, 1982: 101 (Miller et al.,
2013).
Description
Antennae 7 segmented; frontal lobe present. Multilocular pores on venter only, each
with 5 loculi. Macrotubular ducts of two sizes with heavy rim at dermal orifice, scattered
on venter especially on margin, ducts present in high number all over the dorsum. Legs
normal, tibia equal size with tarsus. Claw with denticle. Tarsal and claw digital slightly
capitated. All coxae with spinulae; posterior coxae also with small pores. Hind tibia with
5 setae. Cruciform pores absent, instead of that, heavily sclerotized, oval microtubular
ducts present in a submarginal band. Ventral setae in small number, hair-like. Spine-like
setae present on dorsum in one-two rows on abdomen. Marginal row of spines setose.
Anal lobes well developed; dorsal surface of each lobe with three setose spines, spinulae
present on anal lobe, on cauda, and on derm, ventral surface of each lobe with a long
apical seta and shorter subapical seta. Anal ring sclerotized, well developed with one row
of anal ring pores and with 8 long setae. Microtubular ducts narrow, long, spread all over
dorsum among spines (according to Borchsenius (1960c) with 8-shaped orifice).
Distribution
Palearctic, with 1 species in Far-Eastern subregion.
Biology
Feed on the plant from Fagaceae.
Proteriococcus acutispinatus Borchsenius, 1960 (Fig. 135)

Proteriococcus acutispinatus Borchsenius, 1960c: 916.
Holotype: Female. China (Yunnan Province, 57 km south of Mindu), in a forest,

19.iv.1957, by N. Borchsenius. By original designation. Deposited in ZMAS.
Lit.: Borchsenius, 1960c: 916; Hoy, 1963: 186; Kozár, 2009: 105; Köhler, 1998: 394;
Miller & Gimpel, 2000: 442; Morrison & Morrison, 1966: 162; Tang & Hao, 1995:
516; Wang, 2001: 235 (Miller et al., 2013)
Description
Unmounted female
Body of adult female oval, closed in felted, brown-grey coloured sac.
Mounted female
Body of slide-mounted specimens, oval, about 3 mm long. Antennae 6 or 7 segmented,
terminal segment with 3 sensory falcate setae, and segments V and VI with single falcate
seta. Frontal lobe present, covered with microtrichia. Eyes present on margin. Anal lobes
well developed, strongly sclerotized, with 3 setose spine-like setae. Apical setae short,
strong.
Venter: Labium 3 segmented, basal segment not well developed, with 2 pairs of setae,
total number of setae 9 pairs, apical median setae about as long as the lateral ones;
Legs well developed, tarsal and claw digitules knobbed, longer than claw. All coxae with
spinulae on anterior (ventral) surfaces, hind coxae with translucent pores and spinulae on
posterior (dorsal) surface. Claws with denticles. All legs with few flagellate setae and with
sensory pore at base of each tarsus. Mesothoracic spiracles with several quinquelocular
pores at spiracular opening. Quinquelocular pores, numerous in a submarginal band on
abdomen and scattered in medium numbers on much of surface. Slender short setae
present, scattered. Some longer flagellate setae, present on head and thorax. Cruciform
pores absent, instead of that, heavily sclerotized, oval microtubular ducts present in a
submarginal band. Macrotubular ducts of two sizes, larger form a submarginal band, and
the smaller scattered on midventer. Microtubular ducts absent.
Dorsum: Enlarged setae of different sizes, elongated, conical, sharp-pointed, numerous.
Macrotubular ducts in high number, with sclerotized rim at dermal orifice. Microtubular
ducts long, narrow with oval orifice, scattered. Anal ring well developed, sclerotized
without pores, with 6 strong setae each 20 µm long. Cauda present, covered with
microtrichia, which are also present all over the dorsum (after Borchsenius (1960c) with
modifications).
Acanthococcidae 385
Ecology
Host plant: Lithocarpus sp., Pasania sp., Quercus sp.
Biology: Females on young branches, collected in April had not yet started to lay eggs.
Figure 135. Proteriococcus acutispinatus Borchsenius, 1960, female. After Borchsenius

Genus:
Pseudoacanthococcus Kaydan & Kozár Genus nova
Type sp.: Eriococcus osbeckiae Green, 1922, 353.
Description
Antennae 7 segmented; frontal lobe present. Multilocular pores on venter only, each
with 5 loculi. Macrotubular ducts of one sizes with heavy rim at dermal orifice, scattered
on venter especially on margin, ducts present in high number all over the dorsum. Legs
normal, tibia shorter than tarsus. Claw with a denticle. Tarsal and claw digitules slightly
capitated. All coxae with spinulae; posterior coxae also with small pores. Hind tibia with
4 setae. Cruciform pores few on thorax of venter. Ventral setae in small number, short,
hair-like. Enlarged setae present on dorsum in four longiditual rows, setae with (0-4)
microtubular duct at base. Marginal row of spines enlarged slightly blunted. Anal lobes
well developed; dorsal surface of each lobe with three enlarged setae, microtrichia present
on anal lobe, on cauda and on derm, ventral surface of each lobe with a long apical seta
and shorter subapical seta. Anal ring sclerotized, well developed with one row of anal
ring pores and with 6 long setae. Cauda present. Microtubular ducts narrow, long spread
all over dorsum among spines.
Etymology
The new genus is named after its similarity to the Acanthococcus genus.
Distribution
Oriental and Palaearctic with 2 species, one from Far-eastern subregion of Palaearctic
region.
Biology
Feed on the plant from family Lythraceae and Melastomataceae.
Comments
The genus Pseudoacanthococcus is distinct from all known acanthococcid genera known
in Palaearctic Region, except Orontesicoccus, by the presence of spines with 1 or 2
microtubular ducts associated to their bases. This special character shows an Oriental,
Pacific evolutionary and biogeographic connection. However, Orontesicoccus genus differs
from Pseudoacanthococcus by having spinose dorsal setae on margin and dorsum.
Acanthococcidae 387
Other species designated to this genus:

Pseudoacanthococcus rhodomyrti (Green, 1922). Combination nova.
Eriococcus rhodomyrti Green, 1922: 352. Sri Lanka: Nuera Eliya, Maskelliya; Badulla, at
altitudes of over 6000 feet. Lectotype, female, designated here. Ex coll. EEG, dry type
material from ZALF, DEI, Münchberg, thanks to Eckhard Groll for the kind help with
loan. Mounted by Z. Konczné Benedicty. P. rhodomyrti differs from P. osbeckiae by having
much higher number of small spines on dorsum and equal long spines on anal lobes.
Pseudoacanthococcus osbeckiae (Green, 1922) (Fig. 136) Redescription,

Combination nova
Eriococcus osbeckiae Green, 1922: 353.
Lectotype: Sri Lanka (Nanunakuli), on Osbeskia sp., ?/02/1910. Lectotype and

paralectotype, females and first instar embrios, on the same slide. Designated here. Ex
coll. EEG, dry type material from ZALF, DEI, Münchberg, thanks to Eckhard Groll
for the kind help with loan), mounted by Z. Konczné Benedicty. Other syntypes: Sri
Lanka (Badulla, Nanunakuli), on Osbeckia sp., ?.ii.1910. Syntypes, female. Deposited
in USNM and BMNH. Notes: There are ten adult female syntypes on two slides in
the BMNH (Williams, personal communication, May 15, 1996).
Lit.: Ali, 1970: 77; Hoy, 1963: 105; Ramakrishna, 1926: 452; Wang, 1982b: 142; 2001:
207 (Miller et al., 2013).
Nidularia osbeckiae; Lindinger, 1933a: 116. Change of combination.

Acanthococcus osbeckiae; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus osbeckiae; Tang & Hao, 1995: 448. Revived combination.
Acanthococcus osbeckiae; Köhler, 1998: 381. Revived combination.
Eriococcus osbeckiae; Miller & Gimpel, 2000: 289. Revived combination.
Acanthococcus osbeckiae; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Ovisac of female is creamy white, oblong oval, 2 mm long. Male sac is much smaller, 1
mm long.
Mounted female
Body elongate oval, 1.70 mm long, 1 mm wide. Frontal lobe present. Antennae 7
segmented, 210–225 µm, length of segments: I: 45–60, II: 32.5–35.0, III: 42.5–50, IV:
32.5–35, V: 20, VI: 22.5, and VII: 35 µm; segment III with almost parallel sides; each
segment covered with a few hair-like setae; apical segment with apical seta 65 µm long;
apical segment also with 3 falcate sensory setae, each 27.5–32.5 µm long; two preapical
segments each also with 1 sensory falcate seta 35–40 µm long. Eyes situated on venter near
margin each 17.5–20.0 µm in diameter. Anal lobes strongly developed and sclerotized,
covered by microtrichia, each with three enlarged setae (one outside 15 µm long, two
inside; shorter one 17.5–20 µm, and longer one 30 µm long and 2-4 microtubular ducts
on dorsal surface; apical seta 160–185 µm long.
Venter: Labium 3 segmented, 95.0 µm long, 92.5 µm wide; basal segment not well
developed, but with two different sizes setae present on each side; median setae on apex
of labium strong, 12.5 µm long, much shorter than lateral ones. Stylet loop not very long,
reaching to level with between mesothoracic legs. Legs well developed: anterior legs: coxa:
70–80 µm, trochanter: 30–40 µm, femur: 120–125 µm, tibia: 80 µm; tarsus: 105–110 µm
and claw 35 µm, tarsal digitules: 40–45 µm, claw digitules: 22.5 µm; mesothoracic legs:
coxa: 70.0–82.5 µm, trochanter: 30–40 µm, femur: 115–130 µm, tibia: 85–95 µm; tarsus:
110.0–112.5 m and claw: 25 µm, tarsal digitules: 40.0–42.5 µm, claw digitules: 25 µm;
metathoracic legs: coxa: 80–85 µm, trochanter: 35.0–42.5 µm, femur: 115 µm, tibia: 85–95
µm; tarsus: 110–120 m and claw: 30 µm, tarsal digitules: 45 µm, claw digitules: 25–30 µm,
each knobbed. Mesothoracic and metathoracic coxae with spinulae on anterior surface
and with some small pores. Claw with a denticle. Legs with a few hair-like setae, and
with one sensory pore on tarsus. Tibia each with 4 setae, tarsus with 4 setae. Multilocular
pores each 3–5 µm in diameter and with 5 loculi, distributed in sparse bands and rows
on all abdominal and thoracic segments. Diameter of spiracles 40–45 µm; spiracular
perithreme 20–27.5, posterior spiracle a bit larger than anterior spiracle. Derm with a
sparse covering of scattered, short flagellate hair-like setae, each about 10–60 µm long.
Macrotubular ducts similar sizes; each 6.0–7.5 µm wide and 15–20 µm long; scattered
on abdominal segments. Microtubular ducts absent. Cruciform pores each 2.5–3.0 µm
diameter, situated on submargin.
Dorsum: Marginal setae long, slightly blunted on apex, each 30–35 µm. Dorsal setae in
two sizes; larger one conical slightly blunted, each seta 32.5–35.0 µm long; associated
with 0-4 (generally 1 or 2) microtubular ducts at the base of each setae, arranged in
four logiditual rows across each body segment, rows irregular on head; the row in the
middle not complete, which has smaller enlarged setae on abdominal segments V–VII,
each 5.0-7.5 µm long, other smaller enlarged setae scattered among larger ones each
the same size with on last abdominal segments . Macrotubular ducts wide, and short,
each 7.5–9.0 µm wide and 17.5–20.0 µm long, scattered throughout dorsum, generally
in rows. Microtubular ducts long, each 1.0–1.5 µm wide and 7.5–9.0 µm long, scattered
over dorsum. Anal ring strongly sclerotised, with one complete pore row outside with 28
pores and inner side incomplete row with 8-10 pores, 45 µm in diameter, with 6 setae,
each 110–120 µm long; anal ring situated on margin of dorsum. Cauda present with large
microtrichia, as on dorsal surface (Redescription is based on type material).
Ecology
Host plant: Lagerstroemia sp., Osbeckia sp.
Distribution: China, Sri Lanka.
Biology: Lives on the leaf of plants. Species was collected at an elevation of over 6.000 feet.
Acanthococcidae 389
Figure 136. Pseudoacanthococcus osbeckiae (Green, 1922), female. Original.

Genus:
Rhizococcus Signoret, 1875 Reestablishment in sense of
Koteja (1974)
Type sp.: Rhizococcus gnidii Signoret, 1875: 16, 36, 37, by monotypy and original
designation. Notes: This genus was synonymyzed with Nidularia by Lindinger, 1933a:
107; with Eriococcus by Ferris, 1955: 94, it was a subjective decision (Miller & Gimpel,
2000: 105). Some have treated it as valid (Borchsenius, 1948; Danzig, 1962a: 839;
Kozár, 2008a: 105) while others have treated it as a junior synonym (Hoy, 1963; 132;
Miller & Gimpel, 2000: 105).
Lit.: Borchsenius, 1948: 501; 1949: 351; Danzig, 1962: 520; 1964: 632; 1975: 62; 1980:
205; Dziedzicka, 1977: 5; Dziedzicka & Koteja, 1971: 557; Fernald, 1903: 66; Hoy,
1963: 132; Kosztarab, 1959: 402; Kosztarab & Kozár, 1988: 298; Koteja, 1974b: 47;
Koteja & Zak-Ogaza, 1981: 515; Kozár, 2009: 105; Morrison & Morrison, 1966: 174;
Schmutterer, 2008: 84; Tang & Hao, 1995: Tereznikova, 1981:14 (Miller et al., 2013).
Description
Ovisac elongate-oval, encloses female completely, white to yellowish, 1.5–5.0 µm long,
0.8–2.2 µm wide. Adult female elongate-oval, tapering posteriorly. Antennae 7 (rarely 6)
segmented; eyes usually on ventral margin, close to bases of antennae; labium conical,
3 segmented, with 18 setae; legs well-developed, hind coxae, at least on dorsal surface,
with translucent pores (Dziedzicka, 1977); claws normally with a denticle, tarsal digitules
dilated and longer than claws; claw digitules either capitated and longer than claw or
spinelike and shorter than claw; disc pores on venter only, usually quinquelocular, but
sometimes with 3, 4, 6, 7 or 9-loculi; oval disc pores form a band along ventral margin;
tubular ducts of 3 types: large macrotubular ducts form transverse bands on dorsum
and marginal band on venter, small macrotubular ducts few, found scattered on venter,
except margin, microtubular ducts form transverse bands on tergites and a longitudinal
marginal band on sternites; dorsal enlarged conical setae of varying sizes: normally large
ones form a marginal row, median ones form a submarginal row and small ones form
transverse rows or bands on tergites; ventral setae vary: medium-size spinelike conical
setae form a row along body margin, small ones in an irregular submarginal band, hair-
like setae scattered on venter, but form transverse rows or groups in the median area;
anal ring with pores and 8, rarely 6 setae; anal lobes distinct, each with an apical seta,
3 dorsal conical setae and ventral hair-like setae, cauda usually well seen (Kosztarab &
Kozár, 1988).
In a comparative table (Fig. 137) we studied the differences of enlarged setae variations in
females and first instar nymphs of some species belonging to the genus Rhizococcus. The
types of enlarged setae are quite similar, what indicates the unity of these species in one
genus. However R. cistacearum differs from others by some capitate setae on middorsum
of female and hair-like setae on dorsum of the nymphs, by this character the nymph of
Acanthococcidae 391
Figure 137. Rhizococcus setae range on adult female and first instar.
R. thymeleae, differs as well. In this genus the number of spines, size and shape of studied
species is quite similar and for the identification of species we need also to use some
other characters.
Distribution
Palearctic, with 65 species. Probably part of species known in Nearctic Region and cited
in “Eriococcus” or “Acanthococcus” sensu lato belongs to this genus.
Biology
Most feed on herbaceous plant, some species on grasses.
Key to species
Notes: Several species could not be included into the key, because of the absence of
material, inadequate description, weak drawing: R. guesinus, R. henmii, R. kurdicus, R.
miscanthi, R. multispinosus, R. mumtazi, R. sachalinensis, R. teucricola, R. timidus, R. turkmenicus.
1. – Claw digitules setose, shorter than claw..........................................................................2

– Claw digitules capitated, longer than claw.......................................................................5
2. – Anal lobe with four enlarged setae........................................................R. gavrilovi sp. n.
– Anal lobe with three enlarged setae..................................................................................3
3. – Enlarged setae on body margin on each abdominal segments numbering four or
more, some hair-like setae on venter not capitated.......................................................4
– Enlarged setae on body margin on each abdominal segments numbering one or
two, hair-like setae on venter capitated.....................................................R. artemisiarum
4. – Enlarged setae on body margin in high number forming groups; ventral tubular
ducts in two sizes, smaller tubular ducts in groups on the submarginal area of the
abdomen........................................................................................................... R. arthrophyti
– Enlarged setae number on body margin four or five not in a group; ventral tubular
ducts in one size, smaller tubular ducts not in groups on the submarginal area of
the abdomen........................................................................................................ R. spinifera
5. – Some of enlarged dorsal setae capitated.....................................................R. cistacearum
– Non of enlarged dorsal setae capitated...........................................................................6
6. – Anal lobe setae hair-like................................................................................... R. borchsenii
– Anal lobe setae enlarged, normal spine-like....................................................................7
7. – Enlarged setae on abdominal segment VIII on the sclerotized bar..............R. rugosus
– Enlarged setae on abdominal segment VIII not on the sclerotized bar or absent...8
8. – Enlarged setae on dorsum truncated...............................................................................9
– Enlarged setae on dorsum blunted or sharp pointed..................................................11
9. – Enlarged setae on dorsum as wide as long;........................................................ R. thaleri
– Enlarged setae on dorsum long, 3 or 4 times longer than width;..............................10
Acanthococcidae 393
10. – Enlarged marginal setae on each abdominal segments numbering three, dorsal
enlarged setae generally much smaller than marginal enlarged setae, longest
enlarged dorsal setae present only on mid dorsum . .................................... R. coccineus
– Enlarged marginal setae on each abdominal segments numbering two, dorsal
enlarged setae various sizes generally same size as marginal enlarged setae, in row
on the body segment.......................................................................................R. devoniensis
11. – Dorsal enlarged setae on abdominal segment VIII present.......................................12
– Dorsal enlarged setae on abdominal segment VIII absent.........................................14
12. – Dorsal enlarged setae on abdominal segment VIII numbering 6-8, cruciform
pores a few on venter, only on head................................................................. R. desertus
– Dorsal enlarged setae on abdominal segment VIII numbering two, cruciform
pores present in the submarginal band on venter.......................................................13
13. – Enlarged dorsal setae on abdominal segments in two-three rows, mid-venter
without quiqelocular pores........................................................................ R. ningxianensis
– Enlarged dorsal setae on abdominal segments rare in one row, mid-venter without
quiqelocular pores...............................................................................................R. minimus
14. – Anal lobe with four enlarged setae . ..............................................................................15
– Anal lobe with three enlarged setae................................................................................17
15. – Inner side of anal lobe with three enlarged setae, enlarged setae on dorsum
blunted, some of setae on venter slightly capitated............................... R. saikwanensis
– Inner side of anal lobe with two enlarged setae, enlarged setae on dorsum sharp
pointed, non of setae on venter capitated....................................................................16
16. – Fourth enlarged setae on anal lobe (outher one) thin, almost spine like, enlarged
setae on abdominal segments VI and VII in a single row......................... R. hassanicus
– Fourth enlarged setae on anal lobe (outher one) thin, thick like other enlarged
setae on anal lobe, enlarged setae on abdominal segments VI and VII in double
rows........................................................................................................................R. cantium
17. – Dorsal enlarged setae various sizes but most of them 2/3 smaller than marginal
enlarged setae, small setae scattered all surface ..........................................................18
– Most dorsal enlarged setae almost equal size or slightly shorter than marginal
enlarged setae, if such small setae present restricted on posterior abdominal
segments only....................................................................................................................26
18. – Marginal enlarged setae on abdominal segment VII numbering three.................... 19
– Marginal enlarged setae on abdominal segment VII numbering two.......................21
19. – Quinquelocular pores on venter concentrated on abdominal segments very few on
thorax and head, dorsal enlarged setae slightly curved............................R. saxidesertus
– Quinquelocular pores on venter scattered all over head, thorax and abdomen,
dorsal enlarged setae not curved....................................................................................20
20. – Marginal enlarged setae in big groups on abdominal segment I and thorax with
4-5 rows of enlarged setae, cruciform pores present on submarginal area of
venter.............................................................................................................R. multispinatus
– Marginal enlarged setae not in big groups on abdominal segment I and thorax,
cruciform pores absent or a few on submarginal area of venter ............ R. cactearum
21. – Enlarged setae on venter in submarginal and submedian bands...............................22
– Enlarged setae on venter only in submarginal band....................................................24
22. – Cruciform pores present only on submarginal band on venter,
quinqelocular pores on venter mainly on abdominal segments a few
on head and thorax............................................................................................. R. astragali
– Cruciform pores present on submarginal, submadeian bands and median area
on venter, quinqelocular pores scattered on venter high number on abdominal
segments.............................................................................................................................23
23. – Dorsal enlarged setae in single row on head and thorax, cruciform pores on
midventer.......................................................................................................R. subterraneus
– Dorsal enlarged setae in two-three row on head and thorax, cruciform pores not
on midventer....................................................................................................... R. palustris
24. – Frontal lobe absent at the base of antennae, cruciform pores absent on venter
.............................................................................................................................. R. zygophyli
– Frontal lobe present at the base of antennae, cruciform pores present on venter
.............................................................................................................................................26
25. – Femur with large pores, microtubular ducts on mid-venter.................R. targassonensis
– Femur without large pores, microtubular ducts only on submarginal band
on venter.......................................................................................................R. nedimi sp.n.
26. – Enlarged setae on dorsum equal size with marginal enlarged setae, the size of
setae not reducing through posterior abdominal segment .......................................27
– Enlarged setae on dorsum equal size with marginal enlarged setae, the size of
setae reducing almost ¼ of marginal enlarged setae through posterior abdominal
segment..............................................................................................................................39
27. – Dorsal and marginal enlarged setae sharply pointed...................................................28
– Dorsal and marginal enlarged setae blunted, not sharply pointed............................31
28. – Antennae 6 segmented, dorsal enlarged setae slender three times longer than seta
base.....................................................................................................................................29
– Antennae 7 segmented, dorsal enlarged setae slender two times longer than seta
base.....................................................................................................................................30
29. – With two enlarged setae on the middle of abdominal segment VII on dorsum, on
dorsum abdominal segment VII with 20-24 tubular ducts.............................. R. greeni
– With four enlarged setae on the middle of abdominal segment VII on dorsum, on
dorsum abdominal segment VII with 8-10 tubular ducts................................R. acutus
Acanthococcidae 395
30. – Four or five marginal enlarged setae on abdominal segment VII, frontal lobe
present................................................................................................................. R. echinatus
– Two or three marginal enlarged setae on abdominal segment VII, frontal lobe
absent.................................................................................................................R. crassipinus
31. – Marginal enlarged setae on abdominal segment VII numbering four or more.......32
– Marginal enlarged setae on abdominal segment VII numbering less than four......34
32. – Enlarged dorsal setae on abdominal segment in band (with 2-3 rows), dorsum of
abdominal segment VIII without tubular duct............................................ R. matesovae
– Enlarged dorsal setae on abdominal segment in row, dorsum of abdominal
segment VIII with tubular duct......................................................................................33
33. – Frontal lobe present, some of ventral setae slightly capitated..................R. thymelaeae
– Frontal lobe absent, non of ventral setae capitated.......................................R. erinaceus
34. – Claw slender denticle not visible, antennae 6 segmented, frontal lobe absent
..........................................................................................................................R. ammophilus
– Claw normal denticle visible, antennae 7 segmented, frontal lobe present.............35
35. – Marginal enlarged setae on abdominal segment VII numbering three, enlarged
dorsal setae on abdominal segment V in a single row...................................... R. reynei
– Marginal enlarged setae on abdominal segment VII numbering less than three,
enlarged dorsal setae on abdominal segment V in at least two rows........................36
36. – Cruciform pores absent on venter, enlarged dorsal setae on abdominal segment
VII numbering more than 20 ....................................................................... R. marginalis
– Cruciform pores absent on venter, enlarged dorsal setae on abdominal segment
VII numbering maximum 12-14....................................................................................37
37. – Microtubular ducts on venter a few present only on submarginal area newer on
mid venter . .......................................................................................................... R. lactucae
– Microtubular ducts on venter numareous, present on mid venter............................38
38. – Enlarged dorsal setae on stout two times longer than seta base, microtubular
ducts on venter scattered all surface............................................................. R. brevenniae
– Enlarged dorsal setae on slender at least three times longer than seta base,
microtubular ducts on venter on submarginal area and mid venter........ R. istresianus
39. – Some of hairlike setae on venter slightly capitated......................................................40
– Hairlike setae on venter not capitated............................................................................41
40. – Antennae 6 segmented, cruciform pores on venter a few, present only on thorax
and head.............................................................................................................. R. helichrysi
– Antennae 7 segmented, cruciform pores on venter in a longiditual submarginal
band.......................................................................................................................R. gassinus
41. – Enlarged marginal setae on abdominal segment VII numbering two......................42
– Enlarged marginal setae on abdominal segment VII numbering
more than two...................................................................................................................46
42. – Marginal enlarged setae on abdominal segment VII not in equal size,
one is almost half length of larger one, some of enlarged setae on
dorsum wide and robust............................................................................. R. evinae sp.n.
– Marginal enlarged setae on abdominal segment VII about equal size, none of
enlarged setae on dorsum wide and robust..................................................................43
43. – Frontal lobe absent, enlarged setae on venter in a longiditual row on submarginal
area.............................................................................................................. R. heteroacanthus
– Frontal lobe present, enlarged setae on venter not in a longiditual row on
submarginal area...............................................................................................................44
44. – Hind coxae without large pores, enlarged dorsal setae on abdominal segment VII
numbering 2....................................................................................................R. puymorensis
– Hind coxae without large pores, enlarged dorsal setae on abdominal segment VII
numbering 4-6...................................................................................................................45
45. – Cruciform pores present on mid-venter, enlarged dorsal setae on mid-dorsum in
longiditual row....................................................................................................R. coronillae
– Cruciform pores absent on mid-venter, enlarged dorsal setae on mid-dorsum in
transverse band on segments............................................................................R. caudatus
46. – Frontal lobe present..........................................................................................................47
– Frontal lobe absent...........................................................................................................50
47. – Smallest (reduced) enlarged dorsal setae on abdominal segments VI and VII ......48
– Smallest (reduced) enlarged dorsal setae on abdominal segments IV and VII.......49
48. – Enlarged setae on dorsum of various sizes, three longiditual rows on dorsum,
quinquelocular pores present on mid-venter.................................................... R. munroi
– Enlarged setae on dorsum almost all of same size, three longiditual rows on
dorsum, quinquelocular pores absent on mid-venter...............................R. baldonensis
49. – Enlarged dorsal setae in longiditual band in mid-dorsum, enlarged marginal setae
on abdominal segment VII numbering three.......................................... R. micracanthus
– Enlarged dorsal setae not in longiditual band in mid-dorsum, enlarged marginal
setae on abdominal segment VII numbering more than three (four or five)..........50
50. – Cruciform pores on venter only on marginal area, a few . ............................. R. zernae
– Cruciform pores on venter in submarginal band with high number........................51
51. – Antennae 6 segmented.....................................................................................................52
– Antennae 7 segmented.....................................................................................................53
52. – Microtubular duct on venter present on mid-venter.................................... R. variabilis
– Microtubular duct on venter absent on mid-venter....................................... R. artiguesi
53. – Marginal setae sharp pointed, only quinquelocular pores present on venter
.....................................................................................................................................R. sojae
– Marginal setae blunted, multilocular pores in different size and types present on
venter.........................................................................................................................R. gnidii
Acanthococcidae 397
Rhizococcus acutus (Goux, 1938) (Fig. 138) Redescription, Combination

nova
Eriococcus acutus Goux, 1938: 327.
Holotype: Female. France (Bouches-du-Rhone, Marseille, Ste Marguerite), on

Piptatherum multiflorum, by L. Goux, 25. VI. 1933 (MNHN 8753-01, type no. 666/a).
Paratypes on 65 slides with the same data as the holotype (MNHN 8753-02-66). By
original designation. Deposited in MNHN.
Lit.: Foldi, 2001: 305; 2002: 245; Goux, 1948: 67; Hoy, 1963: 68; Köhler, 1998: 372;
Ouvrard & Kozár, 2009: 130; Pellizzari & Kozár, 2011: 66 (Miller et al., 2013).
Acanthococcus acutus; Kozár & Walter, 1985: 73. Change of combination.

Eriococcus acutus; Miller & Gimpel, 2000: 117. Revived combination.
Acanthococcus acutus acutus; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female is yellowish white and oval.
Mounted female
Body elongate oval, 2.03–2.68 mm long, 0.72–1.06 mm wide. Antennae usually 6
segmented, the size of the segments: I: 16–34, II: 30–37, III: 81–118, IV: 21–25, V:
20–22, and VI: 36–40 µm long; each segment covered with a few, strong hair-like setae
(except third segment); apical segment 25–33 µm long with apical seta and 3 sensory
falcate setae, the longest 28–34 µm long; falcate sensory setae on preapical segment 29–
39 µm long. Frontal lobe absent. Eyes situated on venter near margin. Anal lobes slightly
sclerotized each with two spinose setae along inner margin, and one spinose seta on outer
margin, similar in size to those on dorsum, plus a long apical seta 288–315 µm long.
Venter: Labium 3 segmented, 91–100 µm long; stylet loop as long as labium. Legs well
developed. Lengths of segments of prothoracic legs; coxa: 60–71, trochanter: 40–49,
femur: 141–168, tibia: 113–142, tarsus: 120–136, claw: 28–30 µm long. Lengths of
125–160, tarsus: 120–155, claw: 30–35 µm long. Lengths of segments and digitules of
metathoracic legs; coxa; 84–120, trochanter: 44–49, femur: 148–158, tibia: 130–160,
tarsus: 132–160, tarsal digitules knobbed: 46–60, claw: 28–30 µm, claw digitules: 30–40
µm long, slightly knobbed. Meso- and metathoracic coxae each with spinulae on ventral
surface, posterior coxae with medium number (less than 20) of small pores. The diameter
of anterior spiracles 28–33 µm; tibiae of legs each with 5 setae; tarsi each with 6 setae.
Claw with denticle. With a small number of scattered, hair-like setae and a submarginal
band of macrospines. Quinquelocular pores distributed in sparse bands and rows
on all segments of abdomen and thorax, 4–5 µm in diameter, some trilocular pores
also present. Macrotubular ducts about 4–5 µm wide and 12–14 µm long, scattered.
Microtubular ducts present in a submarginal band. Cruciform pores, or sessile pores
scattered on submarginal band, absent on middle of thorax. Some quinquelocular pores
found around spiracles, 5 µm in diameter.
Dorsum: Dorsal setae all conical, spine-like, each spine 34–78 µm long, with pointed
apex, varying in size and shape. Dorsal spinose setae forming 3 longitudinal medial
bands, except last two segments. Spines in middle part of last segments 34–42 µm long.
On margin of each segments 6 or 7 spines present, 65–78 µm long. Diameter of base of
median spine on penultimate segment 18–19 µm, and on margin 16–20 µm. Macrotubular
ducts similar to those on venter, sparse, present throughout, 6–9 µm wide, and 20–24
µm long. Microtubular ducts each 3 µm long, scattered throughout. Anal ring 52–63 µm
wide, 55–72 µm long, with pores and with 8 hair-like setae, 125–135 µm long. Cauda not
seen (Redescription is based on type material).
Other stages
First and second instar nymphs and winged adul female were described by Goux (1938).
Ecology
Host plant: Brachypodium pinnatum, Dactylis glomerata hispanica, D. hispanica, Piptatherum
multiflorum.
Distribution: France, Italy.
Comments
The drawing of specimens found in Italy differs from the present one, by the absence of
cruciform pores around margin, microtubular ducts not shown, or mixed with smaller
sized macrotubular ducts. Concerning the shape and size of spines, it looks more similar
to R. echinatus (Goux, 1938). The identification of Italian specimens needs further study.
Acanthococcidae 399
Figure 138. Rhizococcus acutus (Goux, 1938), female. Original.

Rhizococcus ammophilus ( Balachowsky, 1933) (Fig. 139) Redescription,

Combination nova
Eriococcus ammophilus Balachowsky, 1933, 46.
Lectotype: Female. France (Corsica, Ile Rousse) (no 4930/4). Paralectotypes: with
the same data of collection (4830/13, 14). Designated here. Notes: There are 14
syntype slides containing approximately 22 specimens (D. Matile-Ferrero, personal
communication, November 20, 1996). Deposited in MNHN.
Lit.: Foldi, 2001: 305; Goux, 1938: 333; Hoy, 1963: 69; Kozár & Walter, 1985: 73;
Köhler, 1998: 373; Lindinger 1936: 156, Ouvrard & Kozár, 2009: 101; Williams, 1969:
Nidularia ammophilus; Lindinger, 1936: 156. Change of combination.

Acanthococcus ammophilus; Kozár & Walter, 1985: 73. Change of combination.
Eriococcus ammophilus; Miller & Gimpel, 2000: 123. Revived combination.
Acanthococcus ammophilus ammophilus; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Ovisac 3.0 mm long, 1.2 mm wide.
Mounted female
Adult female oval, 1.5 mm long, 0.7 mm wide. Antennae 6 segmented, I: 43, II: 29, III:
86, IV: 26, V: 24, VI: 31 µm long; segments with a few hair-like setae; longest seta on
apical segment 27 µm long; apical segment also with 3 sensory falcate setae, each 29 µm
long; two preapical segments each also with 1 sensory falcate seta: on fourth 19, fifth 29
µm long. Frontal lobe and tubercle absent. Eyes situated on venter near margin. Spinulae
on segment margins both on venter and dorsum. Anal lobes well developed, sclerotized,
with 3 blunt, conical spines, ventrally each with two flagellate setae; anal lobe with 298
µm long apical seta.
Venter: Labium 3 segmented, joint length of two apical segments 96 µm; basal segment
with 2 pairs of setae, apical segment with 6 pairs of hair-like setae. Stylet loop reaching
median legs. Width of anterior spiracles 22 µm. Legs well developed, length of segments
and digitules of prothoracic legs: coxa:≈60, trochanter: 48, femur: 139, tibia: 106; tarsus:
115 and claw 34, tarsal digitules 58, claw digitules 34 µm long; lengths of segments and
digitules of mesothoracic legs; coxa: 60, trochanter: 43, femur: 116, tibia: 120, tarsus: 125,
claw: 34, tarsal digitules: 56, claw digitules: 39 µm long; lengths of segments and digitules
of metathoracic legs: coxa: 84, trochanter: 55, femur: 132, tibia: 132, tarsus: 136; claw: 36,
tarsal digitules: 51, claw digitules: 36 µm long, tarsal and claw digitules slightly knobbed,
longer than claw. All coxae with spinulae on anterior surface, more numerous on meso-
and metathoracic coxae; metathoracic coxae and femur also with a few small pores on
Acanthococcidae 401
Figure 139. Rhizococcus ammophilus (Balachowsky, 1933), female. Original.

both dorsal surfaces. Claw without a denticle. Legs with a few hair-like setae; tibia with
5 setae. Multilocular pores with 3 or 5 loculi, mostly 5; each quinquelocular pores 5 µm
in diameter, each trilocular pore 4 µm in diameter; distributed in transverse bands across
abdominal sternites, in sparse rows on thoracic segments and head region. Macrotubular
ducts of two sizes: larger macrotubular ducts each 6 µm wide and 25 µm long, situated on
margin; smaller macrotubular ducts each 4 µm wide and 20 µm long scattered throughout
the venter. Microtubular ducts sparse marginally and submarginally on head and thorax.
Cruciform pores 4 µm long; a few in marginal and submarginal area. A few hair-like setae
present on submedian and submargianal venter, submedian ones longer. Blunt conical
spines in two sizes; larger 19–30 µm long on margin; smaller ones ca. half the –size of
larger ones, present submarginally. Suranal setae hair-like.
Dorsum: Enlarged setae blunt conical, in two sizes; longer setae each 38–43 µm long in
a row on margins, 2 on each segment margin and a few on segments; shorter setae more
numerous, each about 19–30 µm long, forming bands on segments, but absent from
abdominal segment VIII. Twelve dorsal setae on abdominal segment VII. Macrotubular
ducts each 6 µm wide and 25 µm long, scattered among spines. Microtubular ducts
scattered, each 3 µm long. Anal ring situated on margin of dorsum; oval, sclerotized,
62 µm wide, with one row of pores and with 8 setae, each 91 µm long. Cauda not seen
(Balachowsky (1933), with modifications based on the lectotype and paralectotypes).
Ecology
Host plant: Ammophila arenaria arundinacea.
Distribution: France (Corsica).
Biology: From Ammophila arenaria at the beach.
Acanthococcidae 403
Rhizococcus artemisiarum (Matesova, 1976) (Fig. 140) Redescription,

Combination nova
Acanthococcus artemisiarum Matesova, 1976: 22.
Holotype: Female. Kazakhstan (Karaganda Oblast, Balchash), on Artemisia terrae-

albae, 28.v.1956, by original designation. Notes: Holotype should be in ZMAS, but E.
Danzig was unable to locate it. There are approximately 26 paratypes on 18 slides in
ZMAS and several additional slides in AAKA.
Lit.: Kozár & Walter, 1985: 73 (Miller et al., 2013).
Eriococcus artemisiarum; Tang & Hao, 1995: 457. Change of combination.

Acanthococcus artemisiarum; Köhler, 1998: 373. Revived combination.
Eriococcus artemisiarum; Miller & Gimpel, 2000: 135. Revived combination.
Acanthococcus artemisiarum; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Females greenish-brown.
Mounted female
215–230 µm long, length of segments: I: 55–60, II: 30–35, III: 45.0–52.5, IV: 35–45, V:
17.5–22.5, VI: 17.5–25.0 and VII: 30.0–32.5 µm long; each segment covered with a few,
strong hair-like setae (except third segment without setae); apical segment with apical seta
40.0–47.5 µm long; apical segment also with 3 sensory falcate setae, each 22.5–35.0 µm
long; segment VI with 1 sensory falcate seta, 25–32 µm long, segment V with 1 sensory
falcate seta, 17.5 µm long. Frontal lobe absent, frontal tubercle present. Eyes situated
on venter near margin. Anal lobes strongly developed, each with 3 enlarged setae, each
25–35 µm plus 2 or 3 microtubular ducts on dorsal surface; apical seta 240 µm; ventral
hair-like subapical seta long.
short, 10.0–12.5 µm long. Legs well developed; lengths of segments and digitules of
prothoracic legs: coxa 50–60, trochanter 37.5–55, femur 107.5–115, tibia 75.0–82.5,
tarsus 92.5–95; claw 32.5–35.0, trochanther + femur 150–160, tibia + tarsus 165–185,
tarsal digitules 32.5–37.5, claw digitules 10–11 µm long; lengths of segments and digitules
of mesothoracic legs: coxa 57.5–65, trochanter 45–55, femur 100–105, tibia 90.0–92.5,
tarsus 95–100 and claw 30–35, trochanther + femur 147.5–152.5, tibia + tarsus 185–
190, tarsal digitules 37.5–40.0, claw digitules 10–12 µm long; lengths of segments and
digitules of metathoracic legs: coxa 62.5–70.0, trochanter 45–55, femur 110–115, tibia
95.0–102.5, tarsus 105–110 and claw 32.5–35.0, trochanther + femur 155–160, tibia +
tarsus 205–210, tarsal digitules 37.5–40, claw digitules 32.5–35.0 µm long, tarsal digitules
slightly capitated, claw digitules spine like and shorter than claw. Metathoracic coxae each
with spinulae and pores on ventral surface on coxa. Tibiae each with 5 setae (median seta
present), tarsi each with 5 setae. Length of spiracles 50–65 µm; diameter of spiracular
peritreme 25–30 µm, posterior spiracles slightly larger than anterior. Derm with a sparse
covering of scattered slightly blunted hair-like setae, each 12.5–40 µm long. Enlarged
setae, situated on submargin in 1 or 2 rows, each 15.0–17.5 µm long. Multilocular pores
each 5–7.5 µm in diameter and with 5 loculi, distributed all abdominal and thoracic
segments and a few on head. Macrotubular ducts of one size, each 5.0–7.5 µm wide and
17.5–22.5 µm long, scattered on abdomen, thorax and head. Microtubular ducts, each
3–5 µm long scattered over venter. Cruciform pores absent.
Dorsum: Marginal setae broad, truncated, wide at the base, long, each 30–35 µm, forming
a marginal row, each segment with one or two setae and not strongly differentiated
from dorsal setae; other dorsal setae truncated, each setae 12.5–22.5 µm long, those on
or 3 abdominal segments; arranged in transverse row across each body segment, rows
irregular on head. Macrotubular ducts, each 5.0–7.5 µm wide and 17.5–22.5 µm long,
scattered throughout dorsum, generally in segmental bands. Microtubular ducts with 2
sclerotized areas, each 3–5 µm long scattered over dorsum. Anal ring strongly sclerotized,
with on each inner side, 55–65 µm in diameter, with 8 setae, each 95–130 µm long; anal
ring situated on margin of dorsum. Cauda not seen (Redescription is based on type
material).
Ecology
Host plant: Artemisia gurganica, A. terrae-albae.
Biology: On the roots, subterranean part of the stem.
Acanthococcidae 405
Figure 140. Rhizococcus artemisiarum (Matesova, 1976), female. Original.

Rhizococcus arthrophyti (Borchsenius, 1949) (Fig. 141) Redescription,

Combination nova
Acanthococcus arthrophyti Borchsenius, 1949: 341.
Lectotype: Female. Turkmenistan (Repetek), on Haloxylon aphyllum, 23.vii.1944, by N.

Borchsenius. Lectotype female, by subsequent designation Danzig, 1996: 521. Notes:
The type series includes 2 adult females on same slide as lectotype and 2 adult females
on a second slide. Deposited in ZMAS.
Common name: Saxaul felt scale.
Lit.: Danzig, 1996: 574; Hoy, 1963: 72; Köhler, 1998; Lashin, 1956: 114; Tang & Hao,
1995: 452.
Nidularia arthrophyti; Lindinger, 1957: 543. Change of combination.

Eriococcus arthrophyti; Hoy, 1963: 72. Change of combination.
Acanthococcus arthrophyti; Kozár & Walter, 1985: 73. Revived combination.
Eriococcus arthrophyti; Miller & Gimpel, 2000: 135. Revived combination.
Acanthococcus arthrophyti; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Female oval, darkolive coloured, 3.5 mm long, 2.2 mm wide. Egg sac not seen.
Mounted female
Body elongate oval. Antennae 7 segmented, 215–230 µm long, length of segments: I: 54,
II: 62, III: 70, IV: 51, V: 35, VI: 32 and VII: 37 µm long; each segment covered with a few,
strong hair-like setae (except third segment without setae); apical segment with 3 sensory
falcate setae, segment VI and VII with 1 sensory falcate seta. Frontal lobe present. Eyes
situated on venter near margin. Anal lobes not well developed, each with 3 enlarged setae,
40 µm plus 2 or 3 microtubular ducts on dorsal surface; apical seta 165 µm; ventral hair-
like subapical seta 90 µm long.
Venter: Labium wide, median setae on apex of labium short spine-like, stylet loop short,
one and half time longer than labium. Legs well developed, femur 360–415, tibia 300–
340, tarsus, 230–265 µm long, and claw with denticle, tarsal digitules slightly capitated,
claw digitules spine like and shorter than claw. Meso- and metathoracic coxae with
spinulae, metathoracic coxae and femur with large group of pores on ventral surface on
coxa. Anterior and median tibiae with 5 setae (median seta present), on posterior only 4,
tarsi with 5 or 6 setae. Derm with a sparse covering of scattered strong, hair-like setae,
up to 65 µm long. Enlarged setae absent. Multilocular pores with 5 loculi, in bands on
last abdominal segments and a few on thorax and head. Macrotubular ducts of one size,
each 9–11 µm wide and 33 µm long, forming a wide submarginal band and scattered
Acanthococcidae 407
Figure 141. Rhizococcus arthrophyti (Borchsenius, 1949), female. Original.

on midthorax and head. Microtubular ducts, each 3–5 µm long scattered over venter.
Cruciform pores absent.
Dorsum: Marginal setae slightly blunted, wide at the base, long, each 30–55 µm, forming
a marginal band, other dorsal setae blunted, each setae 15–30 µm long, those on thorax,
head and anterior abdominal segments slightly larger than setae on posterior 2 or 3
abdominal segments; arranged in transverse bands across body segments. Macrotubular
ducts, each 11 µm wide, 35 µm long, numerous throughout dorsum, generally in
segmental bands. Microtubular ducts, each 3–5 µm long scattered over dorsum. Anal
ring strongly sclerotized, with partially double row of pores, with 8 setae, 140 µm long.
Cauda developed (after Borchsenius (1949), with modifications and changes based on
type material).
Ecology
Host plant: Haloxylon sp., H. ammodendron.
Distribution: Mongolia, Turkmenistan.
Biology: Lives on root crown and tall roots of black and white saxaul.
Rhizococcus artiguesi (Goux, 1991), (Fig. 142) Redescription, Combination

nova
Eriococcus artiguesi Goux, 1991: 48.
Holotype: Female. France (Hautes-Pyrénées, Artigues), on unidentified Poaceae,

12.viii.1953, by L. Goux. By original designation. Notes: the paratype slide (MNHN
14435-02) contains only eggsacs and eggs. Deposited in MNHN.
Lit.: Foldi, 2001: 305; Kozár et al., 2013: 56; Kozár & Walter, 1985: 73; Miller &
Gimpel, 1996: 598; Ouvrard & Kozár, 2009:101 (Miller et al., 2013).
Acanthococcus artiguesi; Miller & Gimpel, 1996: 598. Change of combination.

Eriococcus artiguesi; Miller & Gimpel, 1996: 598. Revived combination.
Acanthococcus artiguesi; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Unknown.
Mounted female
segments: I: 38, II: 32, III: 97, IV: 26, V: 24 and VI: 38 µm long; each segment covered
with a few, strong hair-like setae (except third segment without setae); apical segment with
Acanthococcidae 409
Figure 142. Rhizococcus artiguesi (Goux,1991), female. Original.

apical seta 35 µm long; apical segment also with 3 sensory falcate setae, longest 26 µm
long; segment IV-V with 1 sensory falcate seta 32 µm long. Frontal lobe absent, frontal
tubercle present. Eyes situated on venter near margin. Anal lobes slightly sclerotised each
with two spinose setae along inner margin, and one spinose seta on outer margin, similar
in size to those on margin of dorsum, apical seta 336 µm long.
Venter: Labium apparently 3-segmented, 96 µm long. Stylet loop not well seen. Legs:
anterior legs: coxa 77, trochanter 53, femur 151, tibia 113, tarsus 125, claw 30 µm long;
middle legs: coxa 79, trochanter 58, femur 139, tibia 127, tarsus 132, claw 30 µm long;
posterior legs: coxa 92, trochanter 60, femur 144, tibia 130, tarsus 140, tarsal digitules
knobbed, 40, claw 34, claw digitules 33 µm long, slightly knobbed. Posterior coxae and
femur with medium number of small pores, without spinulae. Claw with a denticle.
Legs with few hair-like setae, and with one sensory pore on tarsus. Quinquelocular
pores distributed in sparse bands and rows on all segments of abdomen and thorax,
5 µm in diameter, some of them have three loculi around spiracles. The diameter of
anterior spiracles 31 µm. Venter with a small number of scattered, hair-like setae and
two submarginal rows of spines. Microtubular ducts present in a submarginal band.
Macrotubular ducts each about 3 µm wide, length not measurable, scattered. Cruciform
pores 5 µm long, scattered on submarginal band, and on head, absent on middle of
thorax.
Dorsum: All setae spine-like, each seta 19–55 µm long, varying in size and shape, conical
with acute apices. Dorsal spinose setae of one size, forming sparse rows. Spines in middle
part of last segments 19 µm long. On margin of segments 3-4 spines present, 50 µm long.
Macrotubular ducts similar to those on venter, sparse, present throughout, 6 µm wide
and 22 µm long. Microtubular ducts, with 2 sclerotized areas, each 3 µm long, scattered
throughout. Disc pores absent. Anal ring not well seen, 54 µm wide and 80 µm long,
with partially double rows of pores, with 8 hair-like setae, 110 µm long. Cauda not seen
(Redescription based on type materials).
Ecology
Host plant: Poaceae, Thymus glabrescens.
Distribution: Hungary, France.
Biology: Unknown.
Acanthococcidae 411
Rhizococcus astragali Kaydan Species nova (Fig. 143)
Holotype: Female. Turkey (Van-Özalp-Dönerdere, N: 38°41’414’’, E: 044°07’910’’,

2091 m altitude), on Astragalus sp., M.B. Kaydan, 05.vi.2005. Paratypes: 5 adult
females, same data as holotype in same slide; Van-Özalp-Dönerdere, N: 38°41’414’’,
E: 044°07’910’’, 2091 m altitude, on undetermined plant, M.B. Kaydan, 05.vi.2005.
Deposited in KPCT.
Additional material: 2 adult females, Turkey (Van-Başkale-Hakkari road, N: 37°54’
181’’, E: 044°04’945’’, 1913 m altitude), on Silene sp., M.B. Kaydan, 16.vi.2006.
Description
Unmounted female
Unknown.
Mounted female
270–285 µm, length of segments: I: 47.5–65, II: 35–45, III: 55.0–67.5, IV: 45.0–52.5,
V: 25–30, VI: 25.0–27.5, and VII: 32.5–47.5 µm long, each segment covered with a few,
strong hair-like setae (except third segment); apical segment with apical seta 47.5–52.5 µm
VI with 1 sensory falcate seta 30 µm long, segment V with 1 sensory falcate seta 12.5–
15.0 µm long. Frontal lobe absent, frontal tubercle present. Eyes situated on venter near
margin. Anal lobes developed, sclerotized, each with 3 enlarged setae, each 17.5–25.0 µm
plus 4–6 microtubular ducts on dorsal surface; apical seta 255–335 µm; ventral hair-like
subapical seta 77.5–85 µm long.
Venter: Labium 155.0–185 µm long, 105.0–127.5 µm wide, median setae on apex
of labium short, 12.5–15.0 µm long. Legs well developed; lengths of segments and
digitules of prothoracic legs; coxa: 75.0–82.5, trochanter: 52.5–62.5, femur: 130–145,
tibia: 105–130, tarsus: 110–120 and claw: 37.5, trochanther + femur: 182.5–195.0, tibia
+ tarsus: 205–235, tarsal digitules: 35.0–37.5, claw digitules: 25.0–27.5 µm long; lengths
of segments and digitules of mesothoracic legs; coxa: 80–85, trochanter: 55–60, femur:
132.5–137.5, tibia: 110–130, tarsus: 105–120 and claw: 37.5, trochanther + femur: 180.0–
187.5, tibia + tarsus: 220.0–247.55, tarsal digitules: 37.5, claw digitules: 30 µm long;
femur: 130–135, tibia: 130, tarsus 125–135 and claw: 37.5, trochanther + femur: 192.5–
207.5, tibia + tarsus: 245–275, tarsal digitules 37.5, claw digitules capitated, 25.0–32.5 µm
long, and longer than claw. Meso- and metathoracic coxae each with spinulae on ventral
surface and translucent pores on metathoracic coxa (13-17) and femur (4-9). Tibiae each
with 5 setae (median seta present), tarsi each with 6 setae. Length of spiracles 60.0–77.5
µm; diameter of spiracular peritreme 32.5–37.5 µm, posterior spiracles slightly larger than
anterior. Setae on venter spinose, each 12.5–47.5 µm long. Enlarged setae, situated on
submargin in 2 or 3 rows, each 12.5–25.0 µm long. Multilocular pores each 5.0–7.5 µm in

diameter and with 3, 5 and 7 loculi, distributed in sparse bands on all abdominal segments
and scattered on thoracic segments especially around spiracles. Macrotubular ducts of
two size, smaller macrotubular ducts each 3–4 µm wide and 15.0–20.0 µm long, present
on submedian of abdominal segments I–VI, thorax and head, larger macrotubular ducts
each 5.0–7.5 µm wide and 20.0–22.5 µm long, sparse in band on last abdominal segments,
and on submarginal band on abdomen, thorax and head. Microtubular ducts sclerotized,
with oval orifice, present on all venter, 5.0–6.0 µm long; cruciform pores scattered on
thorax and hand on generally on the submedian and median part, each 5.0–6.0 µm in
diameter.
Dorsum: Marginal dorsal setae spine-like each 25.0–52.5 µm forming a marginal row
and differentiated from dorsal setae; other dorsal setae conical, various size, each seta
10–25 µm long, those on thorax, arranged in transverse rows across each body segment,
rows irregular on head. Macrotubular ducts, each 8–10 µm wide and 20.0–22.5 µm long,
scattered throughout dorsum, generally in segmental bands. Microtubular ducts, each
5.0–7.0 µm long with oval dermal orifice, scattered over dorsum. Anal ring strongly
sclerotized, with 17-21 pores on each inner side, 60–75 µm in diameter, with 8 setae, each
95–115 µm long; anal ring situated on margin of dorsum. Cauda present.
Etymology
The new species is named after host plant Astragalus genus.
Ecology
Host plant: Astragalus sp., Silene sp.
Biology: Unknown.
Acanthococcidae 413
Figure 143. Rhizococcus astragali Kaydan sp. n., female.

Rhizococcus baldonensis (Rasina, 1966), (Fig. 144) Combination nova

Acanthococcus baldonensis Rasina, 1966: 18.
Holotype: Female. Latvia (Baldone), on Vaccinium vitis-idaea, 03.viii.1948, by A.

Rasina. By original designation. Deposited in Riga: Museum of the Plant Protection
Institute, Latvia.
Lit.: Danzig, 1975: 43; 1980: 206; Gertsson, 2001: 125; Kozár et al., 2004: 59; Kozár, et
al., 2002: 38; Kozár et al., 2013: 56; Kozár & Walter, 1985; Malumphy, et al., 2010: 258;
Tang & Hao, 1995: 452; Tereznikova, 1981: 21, Vikber, 1991: 4 (Miller et al. 2013).
Eriococcus baldonensis; Tang & Hao, 1995: 459. Change of combination.

Acanthococcus baldonensis; Köhler, 1998: 373. Revived combination.
Eriococcus baldonensis; Miller & Gimpel; 2000: 141. Revived combination.
Acanthococcus baldonensis; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female oval and brown, 2 mm long. Ovisac dirty white, smooth. Eggs pink.
Mounted female
Antennae 7 segmented. Frontal lobe present. Apical labial segment with 6 pairs of long
setae. Anal lobes not sclerotized, dorsally with 3 enlarged setae, ventrally with 2 slender
hair-like setae and with suranal setae.
Venter: Labium wide, median apical setae short. Legs thin; hind coxae and femur with
spinulae on anterior surface, pores present on posterior coxae, tibia with 5 setae elongate,
not enlarged; tarsi longer than tibiae, claws with denticle near tip. Tarsal and claw digitules
longer than claw, slightly capitated. Hair-like setae elongate. Enlarged setae of small size
only, present along body margin in two sparse rows. Macrotubular ducts of 2 kinds:
larger size in high numbers along body margin; smaller size in reduced numbers in medial
or sublateral areas. Microtubular ducts absent. Quinquelocular pores present in sparse
bands on last abdominal segments, and scattered elsewhere. Cruciform pores present
along lateral margin.
Dorsum: Enlarged setae blunted or truncated, normally of 3 sizes, relatively short: 2 or
3 larger setae along margin of abdominal segments and on margin of thorax and head,
on last abdominal segments spines form 1 or 2 irregular rows of shorter spines, in row
or bands on others. Macrotubular ducts present over dorsum, forms a marginal band.
Microtubular ducts numerous, narrow, short. Anal ring dorsal, with 4 pairs of setae.
Cauda present. (After Danzig, 1975; with modifications based on the paratype).
Acanthococcidae 415
Figure 144. Rhizococcus baldonensis (Rasina, 1966), female. After Danzig (1980) with
modifications.
Ecology
Host plant: Calluna vulgaris, Empetrum nigrum, Rhododendron tomentosum, Vaccinium uliginosum,
V. vitis-idaea.
Distribution: Finland, Greece, Hungary, Latvia, Lithuania, Russia, Ukraine.
Rhizococcus borchsenii (Danzig, 1975) (Fig. 145) Combination nova

Acanthococcus borchsenii Danzig, 1975: 67.
Holotype: Female. Russia (Southern Primor'ye, Khasanskiy district), on Artemisia sp.,

30.vii.1949, by N. Borchsenius. By original designation. Deposited in ZMAS.
Lit.: Danzig, 1980: 206; Fang, et al, 2001: 108; Kozár & Walter, 1985: 74; Kwon &
Han, 2003: 156; Tao, 1999: 31-32; Wang, 2001: 208 Miller et al. 2013.
Eriococcus borchsenii; Tang & Hao, 1995: 461. Change of combination.

Acanthococcus borchsenii; Köhler, 1998: 374. Revived combination.
Eriococcus borchsenii; Miller & Gimpel, 2000: 146. Revived combination.
Acanthococcus borchsenii; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female is oval, 2.5 mm long.
Mounted female
Antennae 7 segmented. Frontal lobe present. Apical labial segment with 6 (?) pairs of
long setae. Anal lobes strongly protruding, apically acute, not sclerotized, dorsally with 3
hair-like setae, ventrally with 2 slender hair-like setae and with suranal setae.
Venter: Labium wide, median apical setae short. Legs strong; hind coxae and femora
with spinulae on anterior surface, large group of pores present on posterior coxae, tibia
with 5 setae elongate, not enlarged; tarsi longer than tibiae, claws with denticle near
tip. Tarsal and claw digitules longer than claw, slightly blunted. Hair-like setae elongate.
Enlarged setae of small size only, present along body margin in a sparse band and a row.
Macrotubular ducts of 2 kinds: larger size in high numbers along body margin; smaller
size in reduced numbers in medial or sublateral areas. Microtubular ducts scattered on
margin. Quinquelocular pores present in sparse bands on last abdominal segments, and
scattered elsewhere. Cruciform pores present along lateral margin of thorax.
Dorsum: Enlarged setae blunted or truncated, normally of 3 sizes, 2 or 3 larger setae with
smaller ones forma a marginal band, on last 3 abdominal segments spines form 1 or 2
irregular rows of spines, and bands on others. Macrotubular ducts present over dorsum.
Microtubular ducts with 2 sclerotized areas, numerous, narrow, short. Anal ring dorsal,
Acanthococcidae 417
Figure 145. Rhizococcus borchsenii (Danzig, 1975), female. After Danzig (1980) with
modifications.
with partially double rows of pores, with 4 pairs of setae. Cauda present (after Danzig,
1975; with modifications based on the paratype).
Ecology
Host plant: Artemisia sp.
Distribution: China, Mongolia, North Korea, Russia.
Biology: On the roots, or root crown.
Rhizococcus brevenniae (Goux, 1993) (Fig. 146) Redescription,

Combination nova
Eriococcus brevenniae Goux, 1993: 66.
Holotype: Female. France (Rhône, Courzieu), on Calluna vulgaris, 07.10.1928,

(MNHN 14441), by L. Goux. By original designation. Notes: On slide with holotype,
a paratype female present. Deposited in MNHN.
Lit.: Foldi, 2001: 305; Miller & Gimpel, 1996: 599; Miller & Gimpel, 2000: 149;
Ouvrard & Kozár, 2009: 101 (Miller et al. 2013).
Acanthococcus brevenniae; Miller & Gimpel, 1996: 599. Change of combination.

Eriococcus brevenniae; Miller & Gimpel, 1996: 599. Revived combination.
Acanthococcus brevenniae; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Unknown.
Mounted female
segments: I: 29–30, II: 18–22, III: 38–40, IV: 36–41, V: 14–15, VI: 15–16 and VII: 29–32
µm long; each segment covered with a few, strong hair-like setae (except third segment
without setae); apical segment with apical seta 35–38 µm long; apical segment also with
3 sensory falcate setae, the longest 26–29 µm long; on the preapical segment the falcate
sensory seta 22–25 µm long. Frontal lobes present. Eye visible, situated on venter. Anal
lobes slightly sclerotized each with two enlarged spinosa setae along inner margin, and
one enlarged spinose seta on outer margin, similar in size to those on margin of dorsum,
apical seta 144–150 µm long.
Venter: Labium 96 µm long, with long hair-like setae, styletloop as long as labium. Legs
well developed; lengths of segments of prothoracic legs; coxa: 43–50, trochanter: 42–
52, femur: 104–108, tibia: 81–84, tarsus: 89–90 and claw: 25–29 µm long; lengths of
Acanthococcidae 419
Figure 146. Rhizococcus brevenniae (Goux, 1993), female. Original.

84–90, tarsus: 96–99 and claw: 27–29 µm long; lengths of segments of metathoracic
legs; coxa: 61–67, trochanter: 48–55, femur: 105–110, tibia: 89–90, tarsus: 101–105 and
claw: 29–30; tarsal digitules knobbed, 38–48; claw digitules slightly knobbed 34–35 µm
long. Posterior coxae and femur with small number of small pores, with some spinulae.
Claw with denticle. Tibiae each with 5 setae (median seta present), tarsi each with 5 setae.
Length of anterior spiracles 41–44 µm, posterior spiracles slightly larger than anterior.
Derm with a small number of scattered, hair-like setae and with a submarginal band
of enlarged setae. Quinquelocular pores distributed in sparse bands and rows on all
segments of abdomen and thorax, 5 µm in diameter. Macrotubular ducts about 4–6 µm
wide, length not measurable, scattered. Microtubular duct scattered all over. Cruciform
pores 4 µm diameter, scattered on submarginal band, and on head, absent on middle of
thorax.
Dorsum: All enlarged setae spine-like, conical, each seta 19–31 µm long, varying in size
and shape, slightly rounded, truncated. Dorsal spinose setae forming sparse rows, about
the same sizes. Enlarged setae in middle part of last segments 19–20 µm long. On the
margin of last abdominal segments 2-5 spines present each 29–31 µm long. Macrotubular
ducts similar to those on venter, sparse, present throughout, 6–8 µm wide, length not
measurable. Microtubular ducts each 4 µm long, scattered throughout. Discodial pores
absent. Anal ring not well seen, 53–54 µm wide and 72–80 µm long, with 8 hair-like setae,
74–92 µm long. Cauda not seen (Redescription is based on type material).
Ecology
Host plant: Calluna vulgaris.
Biology: On the roots of plants. Live together with R. ericae.
Acanthococcidae 421
Rhizococcus cactearum (Leonardi, 1918). (Fig. 147)

Eriococcus cactearum Leonardi, 1918: 206.
Holotype: Female. Italy (Bordighera, Liguria), on Cactaceae, ex vetrino 99 della

Collez. Lectotype female, by subsequent designation Tranfaglia & Esposito, 1985:
115. Deposited in IFSP.
Lit.: Borchsenius, 1949: 355; Danzig 1971a: 821; Hoy, 1963: 78; Kosztarab & Kozár,
1988: 292; Leonar1920: 431; Miller & Gimpel, 1996: 599; Ouvrard & Kozár, 2009:
102; Pellizari & Kozár, 2011: 67; Tranfaglia & Esposito 1985: 115 (Miller et al., 2013).
Rhizococcus cactearum; Borchsenius, 1949: 355. Change of combination.

Eriococcus cactearum; Ferris, 1955: 116. Revived combination. Notes: Eriococcus cactearum has
been considered a synonym of Eriococcus coccineus by Lindinger (1931: 114; 1936: 156).
Zimmerman (1948: 287) thought it possibly to be a synonym of E. coccineus as Ferris
(1955: 116), and Hoy (1962: 58) did. Tranfaglia & Esposito (1985: 115) consider the
species distinct.
Rhizococcus cactearum; Kozár & Walter, 1985: 75. Revived combination.
Acanthococcus cactearum; Miller & Gimpel, 1996: 599. Change of combination.
Rhizococcus cactearum; Köhler, 1998 397. Revived combination.
Eriococcus cactearum; Miller & Gimpel, 2000: 152. Revived combination.
Rhizococcus cactearum; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Adult female elongate-oval.
Mounted female
Body elongate oval, 2.56 mm long, 1.42 mm wide. Antennae 7 segmented, 364 µm long,
each segment covered with a few, hair-like setae (except third segment without setae).
Frontal lobe or tubercle not mentioned. Eyes situated on venter near margin. Anal lobes
strongly developed, each with 3 enlarged setae with pointed apex, each 54.4–58 µm apical
seta 393.7 µm; ventral hair-like subapical seta 29–65.3 µm long.
Venter: Labium wide. Legs well developed; lengths of segments of prothoracic legs;
coxa: 50–60, trochanter: 37.5–55, femur+tibia 181.5–196, tarsus 214–218. Metathoracic
coxae with pores. Tibiae each with 5 setae (median seta present). Enlarged setae, situated
on submargin in 1 or 2 rows. Multilocular pores with 5 loculi, distributed all abdominal
and thoracic segments and a few on head. Macrotubular ducts of one size, present on
scattered on abdomen, thorax and head. Microtubular ducts, and cruciform pores not
mentioned.
Dorsum: Marginal setae blunt or rounded, slightly truncated, wide at the base, long, each
50.0–72.6 µm, forming a marginal row, each segment with 3 setae. Dorsal setae very
short; in a sparse transverse row across each body segment, rows irregular on head.
Macrotubular ducts, scattered throughout dorsum, generally in segmental sparse bands.
Microtubular ducts and cauda not mentioned. Anal ring strongly sclerotized, with on
each inner side, 55–65 µm in diameter, with 8 setae, anal ring situated on margin of
dorsum.
Ecology
Host plant: Ariocarpus trigoni, Astrophytum ornatum, Cereus sp., Echinocactus sellowii, Echinopsis
sp., Mammillaria sp.
Distribution: Italy, Italy (Sicily), Malta, Morocco, Russia.
Biology: On surface of plants.
Rhizococcus cantium (Williams, 1985) (Fig. 148) Combination nova

Eriococcus cantium Williams, 1985a: 363.
Holotype: Female. England: Kent, Bearsted, on Brachypodium sylvaticum, 30.vii.1925,

by E.E. Green. By original designation. Deposited in BMNH. In addition to the
holotype there is one adult female paratype on a slide mounted separately in the
BMNH.
Lit.: Kozár, 2009: 91; Kozár, et al., 2004: 59; Kozár et al., 2013: 56; Ouvrard & Kozár,
2009: 101; Tang & Hao, 1995: 449; Williams, 1985a: 363 (Miller et al., 2013).
Anophococcus cantium; Lagowska & Koteja, 1996: 31. Change of combination.

Acanthococcus cantium; Miller & Gimpel, 1996: 599. Change of combination.
Acanthococcus cantium; Köhler, 1998: 374. Revived combination.
Eriococcus cantium; Miller & Gimpel, 2000: 154. Revived combination.
Acanthococcus cantium; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Mounted female
Body elongate oval, almost parallel sides, 2.6 mm long, 1.0 mm wide. Antennae 7
segmented, 300 µm long. Frontal tubercle present, just anterior to basal antennal segment.
Eyes situated on venter near margin. Anal lobes about twice as long as wide, moderately
sclerotised; each lobe with an apical seta 300 µm long, dorsally with 1 inner and 3 outer
enlarged setae and ventrally with 2 slender setae and slender suranal seta shorter than
anal ring setae.
Acanthococcidae 423
Figure 147. Rhizococcus cactearum (Leonardi, 1918), female. After Tranfaglia & Esposito
Venter: Labium 100 µm long, slightly shorter than clypeolabral shield and basal segment
with 2 pairs of setae. Legs well developed; hind trochanther + femur 250 µm long, hind
tibia 150 µm long, hind tarsus 160 µm long, claw curved 40 µm long, with a denticle near
apex. Hind coxa with minute translucent pores on oter half and hind femur with a small
group on mid-anterior margin. Tibiae each with 5 setae (median seta present), tarsi each
with 5 setae. Derm with normal slender setae in median areas. Enlarged setae, same as
on dorsum, scattered on margins of head, thorax and anterior abdominal segments, but
submarginally a few setae that are more slender but stiff. Macrotubular ducts of two sizes;
a larger type, same size as on dorsum around margins and a narrower type in transverse
rows on abdominal segments, and in median to submarginal areas of thorax and head.
Microtubular ducts in small numbers on margins. Discodial pores of quinquelocular
type, numerous across abdominal segments in median areas of thorax and head and near
spiracles. Cruciform pores present in small numbers in a narrow submarginal zone from
about fifth abdominal segment forward to head.
Dorsum: Enlarged conical setae, poited 35–75 µm long, in bands across the segments,
and with two submedian and one median longitudinal band of spines of larger size. On
seventh segment medially a group of 4-5 present and on sixth segment a smillar group
present but this merging with lateral setae. Macrotubular ducts one size about 25 µm
long, the cup narrower than setal bases of largest setae, fairly evenly distributed across
segments. Microtubular ducts not numerous, each about 4 µm long, with short collar,
ampulla and swollen inner end to tube. Cauda not not prominent, rounded, nodulose
(after Williams (1985a) with modifications).
Ecology
Host plant: Brachypodium sylvaticum.
Distribution: Hungary, Poland, United Kingdom.
Biology: On the leaves of plants.
Comments
This species probably present widely in Russia (Kaliningrad, Siberia, Far East, Sachalin)
under the name of R. greeni, according to Danzig (1985) who found spcimens with four
spines on anal lobes.
Acanthococcidae 425
Figure 148. Rhizhococcus cantium (Williams, 1985), female. After Williams (1985) with
modifications.
Rhizococcus caudatus (Tang & Hao, 1995) (Fig. 149) Combination nova
Eriococcus caudatus Tang & Hao, 1995: 463.
Holotype: Female China (Inner Mongolia), on Artemisia gmelinii, 30.vii.1989. By

original designation. Deposited in Sanxi.
Lit.: Tao, 1999: 32; Wang, 2001: 208 (Miller et al., 2013).
Acanthococcus caudatus; Miller & Gimpel, 1996: 599. Change of combination.

Eriococcus caudatus; Miller & Gimpel, 2000: 157. Revived combination.
Acanthococcus caudatus; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Mounted female
Body elongate oval. 2.2–3.0 mm long, 1.8–2.0 mm wide. Antennae 7 segmented, length
of segments: I: 40, II: 41, III: 52, IV: 40, V: 20, VI: 20 and VII: 35 µm long; each segment
covered with a few, hair-like setae. Frontal lobe present. Eyes situated on venter near
margin. Anal lobes sclerotised and with three dorsal spines, all like slender setae, except
sometimes for the outward one.
Venter: Mouth parts developed. Stylet loop longer than labium. Legs well developed,
medium size tibia and tarsus are equal length, but sometimes the former shorter than the
latter; claw with denticle; tarsal and claw digitules surpassing the claw; and hind coxa with
transparent pores in quantity. Derm with a sparse covering of scattered hair-like setae,
Enlarged setae, situated on submargin in 1 or 2 rows. Quinquelocular pores distributed
all abdominal and thoracic segments and a few on head. Macrotubular ducts of two sizes,
larger one present on scattered on abdomen, thorax and head, smaller one present on
the middle part of abdominal segment only. Microtubular ducts, situated marginal area
of body surface. Cruciform pores present on submarginal zone of head, thorax and first
abdominal segments.
Dorsum: Marginal setae slightly truncated, wide at the base, long, each 30–35 µm, forming
a marginal row, no presence on eighth abdominal segment, the sixth and seventh each
with one transverse row, other segments each with a band, all approximately same size.
Macrotubular ducts of one size, scattered throughout dorsum, generally in segmental
bands. Microtubular ducts with 2 sclerotized areas; scattered over dorsum. Anal ring
sclerotized, with pores in rows, with 7 or 8 setae, anal ring situated on margin of dorsum.
Cauda well developed (after Tang & Hao (1995) with modifications).
Acanthococcidae 427
Figure 149. Rhizococcus caudatus (Tang & Hao, 1995), female. After Tang & Hao (1995)
with modifications.
Ecology
Host plant: Artemisia sp., A. gmelini.
Biology: Unknown.
Rhizococcus cistacearum (Goux, 1936) (Fig. 150) Redescription, Revived

combination
Eriococcus cistacearum Goux, 1936: 344.
Holotype: Female. France (Bouches-du-Rhone, Marseille, La Madrague de

Montredon), on Cistus albidus, by L. Goux, 30. ix. 1935 (MNHN 14449-01).
Paratypes: 35 slides are with the same data as holotype (MNHN 14449-02-36).
Notes: Specimens on 24 slides are from the same place and host plant, but they were
collected in 12.x.1934 (MNHN 14450-01-24), specimens on 4 slides were collected in
20. XII. 1934 (MNHN 14451-01-04), specimens on 8 slides were collected 21.x. 1931
(MNHN 14452-01-08). Deposited in MNHN.
Lit.: Foldi, 2001: 305; 2003: 149; Kozár et al., 2004: 60; Kozár & Walter, 1985: 74;
Köhler, 1998: 374 (Miller et al., 2013).
Nidularia cistacearum; Lindinger, 1943: 223. Change of combination.

Eriococcus cistacearum; Hoy, 1963: 79. Revived combination.
Acanthococcus cistacearum; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus cistacearum; Miller & Gimpel 2000: 166. Revived combination.
Rhizococcus cistacearum; Ouvrard & Kozár, 2009: 102. Change of combination.
Rhizococcus veyrensis; (Goux, 1991). Synonym and combination nova.
Holotype: Female. France (Buches-du-Rhone, Marseille, Marseilleveyre, Mazargues),
on Helianthemum sp. by L. Goux, 05.VI. 1932, MNHN 14528-01 (Holotype), MNHN
14528-02 (paratype), MNHN 14528-03 (paratype, nymph).
Acanthococcus? cistacearum; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female elongate-oval, and red. Ovisac is ovoid, white at time of secretion, but
rapidly turning yellowish cream. Adult male is red.
Mounted female
the length of the segments: I: 24–32, II: 20–31, III: 38–50, IV: 38–46, V: 20–22 and VI:
20–22 µm long and VII: 30.0–32.5 µm long; each segment covered with a few, strong
hair-like setae (except third segment without setae); apical segment with apical seta 34–38
Acanthococcidae 429
Figure 150. Rhizococcus cistacearum (Goux, 1936), female. Original.

µm long; apical segment also with 3 sensory falcate setae, the longest one 25–31 µm long.
Frontal lobe present. Eyes situated on venter near margin. Anal lobes slightly sclerotized
each with two blunted setae along inner margin, and one seta on outer margin, similar in
size to those on dorsum, plus a long apical seta 280–320 µm long.
Venter: Labium 3 segmented, 110–140 µm long, stylet loop long five times longer
than labium, reaching the posterior coxae. Legs well developed; lengths of segments
and digitules of prothoracic legs; coxa: 50–73, trochanter: 45–56, femur: 113–138,
tibia: 89–103, tarsus: 110–128 and claw: 30–34, tarsal digitules: 40–46, claw digitules:
30–35 µm long; lengths of segments and digitules of mesothoracic legs; coxa: 60–66,
trochanter: 50–52, femur: 105–125, tibia: 87–102, tarsus: 110–120 and claw: 33–36, tarsal
digitules: 39–47, claw digitules: 34–39 µm long; lengths of segments and digitules of
metathoracic legs; coxa: 65–78, trochanter 42–52, femur: 122–148, tibia: 104–108, tarsus:
124–135 and claw: 32.5–35.0, tarsal digitules: 42–45, claw digitules: 34–39 µm long, tarsal
digitules slightly knobbed, claw digitules slightly knobbed and longer than claw. Posterior
coxae without spinulae, with high number (more than 20) of large pores. Claw with
denticle. Tibiae each with 5 setae (median seta present), tarsi each with 5 setae. Diameter
of spiracles 31–40 µm; posterior spiracles slightly larger than anterior. Venter with a
small number of scattered, hair-like setae, some of them capitated, and two submarginal
rows of enlarged setae. On several specimens the submarginal spines setose, slightly
knobbed, 30–50 µm long, on others enlarged spine-like, 17–30 µm long. Quinquelocular
pores distributed in sparse bands and rows on all segments of abdomen and thorax, 5
µm in diameter. Macrotubular ducts about 4–6 µm wide and 20–22 µm long, scattered.
Dorsum: Dorsal enlarged setae elongate, conical with acute apices; marginal setae
noticeably longer than other setae on dorsum, with one large lateral seta on margin of
each abdominal segment, each setae 8–52 µm long, with blunted apex, varying in size
and shape enlarged setae scattered, except last three segments. Some other enlarged setae
hair-like, slightly knobbed, 14–17 µm long. On margin of segments setae 36–52 µm long.
and 20–24 µm long. Microtubular ducts with 2 sclerotized areas, 4–6 µm long, scattered
throughout. Discodial pores absent. Anal ring 50–64 µm wide, 60–70 µm long, with
pores and with 8 hair-like setae, each 120–130 µm long. Cauda present (Redescription is
Other stages
Adult winged male, as well as immatures were described by Goux (1936).
Acanthococcidae 431
Ecology
Host plant: Arthrocnemium macrostachyum, Cistus albidus, C. monpeliensis, Helianthemum
polifolium.
Biology: This species has two generations per year. The eggs of the first generation
hatching in May or June and eggs of the second generation appearing at the end of
September. Overwinters as eggs (Goux, 1936).
Comments
Among the studied materials two kind of females were found, in one case the large spines
on submargin of venter are spinose, while on others they are setose and longer. There are
also intermediate forms with these two kinds of spines altogether. All of these variations
were found on the same locality and host plants. Further studies should be done to
understand if they are separate species, or only different morphological or ecological
forms. The presence of setose spines among true spines on dorsum needs further special
studies. This kind variation of characters is very sparse among genera and species of
Acanthococcidae. The detailed study of the materials of Rhizococcus veyrensis (Goux, 1991)
shows that this species belongs to R. cistacearum (Goux, 1936) and synonymised here.
Records form Hungary under name of R. cistacearum belongs to the new described species
An. pannonicus.
Rhizococcus coccineus (Cockerell, 1894), (Fig. 151) Revived combination

Eriococcus coccineus Cockerell, 1894a: 204.
Lectotype: Female. USA (Nebraska, Lincoln), on a cactus in a greenhouse, 23.v.1894,

by Prof. Bruner. Designated by Miller & Miller, 1992: 19-22. Deposited in USNM.
Common name: Cactus eriococcin, cactus mealybug, cactus spine scale, spine
mealybug. woolly cactus scale.
Lit.: Ben-Dov, 1985a: 187; Cockerell, 1894a: 204; Cook & Gullan, 2004: 444; Fernald,
1903: 72; Ferris, 1955: 97; Foldi, 2000: 81; 2001: 305; Gill, 1993: 157; Hardy et al.,
2008: 369; Hodgson & Miller, 2010 99; Hoy, 1963: 80; Kawai, 1980: 128; Koteja,
1974b: 76; Kosztarab & Kozár, 1988: 275; Kozár, 2009: 106; Kozár & Walter, 1985:
74; Köhler, 1998: 374; Kwon & Han, 2003, Masten Milek & Simala, 2008: 105; Miller,
1996: 79; 2005: 491; Miller & Gimpel, 2000: 166; Miller & Miller, 1992: 19; 1993: 23;
Pellizzari & Kozár, 2011: 67; Tang & Hao, 1995: 520; Zahradník, 1968: 11 (Miller et
al., 2013).
Eriococcus coccineus lutescens; Cockerell, 1894a: 204. Synonymy by Ferris, 1955: 116. Notes:
Although Cockerell (1894a) considered this to be a color form of Eriococcus coccineus, it
has been treated as a subspecies or variety several times (Cockerell 1896 and Ferris 1955)
and based on the rules of the International Code of Zoological Nomenclature must be
considered a subspecies.
Dactylopius mammillariae; Lindinger, 1931: 114. Incorrect synonymy.
Eriococcus cactearum; Lindinger, 1931: 114. Incorrect synonymy.
Nidularia coccinea; Lindinger, 1933a: 108. Change of combination requiring emendation
of specific epithet for agreement in gender.
Rhizococcus multispinosus; Lindinger, 1933a: 114. Incorrect synonymy; discovered by Hoy,
1963: 103.
Acanthococcus coccineus; Miller & Miller, 1992: 19-22. Described: female. Illust. Change of
combination.
Rhizococcus coccineus; Kozár, 2009: 106. Change of combination.
Acanthococcus coccineus; Hodgson & Miller, 2010: 99-100. Revived combination.
Eriococcus saboteneus; Kuwana & Tanaka, 1922: 215.
Syntypes: Female. Japan. Unknown type status, type designation unknown. Notes:
Morrison (1952) states that much of Kuwana's material was destroyed in the 1923
earthquake. In Kawai (1980: 128) it was cited under the name E. coccineus. According
to a current communication of Professor Kawai (e-mail from 02.25.2011) he consider
it as synonym of E. coccineus.
Acanthococcus saboteneus; Miller & Gimpel, 1996: 604. Change of combination.
Eriococcus saboteneus; Miller & Gimpel, 2000: 166. Revived combination.
Acanthococcus saboteneus; Kozár, 2009: 93. Revived combination.
Acanthococcidae 433
Description
Unmounted female
Adult female elongate oval. Body usually violet purple with central yellow band,
occasionally crimson red or dull yellow; color changes due to period of development and
perhaps amount or lack of feeding. Ovisac loose white.
Mounted female
Slide-mounted adult female 1.42–3.17 mm long, 0.81–2.3 mm wide. Antennae 6 or 7
segmented, when 6 segmented third segmented longest, when 7 segmented third and
fourth segments subequal. Apical segment with 3 sensory setae. Frontal lobe and
frontal tubercle not seen. Eyes situated on venter near margin. Anal lobes protruding,
moderately sclerotised; each lobe dorsally with 3 enlarged setae (posteromedial setae
largest, anteromedial seta smallest, lateral setae intermediate), with 4–7 microtubular
ducts; each lobe ventrally with 4 slender body setae and 5 or 6 discodial pores (Miller &
Miller, 1992).
Venter: Legs with hind coxae dorsally with 12-29 pores, ventral surface with these pores
absent; hind femora dorsally with 2-6 pores. Tibia with 5 setae; inner apical, tibial setae
robust; tibiae and tarsi nearly equeal in length (hind tibia/tarsus ratio from 0.84–0.96);
claws with inconspicuous denticle near apex. With normal flagellate setae moderate in
length (longest seta on abdominal segment VII from 23–34 µm long, on segment II
from 47–66 µm long), medial setae capitate. Enlarged setae present in two areas: from
abdominal segment VII through head in lateral row; also in sublateral row from abdominal
segment VI or V through prothorax just below anterior spiracles, this row present or
absent. Setae on both areas of two sizes: most with blunt apices; a few, generally smaller
and with rounded apices. Discodial pores with 3 or 5 loculi. Quinquelocular pores most
numerous on posterior abdominal segment, sparse on thorax; trilocular pores usually
present mainly anterior abdomen, thorax and head. Macrotubular ducts of two sizes;
larger ones restricted to lateral areas; smaller size present on medial areas of venter.
Microtubular ducts sparse, present only near lateral margins. Cruciform pores present on
lateral margins from abdominal segments VI or V through head, rarely present in medial
areas (Miller & Miller, 1992).
Dorsum: With enlarged setae of primary two sizes: setae along body margin and in medial
area of thorax large, 53–62 µm long, straight with truncated at apex; others on dorsum
noticeably small (7–10 µm long) and inconspicuous, being no larger in diameter than that
of most ventral body setae; rounded at apex. Normally with 3 large setae along body
margin of each abdominal segment, becoming more numerous head and thorax. Larger
setae in medial area of thorax variable in number; within same population 0-15 present.
Macrotubular ducts, present on over surface. Discodial pores absent. Microtubular ducts,
4–6 µm long, numerous over surface. Anal ring, with 4 or 5 setae, usually 4 (Miller &
Miller, 1992).
Ecology
Host plant: Astrophytum sp., Cereus sp., Cleistocactus sp., Echinocactus sp., Echinocereus sp.,
Echinopsis sp., Hylocereus sp., Mammillaria sp., Neobuxbaumia polylopha, Opuntia sp., O. ficus
barbarica, Pelecyphora sp., Rebutia sp., Rhipsalis sp., Selenicereus sp., Thelocactus bicolor, Wilcoxia
sp.
Distribution: Australia, Brazil, Croatia (former Yugoslavia), former Czechoslovakia,
Egypt, France, Germany, Israel, Italy, Japan, New Zealand, Mexico, South Africa, Spain
(Canary Islands), United Kingdom, USA (Hawaiian Islands), former Yugoslavia. (Notes:
Malumphy, 2006 states that all known outbreaks of R. coccineus in the UK have been
eradicated). This species appears to be a native of Mexico and southern parts of Texas,
New Mexico, Arizona and California. R. coccineus infests cacti, whose transportation
all over the world has spread the species, which is now cosmopolitan in nurseries and
greenhouses worldwide.
Biology: This species apparently has continuous overlapping generations (Gill, 1993).
Early instars remain attached to fleshy parts of cactus; adult female often migrates to
spines of a cactus just before oviposition. Male sacs also normally produced on cactus
spines (Miller & Miller, 1992). This species known in indoor places in Palaearctic Region
only.
Acanthococcidae 435
Figure 151. Rhizococcus coccineus (Cockerell, 1894), female. After Miller & Miller (1992)
with modifications.
Rhizococcus coronillae (Tereznikova, 1977) (Fig. 152) Combination nova

Acanthococcus coronillae Tereznikova, 1977: 573.
Holotype: Female. Ukraine (Krasnodar Kray), on Coronilla varia, 05.vi.1963, E.

Tereznikova. By original designation. Deposited in Kiev. Notes: 1 paratype in ZMAS.
Lit.: Kozár & Walter, 1985: 74; Tereznikova, 1981: 24, 25; Ter-Grigorian, 1983: 879
Eriococcus coronillae; Tang & Hao, 1995: 465. Change of combination.

Acanthococcus coronillae; Köhler, 1998: 375. Revived combination.
Eriococcus coronillae; Miller & Gimpel, 2000: 176. Revived combination.
Acanthococcus coronillae; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Body oval, yellow, or pinkish. Egg sacs white, felted.
Mounted female
Body elongate oval. 2.0–2.2 mm long, 1–1.2 mm wide. Antennae 7 segmented. Frontal
lobe present. Eyes situated on venter near margin. Anal lobes developed, each with 3
enlarged setae.
Venter: Labium well developed, median setae on apex short. Stylet loop short not
reaches median coxae. Legs with normal; metathoracic coxae with large pores on ventral
surface on coxa. Tarsal and claw digitules longer than claw. Tibiae with 5 setae (median
seta present), tarsi with 4 setae. Derm with a sparse covering of scattered hair-like setae.
Enlarged setae, situated on submargin in 1 or 2 rows. Multilocular pores with 5 loculi,
distributed all abdominal and thoracic segments and a few on head. Macrotubular ducts
of two sizes, scattered on abdomen, thorax and head. Microtubular ducts not seen on
venter. Cruciform pores in a submarginal band.
Dorsum: Marginal setae sharp pointed wide at the base, forming a marginal band, setae
much longer than dorsal setae; which are wide and short on abdomen, some much longer
in median part of thorax. Setae arranged in transverse row across body segments, rows
irregular on head. Macrotubular ducts, scattered throughout dorsum, generally in sparse
segmental bands. Microtubular ducts scattered over dorsum. Anal ring with partially
double row of pores. Cauda present (after Tereznikova (1977), with modifications).
Ecology
Host plant: Coronilla varia.
Distribution: Russia, Ukraine.
Acanthococcidae 437
Figure 152. Rhizococcus coronillae (Tereznikova, 1977), female. After Tereznikova (1977)
with modifications.
Rhizococcus crassispinus (Borchsenius, 1949) (Fig. 153) Combination

nova
Acanthococcus crassispinus Borchsenius, 1949: 334.
Lectotype: Female. Armenia (Megri), on roots of Artemisia sp., 25.v.1947, by N.

Borchsenius. Designated by Danzig, 1996: 521. Deposited in ZMAS. Notes: There is
an additional female on the slide with the lectotype and there are 3 other paralectotypes
on 2 slides with same label data as lectotype (Danzig, 1996).
Lit.: Köhler, 1998: 375. Tao, 1999: 32; Ter-Grigorian, 1983: 877; Wang, 2001: 209
Nidularia crassispinus; Lindinger, 1957: 543. Change of combination.

Eriococcus crassispinus; Hoy, 1963: 83. Change of combination.
Acanthococcus crassispinus; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus crassispinus; Tang & Hao, 1995: 452. Revived combination.
Acanthococcus crassispinus; Danzig, 1996: 574. Revived combination.
Eriococcus crassispinus; Miller & Gimpel, 2000: 178. Revived combination.
Acanthococcus crassispinus; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Adult female oval.
Mounted female
Body elongate oval. 1.6 mm long, 1 mm wide. Antennae 7 segmented, length of segments:
II: 37, III: 56, IV: 56, V: 28, VI: 25 and VII: 35 µm long; apical segment also with 3 sensory
falcate setae, segment V and VI with 1 sensory falcate. Frontal lobe present. Eyes situated
on venter near margin. Anal lobes strongly developed, sclerotized, each with 3 enlarged
setae, 28–40 µm long, apical seta 300 µm long and ventral hair-like subapical 100 long.
Venter: Labium strong, median setae on apex of labium short. Legs well developed;
femur 140, tibia 120, µm long; tarsal and claw digitules slightly capitated, longer than
claw. Metathoracic coxae with spinulae and pores on ventral surface on coxa. Tibiae with
5 setae (median seta present), tarsi each with 5 setae. Some enlarged setae, situated on
submargin. Quinquelocular pores distributed all abdominal and thoracic segments, with
groups around spracles, and a few on head. Macrotubular ducts of one size scattered on
abdomen, thorax and head. Microtubular ducts not seen. Cruciform pores in a sparse
submarginal band.
Dorsum: Marginal setae sharp pointed, wide at the base, long, each 28–42 µm long,
forming a wide longitudinal, middorsal band, marginal row, or band absent, dorsal setae
quite equal long; arranged in transverse rows and bands across each body segment.
Macrotubular ducts, each 9 µm wide and 24 µm long, scattered throughout dorsum,
Acanthococcidae 439
Figure 153. Rhizococcus crassispinus (Borchsenius, 1949), female. After Tang & Hao
generally in segmental bands. Microtubular ducts, each 3–5 µm long scattered over
dorsum. Anal ring with partly double rows of pores, with 8 setae, 125 µm long; anal ring
situated on margin of dorsum. Cauda not seen.
Ecology
Host plant: Artemisia sp., A. sphaerocephala, Youngia tenuicaulis.
Distribution: Armenia, China.
Rhizococcus desertus (Matesova, 1957) (Fig. 154) Redescription,

Combination nova
Acanthococcus desertus Matesova, 1957: 168.
Syntype: Female. Kazakhstan (Bank of Ili River near Iliysk), on Camphorosma

monspeliacum roots, 31.05.1952, G. Matesova. By original designation. Deposited in
ZMAS. Notes: Although there is one female on a slide labeled as a holotype, the
entire series must be considered as syntypes because there is no mention of the word
holotype or type in the original description.
Common name: Desert felt scale.
Lit.: Danzig, 1977: 200; 1982: 147; Hoy, 1963: 84; 1988: 279; Kozár, 1980: 67; Kozár
et al., 2013: 56; Kozár & Walter, 1985: 74; Myartseva, 1980: 64; Ouvrard & Kozár,
2009: 101 (Miller et al., 2013).
Eriococcus desertus; Hoy, 1963: 84. Described: female. Change of combination.

Acanthococcus desertus; Kosztarab & Kozár, 1978: 71. Revived combination.
Eriococcus desertus; Tang & Hao, 1995: 452. Revived combination.
Acanthococcus desertus; Köhler, 1998: 375. Revived combination.
Eriococcus desertus; Miller & Gimpel, 2000: 188. Revived combination.
Acanthococcus desertus; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Ovisac smooth, felted, cream-colored. Adult female oval, constricted toward posterior
end, light pink.
Mounted female
Body elongate oval. 2.12 mm long, 0.84 mm wide. Antennae 7 segmented, 220–230 µm
long, length of segments: I: 52.5, II: 35–45, III: 50–80, IV: 32.5, V: 25, VI: 20–22.5 and
VII: 35 µm long; each segment covered with a few, strong hair-like setae; apical segment
Acanthococcidae 441
Figure 154. Rhizococcus desertus (Matesova, 1957), female. Original.

with apical seta 40–45 µm long; apical segment also with 3 sensory falcate setae, each
25–35 µm long and with coeloconic sensilla; segment VI with 1 sensory falcate seta 25.0–
27.5 µm long, segment V with 1 sensory falcate seta 15.0–17.5 µm long. Frontal lobes
and frontal tubercle present. Eyes situated on venter near margin. Anal lobes strongly
developed, each with 3 enlarged setae, each 35–60 µm plus 2 or 3 microtubular ducts on
dorsal surface; apical seta 325 µm; ventral hair-like subapical seta 60 µm long.
Venter: Labium 150–160 µm long, 85–95 µm wide, median setae on apex of labium short
and strong, 10–11 µm long. Stylet loop reaches the median coxae. Legs well developed;
lengths of segments and digitules of prothoracic legs; coxa: 75–100, trochanter: 50–
60, femur: 115–125, tibia: 100–105, tarsus: 105.0–122.5 and claw: 30–35, trochanther
+ femur: 255–280, tibia + tarsus: 205.0–222.5, tarsal digitules 40.0–42.5, claw digitules
27.5 µm long; lengths of segments and digitules of mesothoracic legs; coxa: 85.0–87.5,
trochanter: 50–60, femur: 115–120, tibia: 110–115, tarsus: 110–115 and claw: 30.0–32.5,
trochanther + femur: 255–280, tibia + tarsus: 220–230, tarsal digitules 42.5, claw digitules
25–30 µm long; lengths of segments and digitules of metathoracic legs; coxa: 85–90,
trochanter: 50–60, femur: 120–135, tibia: 110.0–122.5, tarsus: 110–130 and claw: 30–
32.5, trochanther + femur: 275–295, tibia + tarsus: 220–252.5, tarsal digitules 37.5–45.0,
claw digitules 25–30 µm long. Metathoracic coxae each with spinulae on ventral surface
and translucent pores on coxa. Tibiae each with 5 setae (median seta present), tarsi each
with 5 setae. Length of spiracles 50–55 µm; diameter of spiracular peritreme 30 µm,
posterior spiracles slightly larger than anterior. Derm with a sparse covering of scattered
flagellate hair-like setae, each 22.5–70.5 µm long. Enlarged setae, situated on submargin
in 2 or 3 rows, each 17.5–25.0 µm long. Multilocular pores each 4–5 µm in diameter
and with 3–5 loculi, distributed in sparse bands on all abdominal and thoracic segments
and head. Macrotubular ducts of one size, each 5–6 µm wide and 17.5–22.5 µm long,
present on scattered on abdomen, thorax and head. Cruciform pores few, each, 3–4 µm
in diameter, distributed on the submargin of prothorax and head.
Dorsum: Marginal slightly truncated, long, each 22.5–35 µm, forming a marginal row
and not strongly differentiated from dorsal setae; other dorsal setae conical, slightly
truncated, each seta 22.5–32.5 µm long, those on thorax, head and anterior abdominal
segments slightly larger than setae on posterior 2 or 3 abdominal segments; arranged
in transverse rows across each body segment, rows irregular on head. Macrotubular
ducts, each 8–10 µm wide and 15–20 µm long, scattered throughout dorsum, generally in
segmental bands. Microtubular ducts, each 6.0–7.0 µm long scattered over dorsum. Anal
ring strongly sclerotized, with partially double rows of pores, 67.5 µm in diameter, with
6 strong setae, 130–160 µm long; anal ring situated on margin of dorsum. Cauda present
(Redescription is based on type material).
Ecology
Host plant: Artemisia sphaerocephala, Youngia tenuicaulis, Minuartia sp., Camphorosma lessingii,
C. monspeliacum, Kochia prostrata, Scabiosa sp., Punica granatum.
Distribution: Bulgaria, China, Hungary, Kazakhstan, Mongolia, Turkmenistan.
Acanthococcidae 443
Biology: This is a steppe inhabiting xerophilous species. In Kazakhstan, adults appear

in May or June. Egg production (12-86) probably depends on host plant. Young females
and molting last-stage nymphs were found at the end of May. It was at this time that
the flight of the males began (Matesova, 1957). The ant Tetramorium semilaevi has been
associated with this species. This species is found on the bases of leaves and root crowns
of its host (Kosztarab & Kozár, 1988).
Rhizococcus devoniensis Green, 1896 (Fig. 155) Revived combination

Rhizococcus devoniensis Green, 1896: 260.
Lectotype: Female. United Kingdom (England, Budleigh-Salterton), on Erica cinerea,

?.viii.1896, by S. Devon. By subsequent designation Williams, 1985a: 365. Deposited in
BMNH. Notes: In addition to the lectotype there are two adult female paralectotypes
on the same slide in the BMNH.
Common name: Heather scale.
Lit.: Balachowsky, 1937b: 5; Bodenheimer, 1935: 260; Borchsenius, 1949: 337;
Fernald, 1903: 74; Hoy, 1963: 85; Jancke, 1955: 285; Jansen, 2001: 200; Kaydan, et
al., 2007: 90; Koteja, 1974b: 76; Kozár, 1986: 172; Kozár et al., 2013: 56; Kozár &
Walter, 1985: 74; Kozarzhevskaya & Reitzel, 1975: 16; Köhler, 1998: 375; Miller
& Gimpel, 2000: 190; Morrison & Morrison, 1922: 24; Ouvrard & Kozár, 2009:
101; Schmutterer, 1952: 68; Tang & Hao, 1995: 469; Williams, 1973: 83; 1985a: 365;
Zahradník & Rozkosny, 1995: 205 (Miller et al., 2013).
Eriococcus devoniensis; Cockerell, 1897: 588. Change of combination.

Eriococcus ericae; Lindinger, 1911: 357. Incorrect synonymy; discovered by Hoy, 1963: 87.
Acanthococcus devoniensis; Borchsenius, 1949: 337. Change of combination.
Acanthococcus devoniensis; Kosztarab & Kozár, 1988: 277. Revived combination.
Eriococcus devoniensis; Miller & Gimpel, 2000: 190. Revived combination.
Acanthococcus devoniensis; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Adult female is oval and purple. Ovisac strongly convex, creamy-white; up to 2.5 mm
long, 1.8 mm wide. Eggs are oval, pink or pinkish yellow and dusted with white wax
powder.
Mounted female
Body broadly oval. 2.0 mm long, 1.3 mm wide, body nodulose. Antennae 7 segmented,
190–210 µm long. Frontal lobes conspicuous and larger than a basal antennal segment.
Eyes situated on venter near margin. Anal lobes sclerotised, rather wide and pointed,
triangular, about as wide as long at base in older specimens, but conical and rounded
in young specimens; each lobe with an apical setae 180 µm long and dorsally 2 inner
submarginal enlarged setae and 1 outer marginal seta; ventrally each lobe with 2 slender
setae and a suranal seta shorter than anal ring setae.
Venter: Labium 130–150 µm long, usually slightly shorter than clypeolabral shield, basal
segment with 2 pairs of setae. Legs well developed; hind trochanter + femur 180–200
µm long, hind tibia 100–115 µm long, hind tarsus 110–120 µm long, the tibia + tarsus
longer than trochanter + femur; claw stout and curved, 30 µm long, with denticle near
apex. Hind coxa with a few noticeable translucent pores and hind femur with a few on
anterior edge. Derm with normal slender setae in median areas and enlarged setae smilar
to on dorsum, in a marginal zone, on thorax, reaching almost to spiracles. Discodial
pores predominantly quinquelocular, numerous on abdomen and present in median
areas of thorax and around spiracles. Cruciform pores few, in submargins of head and
thorax. Macroducts of 2 sizes; a large type, same as those on dorsum, in a marginal
zone, interspersed with enlarged setae; a narrower type present across median areas of
abdominal segments and in groups in median areas of thorax and head. Microducts few,
on margins only.
Dorsum: Derm with an even distribution parse covering of enlarged setae of various sizes,
28–40 µm long, each truncate, the sides straight or only slightly concave, and tapering
gradually. Macrotubular ducts of one size, about 20 µm long, the cup about same size or
smaller than diameter of setal base of larger setae, fairly numerous. Microtubular ducts
with 2 sclerotized areas, about 4 µm long, with ampulla, inner end of tube swollen, but
without internal collar, opening to exterior with minute orifice, in moderate numbers
over entire surface (Williams, 1985a).
Other stages
First instar described by Schmutterer (1952) as R. devoniensis probably belongs to R. munroi.
Adult male described by Goux (1931) and Jancke (1955).
Ecology
Host plant: Achillea sp., Acroptilon repens, Agropyron repens, Artemisia sp., Calluna vulgaris,
Cichorium intybus, Dianthus sp., Erica sp., E. arborea, E. cinerea, E. multiflora, E. tetralix, Erodium
sp., Ilex sp., Lavandula sp., Salicornia sp., S. fruticosa, Salvia sp., Santolina sp., Taraxacum sp.,
Thymelaea sp., Thymus sp., Ulex europaeus, Vaccinium sp., Veronica multifida.
Distribution: Algeria, Austria, Denmark, France, Germany, Hungary, Ireland, Israel,
Italy, Malta, Netherlands, Poland, Spain, Sweden, Switzerland, Turkey, United Kingdom.
Biology: This species overwinters as eggs and hatches during the first third of May with
adults appearing by the end of June or the beginning of July. Females lay 28-82 eggs in
late July. Infested shoots of Erica are often distorted (Kosztarab & Kozár, 1988).
Acanthococcidae 445
Figure 155. Rhizococcus devoniensis (Green, 1896), female. After Williams (1985) with
modifications.
Rhizococcus echinatus Goux, 1938 (Fig. 156) Redescription, Combination

nova
Eriococcus acutus echinatus Goux, 1938: 337.
Holotype: Female. France (Var, La Seyne, Tamaris-sur-Mer), on Brachypodium

pinnatum, by L. Goux, 26. vii. 1934 (MNHN 14432-01). Paratypes on two slides with
the same data as the holotype (MNHN 14432-02-03). Deposited in MNHN.
Lit.: Foldi, 2001: 305; Hoy, 1963: 68; Miller & Gimpel 2000: 117; Kozár et al., 2013:
56; (Miller et al., 2013).
Acanthococcus echinatus; Ouvrard & Kozár, 2009: 103. Change of combination and status.
Acanthococcus (acutus) echinatus; Kozár, 2009: 92. Change of combination and status.
Description
Unmounted female
Egg sacs oval, white, 4.5 mm long, 1.5 mm wide.
Mounted female
length of segments: I: 36–52, II: 36–41, III: 46–60, IV: 81–100, V: 24–38, VI: 24–31 and
VII: 41–43 µm long; apical setae of antenna broken, on apical segment three sensory
falcate setae are found, the longest 31 µm long. On the preapical segment the falcate
sensory seta 26 µm long. The segments of the antenna are covered with few hair-like
setae. Frontal lobes present. Eyes situated on venter near margin. Anal lobes slightly
sclerotized each with two enlarged setae along inner margin, and one enlarged seta on
outer margin, similar in size to those on dorsum, apical seta 288–320 µm long.
Venter: Labium 3 segmented, 103–116 µm long; stylet loop as long as the labium. Legs
well developed; lengths of segments of prothoracic legs; coxa: 89–92, trochanter: 54–61,
femur: 173–179, tibia: 146–160, tarsus: 141–151 and claw: 31–36 µm long; lengths of
legs; coxa: 108–120, trochanter: 72–78, femur: 185–198, tibia: 180–181, tarsus: 170–173
and claw: 36–38 µm long; tarsal digitules knobbed, 43–46, claw digitules, 35–37 µm
long, slightly knobbed; claw with denticle. Metathoracic coxae without spinulae, with
medium number (less than 20) of translucent pores. Tibiae each with 5 setae (median
seta present). Length of anterior spiracles 36–44 µm; posterior spiracles slightly larger
than anterior. Derm with a small number of scattered, hair-like setae and a marginal
row of enlarged setae. Quinquelocular pores distributed in sparse bands and rows on
all segments of abdomen and thorax, 5 µm in diameter. Macrotubular ducts about 4–7
µm wide and 19–24 µm long, scattered. Cruciform pores 3–5 µm long, scattered in a
submarginal band, absent on middle of thorax.
Acanthococcidae 447
Figure 156. Rhizococcus echinatus (Goux, 1938), female. Original.

Dorsum: Dorsal enlarged setae all spine-like, with pointed apex, very wide and strong,
varying in size and shape. Other enlarged setae forming 3 longitudinal medial bands,
except last two segments; setae in middle part of last segments 24–36 µm long. On margin
of segments 6 or 7 spines present, 64–65 µm long. Diameter of base of median enlarged
setae on penultimate segment 15–18 µm, and on margin 20–28 µm. Macrotubular ducts
similar to those on venter, sparse, present throughout, 6–10 µm wide, and 22–30 µm
long. Microtubular ducts with bifid orifice 4–6 µm long, scattered throughout. Discodial
pores absent. Anal ring 62–70 µm wide, 75–77 µm long, with pores rows and with 8 hair-
like setae, each 113–130 µm long. Cauda absent (Redescription is based on type material).
Ecology
Host plant: Brachypodium pinnatum.
Distribution: France, Hungary.
Biology: One generation in a year, ovisac appears in June.
Comments
According to the drawing of specimens found in Italy concerning the shape and size
of spines it looks similar to R. echinatus (Goux, 1938), however it differs by absence of
cruciform pores around margin, microtubular ducts not shown, or mixed with smaller
sized macrotubular ducts. The identification of Italian specimens needs further study.
Rhizococcus erinaceus (Kiritchenko, 1940) (Fig. 157) Redescription,

Combination nova
Eriococcus erinaceus Kiritchenko, 1940: 128
Lectotype: Female. Ukraine (Kherson District, Kopani, near the Dniepr river), on
Achillea millefolia. By subsequent designation Danzig, 1996: 521. Deposited in ZMAS.
Notes: There also is 1 paralectotype female in ZMAS.
Common name: Woolly felt scale.
Lit.: Babaev, 1980: 57; Borchsenius, 1949: 332; Danzig, 1996: 574; Fetykó, et al., 2010:
295; Kosztarab & Kozár, 1988: 277; Lindinger, 1957: 543; Tereznikova, 1975: 29;
1981: 27.
Acanthococcus erinaceus; Borchsenius, 1949: 332. Change of combination.

Nidularia erinaceus; Lindinger, 1957: 543. Change of combination.
Eriococcus erinaceus; Hoy, 1963: 88. Revived combination.
Acanthococcus erinaceus; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus erinaceus; Tang & Hao, 1995: 448. Revived combination.
Acanthococcus erinaceus; Köhler, 1998: 376. Revived combination.
Acanthococcidae 449
Figure 157. Rhizococcus erinaceus (Kiritchenko, 1940), female. Original.

Eriococcus erinaceus; Miller & Gimpel, 2000: 200. Revived combination.

Acanthococcus erinaceus; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Ovisac pyriform, strongly convex, red, or rusty red, cream-colored, up to 2 mm long, 1
mm wide, female 1.0–1. 3 mm long, 0.5–0.8 mm wide.
Mounted female
Body elongate oval. 1.55 mm long, 0.77 mm wide. Antennae 7 segmented, 230 µm long.
On apical segment three sensory falcate setae are found. The segments of the antenna
are covered with few hair-like setae. Frontal tubercle present. Eyes situated on venter
near margin. Anal lobes slightly sclerotized each with two enlarged setae along inner
margin, and one enlarged seta on outer margin, similar in size to those on dorsum, apical
seta 350, preanal setae 45 µm long.
Venter: Labium 3 segmented, with long hair-like setae, stylet loop reaches middle line
between first and second coxae. Legs well developed; tarsal and claw digitules slightly
knobbed, longer than claw; claw with denticle. All coxae with spinulae and on the
posterior medium number (less than 20) of translucent pores. Tibiae each with 5 setae
(median seta present). Derm with a small number of scattered, hair-like setae and a
marginal row of enlarged setae. Quinquelocular pores distributed in sparse bands and
rows on all segments of abdomen and thorax. Macrotubular ducts one size, scattered
in marginal and submarginal bands and transverse rows. Cruciform pores scattered in a
submarginal band from head to thirth abdominal segment; absent on middle of thorax.
Dorsum: Dorsal enlarged setae all with blunted apex, wide and strong, varying in size and
shape; form 5 transverse bands on prosoma, a narrower transverse band on abdominal
tergites I-VI, a row on tergite VII; setae dense, distance between them normally less than
or equal to length of setae; marginal band of large setae and 3 longitudinal bands in
middle line of dorsum not distinct; setae in middle part of last segments 24–27 µm long.
On margin of segments 3 or 4 enlarged setae present, 36–45 µm long. Macrotubular ducts
sparse, present throughout. Microtubular ducts scattered throughout. Anal ring with
pore rows and with 8 hair-like setae, each 125 µm long. Cauda not seen (Redescription is
Ecology
Host plant: Achillea millefolium.
Distribution: Romania, Ukraine.
Biology: On the upper surface of the leaves. Apparently a rare steppe species.
Acanthococcidae 451
Rhizococcus evinae Kaydan Species nova (Fig. 158)
Holotype: Female. Turkey (Van- Hakkari Road, N: 38°22’248’’, E: 043° 35’176”,

1859 m altitude), on Euphorbia sp., M.B. Kaydan, 06.vi.2009 (KPCT: 4504). Paratypes:
4 adult females, same data as holotype (KPCT: 2237). Deposited in Kaydan’s personal
collection KPCT.
Common name: Euphorbia felt scale.
Description
Unmounted female
Adult females redish; found at rootcrown host plant, felt-like test white.
Mounted female
250–290 µm, length of segments: I: 32.5–57.5, II: 37.5–57.5, III: 92.5–112.5, IV: 25–
27.5, V: 22.5–27.5, and VI: 25–35 µm long, each segment covered with a few, strong
hair-like setae; apical segment with apical seta 47.0–52.5 µm long; apical segment also
with 3 sensory falcate setae, each 20–40 µm long; segment V with 1 sensory falcate seta
32.5–37.5 µm long, segment IV with 1 sensory falcate seta 17.5 µm long. Frontal lobes
absent, frontal tubercle present. Eyes situated on venter near margin. Anal lobes strongly
developed, each with 3 enlarged setae, each 32.5–40.0 µm plus 2 or 3 microtubular ducts
on dorsal surface; apical seta 330–340 µm; ventral hair-like subapical seta 70–95 µm long.
Venter: Labium 155.0–187.5 µm long, 87.5–120 µm wide, median setae on apex of
labium short, 15–20 µm long. Legs well developed; lengths of segments and digitules
of prothoracic legs; coxa: 70–85, trochanter: 55–60, femur: 130–155, tibia: 107.5–120.0,
tarsus: 110–120 and claw: 32.5–35.0, trochanther + femur: 182.5–195.0, tibia + tarsus:
215–235, tarsal digitules 37.5–45, claw digitules 27.5–35.0 µm long; lengths of segments
and digitules of mesothoracic legs; coxa: 70–85, trochanter: 55–60, femur: 125–145,
tibia: 127.5–147.5, tarsus: 110–125 and claw: 30.0–37.5, trochanther + femur: 185–205,
tibia + tarsus: 215–240, tarsal digitules 37.5–42.5, claw digitules 30–35 µm long; lengths
of segments and digitules of metathoracic legs; coxa: 85.0–92.5, trochanter: 52.5–
67.5, femur: 127.5–152.5, tibia: 120.0–147.5, tarsus: 120.0–137.5 and claw: 32.5–37.5,
trochanther + femur: 187.5–210, tibia + tarsus: 245–270, tarsal digitules 35–40, claw
digitules 25.0–32.5 µm long. Tarsal and claw digitules knobbed. Meso- and metathoracic
coxae each with spinulae on ventral surface and translucent pores on metathoracic coxa
and femur. Tibiae each with 5 setae (median seta present), tarsi each with 5 setae. Length
of spiracles 62.5–72.5 µm; diameter of spiracular peritreme 30–35 µm, posterior spiracles
slightly larger than anterior. Derm with a sparse covering of scattered short flagellate
setae, almost spine like, each 12–45 µm long. Enlarged setae, situated on submargin in
2 or 3 rows, each 24.5–40.0 µm long. Multilocular pores each 5.0–7.5 µm in diameter
and with 5 loculi, distributed in sparse bands on all abdominal and thoracic segments
and head. Macrotubular ducts of two sizes, smaller macrotubular ducts each 5.0–7.5 µm
wide and 17.5–25.0 µm long, present on submedian of abdominal segments I-VI, thorax
and head, larger macrotubular ducts each 8.5.0–10.0 µm wide and 22.5–27.5 µm long
scattered elsewhere on abdomen, thorax and head. Microtubular ducts sclerotized, with
oval orifice, present on all venter, 5.0–7.0 µm long; cruciform pores few on thorax only,
each 5.0–6.0 µm in diameter.
Dorsum: Marginal dorsal setae strong, spine-like, blunted, various size, most of them
long, some of them shorter than half size of longer one, each 17.5–65.0 µm, forming
a marginal row and slightly differentiated from dorsal setae; other dorsal setae conical,
spine-like, various size, each seta 12.5–40.0 µm long, those on thorax, head and anterior
abdominal segments larger than setae on median part of posterior 2 or 3 abdominal
segments; arranged in transverse rows across each body segment, rows irregular on head.
Macrotubular ducts, each 7–10 µm wide and 20.0–27.5 µm long, scattered throughout
oval dermal orifice, scattered over dorsum. Anal ring strongly sclerotized, with 21-28
pores on each inner side, 57.5–72.5 µm in diameter, with 8 setae, each 87.5–105 µm long;
anal ring situated on margin of dorsum. Cauda present.
Etymology:
The new species is named in after Evin Polat Akköprü, who collected felt scale samples
for this study.
Ecology
Host plant: Euphorbia sp.
Biology: On the root and root crown of the host plant.
Acanthococcidae 453
Figure 158. Rhizococcus evinae Kaydan sp. n., female.

Rhizococcus festucae (Kuwana & Fukaya, 1914) (Fig. 159) Combination

nova
Eriococcus festucae Kuwana & Fukaya, 1914 (in Kuwana, 1914: 2).
Syntypes: Female. Japan: (Honshu, Nishigahara, Tokyo). Deposited in Ibaraki-ken:

Insect Taxonomy Laboratory, National Institute of Agricultural Environmental
Sciences, Kannon-dai, Yatabe, Tsukuba-shi. Notes: Eriococcus festucae Kuwana (1914)
is a junior primary homonym of Eriococcus festucae Targioni Tozzetti (1869). The
latter species is now considered to be a member of the genus Eriopeltis in the family
Coccidae (Lindinger, 1935).
Lit.: Hoy, 1963: 89; Kawai, 1980: 131; Kozár & Walter, 1985: 74; Kuwana, 1917: 138;
Lindinger, 1932: 201; 1933a: 108; Miller & Gimpel, 1996: 600; Signoret, 1869: 853;
Tereznikova, 1959d: 179 (Miller et al., 2013).
Eriococcus festucarum; Lindinger, 1932: 201. Replacement name for Eriococcus festucae
Kuwana & Fukaya 1914.
Nidularia festucarum; Lindinger, 1933a: 108. Change of combination.
Acanthococcus festucarum; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus festucae; Tang & Hao, 1995: 451. Revived combination.
Acanthococcus festucarum; Köhler, 1998: 376. Revived combination.
Eriococcus festucarum; Miller & Gimpel, 2000: 207. Revived combination.
Acanthococcus (Rhizococcus?) festucarum; Kozár, 2009: 92. Change of combination.
Description
Unmounted female
Female sac is closely felted and tough, pale tan or white in color, with many transverse
ridges, caudal extremity with a small openings. Length 3.5 mm width 1.3 mm. Adult
female is elongate, pale yellow and turns pink when treated with KOH (Kuwana, 1914).
Mounted female
Body elongate oval, 2.0 mm long, 1.1 mm wide. Antennae 6, or seven segmented, with
long hair-like setae. Anal lobes normal, with one long enlarged setae and two short.
Venter: Labium small, rostral setae short. Legs well developed; tarsus slightly longer
than tibia Tarsal and claw digitules knobbed, slightly longer than claw.
Dorsum: Marginal dorsal enlarged setae very strong, conical spine-like, pointed, each
abdominal segment with 3 or 4 enlarged setae, other dorsal enlarged setae much smaller.
Anal ring with 4 pairs setae (Kuwana, 1914).
Acanthococcidae 455
Ecology
Host plant: Festuca parvigluma.
Biology: On the leaves. Eggs are yellow in color. Female collected in June.
Comments
According to Authors the species near to R. greeni and An. insignis, but differs by
arrangement of spines of abdominal segments. According to the presence of enlarged
setae on dorsal segments, shown by Kawai (1980) the similarity with An. insignis can be
excluded.
Figure 159. Rhizococcus festucae (Kuwana & Fukaya, 1914), female. After Kawai (1980)
with modifications.
Rhizococcus gassinus ( Goux, 1992). (Fig. 160) Redescription, Combination

nova
Eriococcus gassinus Goux, 1992: 41.
Holotype: Female. France (Var, Gassin), on Lotus cystisoides, 06.vi.1933, by L. Goux.

By original designation. Deposited in MNHN.
Lit.: Foldi, 2001: 305; Ouvrard & Kozár, 2009 : 102-118 (Miller et al., 2013).
Acanthococcus gassinus; Miller & Gimpel, 1996: 601. Change of combination.

Eriococcus gassinus; Miller & Gimpel, 2000: 212. Revived combination.
Acanthococcus gassinus; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Body oval, 2.3 mm long, 1.4 mm wide.
Mounted female
segments: I: 42, II: 34, III: 58, IV: 60, V: 21, VI: 20 and VII: 33 µm long, each segment
covered with a few, strong hair-like setae; apical segment with apical seta 46 µm long;
apical segment also with 3 sensory falcate setae, longest 27 µm long; on the preapical
segment the falcate sensory seta 32 µm long. Frontal lobes present. Eyes situated on
venter near margin. Anal lobes slightly sclerotized each with two spinose setae along
inner margin, and one spinose seta on outer margin, similar in size to those on dorsal
margin, apical seta 280 µm long.
Venter: Labium 3 segmented 120 µm long; stylet loop almost reaching the coxae
of median legs. Legs well developed; lengths of segments prothoracic legs; coxa: 60,
trochanter: 40, femur: 145, tibia: 121, tarsus: 115 and claw: 29 µm long; lengths of
segments of mesothoracic legs; coxa: 70, trochanter: 44, femur: 120, tibia: 128, tarsus:
122 and claw: 35; lengths of segments of metathoracic legs; coxa: 65, trochanter: 40,
femur: 138, tibia: 130, tarsus: 132 and claw 35 µm long, tarsal digitules knobbed 50, claw
digitules 30 µm long, slightly knobbed. Meso- and metathoracic coxae each with spinulae
on ventral surface and translucent pores on metathoracic coxa and femur. Tibiae each
with 5 setae (median seta present), tarsi each with 5 setae. Claw with denticle. Length of
spiracles 38 µm; posterior spiracles slightly larger than anterior ones. Derm with a small
number of scattered, hair-like setae and with about 5 group of enlarged setae including
5-10 spines on submargin and in addition farther inside a submarginal rows of spines.
Multilocular pores generally quinqueloculars; distributed in sparse bands and rows on
all segments of abdomen and thorax, 5 µm in diameter. Macrotubular ducts about 6
µm wide and 17 µm long scattered. Microtubular duct present in a submarginal band.
Cruciform pores 5 µm long, very few scattered on submarginal band, and on middle of head.
Acanthococcidae 457
Figure 160. Rhizococcus gassinus (Goux, 1992), female. Original.

Dorsum: Enlarged setae on margin 30–60 µm long, varying in size and shape, blunted.
Other dorsal enlarged setae forming sparse bands, except last three segments, where only
a sparse row present. Setae in middle part of last segments very short, 10–14 µm long,
on thorax they are somewhat longer, 25–40 µm. On margin of abdominal segments
2 or 3 spines present. Macrotubular ducts similar to those on venter, sparse, present
throughout, 7–8 µm wide and 18–22 µm long. Microtubular ducts apparently with 2
sclerotized areas, with bifid orifice, each 4 µm long, scattered throughout. Anal ring well
seen, 54 µm wide and 82 µm long, with 8 hair-like setae, 110 µm long. Cauda present, flat
Ecology
Host plant: Lotus cystisoides.
Biology: Unknown.
Rhizococcus gavrilovi Kozár & Kaydan, Species nova (Fig. 161)
Holotype: Turkmenistan, Artemisia sp., x. 1988, G. Harchenko with one paratype on

the same slide (45=69). Paratypes: 3 females on two slides, Turkmenistan, Artemisia
sp. Kara Kala ush., Shovlan Baba 23. IX. 1976, C. Myartzeva 2-79.
Description
Unmounted female
Unknown.
Mounted female
Body elongate oval, 2.8–3.2 mm long, 1.8–2.2 mm wide. Antennae 7 segmented, the size
of the segments: I: 36–43, II: 26–34, III: 43–50, IV: 40–46, V: 19–-26, VI: 19–21 and
VII: 36–38 µm long; each segment covered with a few, strong hair-like setae (except third
and fifth segments); apical segment 36–38 µm long with apical seta 38–46 µm long and
3 sensory falcate setae, each 24–29 µm long; falcate sensory setae on preapical segments
12–29 µm long. Frontal lobe present, frontal tubercle 5 µm wide. Eyes situated on venter
near margin. Anal lobes well developed, sclerotized, with 4 blunt, conical enlarged setae,
ventrally each with two flagellate setae; anal lobe with 204–216 µm long apical seta.
Venter: Labium 3 segmented, 127–139 µm long; basal segment only 1 pairs of setae
seen, apical segment with 6 pairs of hair-like setae, stylet loop reach the median legs.
Legs well developed; lengths of segments and digitules of prothoracic legs; coxa: 65–
80, trochanter: 55–72, femur: 134–144, tibia: 108–120, tarsus: 108–122, claw: 48, tarsal
digitules: 55–58 µm, claw digitules 17 µm long; lengths of segments and digitules of
Acanthococcidae 459
Figure 161. Rhizococcus gavrilovi Kozár & Kaydan sp. n., female.
tarsus: 115–120, claw: 41–48, tarsal digitules: 55–57 µm, claw digitules: 14–17 µm long;
femur: 148–180, tibia: 132, tarsus: 128–137, claw: 48–53, tarsal digitules 53–62, claw
digitules 14–19 µm long, tarsal slightly knobbed; claw digitules spine-like, shorter than
claw. Meso- and metathoracic coxae with spinulae on anterior surface; metathoracic coxae
and femur also with small pores on both anterior and posterior surfaces. Claw with a
denticle. Legs with a few hair-like setae, and with one sensory pore on tarsus; tibia with 5
setae. Multilocular pores with 3 or 5 loculi, mostly quinquelocular each 6 µm in diameter,
3-locular pores 5 µm in diameter; distributed in transverse bands across abdominal
sternites, in sparse rows on thoracic segments and head region. Macrotubular ducts each
5–7 µm wide and 18–24 µm long, scattered throughout the venter. Microtubular ducts
sparse marginally and submarginally. A few cruciform pores on head and in submarginal
area of thoracic segments. On submedian venter a few hair-like setae of 3 kinds, some
of them are capitated; submarginally a few short hair-like setae. Blunt conical spines in
a wide band on margin, each 27–34 µm long. A ventral cauda like structure also seen, it
was not studied by authors earlier. Suranal setae hair-like.
Dorsum: Enlarged setae blunt, conical; longer setae straight or slightly curved, each
48–53 µm long sparse bands on thoracic and abdominal segments, and groups on
head; shorter setae more numerous, each about 27–34 µm long, forming wide bands
on thoracic and abdominal segments, and groups on the head region, but absent from
eighth abdominal segment. More than thirty dorsal setae on seventh abdominal segment.
Macrotubular ducts each 5–7 µm wide and 18–24 µm long, scattered among enlarged
setae. Microtubular ducts scattered, each 5 µm long. Anal ring situated on margin of
venter; oval, sclerotized, 74–86 µm wide, 77–91 µm long, with one row of pores and with
8 setae each 134–156 µm long. Cauda with serrated distal margin.
Etymology
The species named after Dr. Ilya Gavrilov-Zimin, acknowledging his help for us loaning
the specimens and to clarify the situation of this species.
Ecology
Biology: Unknown.
Comments
The species differs from other members of the genus by four spines on the anal lobes
and spinelike claw digitules shorter then claw. These specimens were under the name of
R. turkmenicus (Archangelskayae) in ZMAS, but these specimens are not related with that
species. See remarks under R. turkmenicus.
Acanthococcidae 461
Rhizococcus gnidii Signoret, 1875 (Figs. 162, 163) Revived combination

Rhizococcus gnidii Signoret, 1875: 37.
Type material: Female. France (Estrelle Mountains, near Cannes), on Daphne gnidium.
Unknown type status. Described: female. Synonymy by Williams, 1985a: 382. Notes:
Matile-Ferrero & Danzig visited the Naturhistorisches Museum, Vienna in June
of 1997 and were unable to locate type material, but did find specimens labeled as
follows: "Cannes/Gall. Mer./Daphne gnidium/gnidii det. Signoret/n°4/coll. Nat.
Mus. Wien/Rhizococcus gnidii Sign." Since this label does not contain any indication
that it was collected in the Estrelle Mountains which is part of the type locality, this
material is probably not part of the type series and Var, Agay, on D. gnidium, 4.v.1908
(P. Marchal). From that time some more notes from Danielle Matile-Ferrero: Paul
Marchal collected E. gnidii on the West part of the corniche de l'Esterel (massif de
l'Esterel or Esterel), close to the locality of Agay (departement of Var) where he
had a house. Victor Signoret collected E. gnidii on the East part of the corniche de l'
Esterel (massif de l'Esterel or Esterel) close to the locality of Cannes (department of
Alpes-Maritimes) where he had a house. The material in Wien is probably part of the
type material collected by Signoret. The collection by Marchal is from the type area.
Daphne gnidium is everywhere in the massif de l'Esterel.
Lit.: Borchsenius, 1948: 501; Danzig, 1975: 62; Dziedzicka & Koteja, 1971: 558;
Foldi, 2000: 81; Fernald, 1903: 79; Ferris, 1957b: 85; Hoy, 1963: 92, 120; Koteja &
Zak-Ogaza, 1981: 501; Kozár et al., 2013: 56; Kozár & Walter, 1985: 74; Köhler, 1998:
384; Marcha1, 1909: 198; Miller & Gimpel, 2000: 360; Ouvrard & Kozár, 2009: 102;
Tang & Hao, 1995: 519, 541, 653; Williams, 1985a: 382 (Miller et al., 2013).
Eriococcus gnidii; Lindinger, 1931: 43. Change of combination.

Nidularia gnidii; Lindinger, 1933: 166. Change of combination.
Rhizococcus gnidii; Borchsenius, 1949: 351. Revived combination.
Eriococcus gnidii; Ferris, 1955: 94. Revived combination.
Rhizococcus gnidii; Danzig, 1962a: 839. Revived combination.
Eriococcus gnidii; Williams, 1985a: 382. Revived combination.
Acanthococcus gnidii; Kozár, 1988: 298. Revived combination.
Acanthococcus? gnidii; Kozár, 2009: 92. Change of combination.
Description
Unmounted female
Body oval, 1 mm long, 0.5–1 mm wide, ovisac white.
Mounted female
Body elongate oval, 2.4 mm long, 1.6 mm wide. Antennae 7 segmented, 260 µm, length
of segments: I: 50–53, II: 29–31, III: 50–52, IV: 38–41, V: 14–21, VI: 14–16 and VII:
28–33 µm long, each segment covered with a few, strong hair-like setae; apical segment
with apical seta 48 µm long; apical segment also with 3 sensory falcate setae, longest 37
falcate seta 17 µm long. Frontal tubercle present. Eyes situated on venter near margin.
Anal lobes strongly developed, each with 3 enlarged setae, 30–61 µm plus on dorsal
surface; apical seta 270–285 µm; ventral hair-like subapical seta 110 µm long.
Venter: Labium 3 segmented, 130–139 µm long, stylet loop reach, between meta and
meso coxa. Legs well developed; lengths of segments of prothoracic legs; coxa: 55–60,
trochanter: 53, femur: 140–150, tibia: 95–103, tarsus: 100 and claw: 40 µm long; lengths
of segments of mesothoracic legs: coxa: 55–70, trochanter: 50, femur: 127–130, tibia:
legs: coxa: 68–72, trochanter: 55, femur: 110–132, tibia: 112–118, tarsus: 119–120 and
claw: 35–41, tarsal digitules: 46–49, claw digitules: 35–41 µm long. Tarsal and claw
digitules knobbed. Meso- and metathoracic coxae each with spinulae on ventral surface
and translucent pores on metathoracic coxa and femur. Tibiae each with 5 setae (median
seta present), tarsi each with 5 setae. Length of spiracles 22–28 µm; posterior spiracles
slightly larger than anterior. Derm with a sparse covering of scattered flagellate setae.
Enlarged setae, situated on submargin in 2 rows. Multilocular pores each 5.0–6.0 µm
in diameter and with 3, 5 and 7 loculi, distributed in sparse bands on all abdominal
and thoracic segments and head. Macrotubular ducts of one size, smaller macrotubular
ducts each 5.0–7.5 µm wide and 17.5–25.0 µm long, present on submedian of abdominal
segments I-VI, larger macrotubular ducts, 6–7 µm wide and 25 µm long scattered in
submarginal band on abdomen, thorax and head. Microtubular ducts sclerotized, with
oval orifice, present on submarginal band. Cruciform pores 6 µm long, very few scattered
on submarginal band, and on middle of head.
Dorsum: Marginal enlarged setae spine-like, slightly blunted, various size, most of
them long, some of them shorter than half size of longer one, 28–58 µm, forming a
marginal row and slightly differentiated from dorsal setae; other dorsal enlarged setae
conical, spine-like, various size, each seta 12.5–42.0 µm long, those on thorax, head and
anterior abdominal segments larger than setae on median part of posterior 2 abdominal
segments; arranged in transverse rows across each body segment, rows irregular on
head. Macrotubular ducts, each 6–10 µm wide and 25–28 µm long, scattered throughout
dorsum, generally in segmental bands. Microtubular ducts, each 6–7 µm long with oval
dermal orifice, scattered over dorsum. Anal ring strongly sclerotized, with 8 setae, each
130–140 µm long; anal ring situated on margin of dorsum. Cauda present (Redescription
is based on material from MNHM, Paris and Naturhistorisches Museum, Vienna
collection).
Ecology
Host plant: Daphne gnidium, Seseli sp., Galium sp., Veronica sp., Carex sp., Poaceae, Koeleria
marchantha, Thymus glabrescens.
Acanthococcidae 463
Figure 162. Rhizococcus gnidii Signoret, 1875, female. Original.

Comments
From the curtesy of Daniele Matile-Ferrero we have possibility to add the excellent, very
detailed original drawing (Fig. 163) made by M. V. Signoret. It could be seen, that his
drawing quite well fit to our drawing. It covers most of the characters which are in use
to day and represent the genus Rhizococcus. The only differences what could be seen is
the number of enlarged setae on the margin of segments, he shown only two, but some
could be presented on ventral side of margin.
In the revision of slides in the collection of PPI, specimens from Budaörs, Piliscsaba,
Nagykovácsi, Tatabánya, Isaszeg (Hungary), appeared to be R. gnidii, which are a new
record for the this country.
Rhizococcus greeni (Newstead, 1898) (Fig. 164) Combination nova

Eriococcus greeni Newstead, 1898: 96.
Lectotype: Female. United Kingdom (England, Devon, Budleigh Salterton),

20.ix.1896, by E.Green. By subsequent designation Williams, 1985a: 367-370.
Deposited in BMNH.
Common name: Newstead's felt scale.
Lit.: Danzig, 1980: 212; Fernald, 1903: 75; Fetykó, et al., 2010: 295; Foldi, 2001: 305;
Gertsson, 2001: 125; Kaydan et al., 2005: 399; Kosztarab & Kozár, 1988: 281; Koteja,
1971: 322; Koteja & Zak-Ogaza, 1983: 476; Kozár et al., 2013: 56; Kozár & Walter,
1985: 74; Köhler, 1998: 377; Milonas, et al., 2008: 143-147; Ouvrard & Kozár, 2009:
102; Schmutterer, 1955: 160; Tang & Hao 1995: 449, 645; Tereznikova, 1981: 27;
Williams, 1985a: 367 (Miller et al., 2013).
Acanthococcus greeni; Borchsenius, 1949: 340. Change of combination.

Eriococcus greeni; Lindinger (1912) incorrectly considered to be a junior synonym of An.
insignis (Williams, 1985a).
Eriococcus greeni; Hoy, 1963: 93. Revived combination.
Anophococcus greeni; Lagowska and Koteja, 1996: 31. Change of combination.
Acanthococcus greeni; Kosztarab & Kozár, 1978: 72. Revived combination.
Eriococcus greeni; Miller & Gimpel, 2000: 222. Revived combination.
Acanthococcus greeni; Kozár, 2009: 92. Revived combination.
Eriococcus variabilis; Goux, 1948: 69. Incorrect synonymy. Notes: Goux (1948) considered
E. variabilis to be a junior synonym of E. greeni. However, in Goux (1948), E. variabilis
was regarded as a separate and distinct species.
Acanthococcus (Eriococcus) sachalinensis; (Siraiwa, 1939); Danzig (1986) states that E.
sachalinensis is probably a synonym of E. greeni.
Eriococcus brucius Goux, 1995. Synonym nova.
Holotype: France: Savoie, La Bridorie, by L. Goux, on Gramineae, 10. X. 1932.
(MNHN 14442). Notes: According to the study of the holotype, this species belongs
Acanthococcidae 465
Figure 163. Rhizococcus gnidii Signoret, 1875, female. Original drawing of Signoret.
to R. greeni (Newstead, 1898), and considered here as synonym nova. In contrary to

the original drawing one antennae has 6, the other 7 segments.
Acanthococcus brucius; Kozár 2009: 91. Combination nova.
Description
Unmounted female
Ovisac is elongate-oval, tough, coarse, felted wax and creamy white, 3.5–5 mm long,
1–1.7 mm wide. Adult female is elongate, white or yellowish in color, 2.5–3.4 mm long.
Mounted female
Body elongate oval, 3.0 mm long, 1.6 mm wide, body nodulose. Antennae 6 or 7
segmented, 220–300 µm. Frontal tubercle present. Eyes situated on venter near margin.
Anal lobes about twice as wide, conical, tending to be pointed, moderately sclerotised;
with an apical seta 270–350 µm long and dorsally with 1 outer and 2 inner enlarged setae
and ventrally with 2 slender setae and a slender suranal seta 69 µm long much shorter
than anal ring setae.
Venter: Labium 3 segmented, 110–120 µm long, shorter than clypeolabral shield, stylet
loop as long as labium. Legs well developed; hind leg 680–740 µm long; hind trochanter
and femur: 240–270, hind tibia: 150–270, hind tarsus: 150–170 µm long, tibia + tarsus
always longer than trochanter + femur; claw curved, 35 µm long, with a minute denticle
near apex; hind coxa with translucent pores on outer half and hind femur with a small
group on mid-anterior edge. Tibiae with 5 setae (median seta present). Derm with with
normal slender setae in median areas, marginally with a few enlarged setae smilar to
dorsal setae, 12–20 µm long, submarginally with more slender but stiff setae. Macroducts
of two sizes; a large type same on dorsum, in small numbers around margins; a narrower
type in bands across abdominal segments, in median areas of thorax and head, extending
in submargins almost to marginal macroducts. Micruducts not numerous, in marginal
zone from anterior abdominal segments to head. Quinquelocular pores numerous on
abdomen, in median areas of thorax and around spiracles. Cruciform pores few, in
submarginal areas only of head, thorax and first abdominal segments.
Dorsum: Derm with pointed enlarged conical setae of two sizes, 24–70 µm long, large
size forming 3 longitudinal lines on each side of body, small size scattered over surface,
arrangement of the setae not regular but always some of the longest in the marginal
groups; on the mid dorsum of seventh abdominal segment, there is usually a group of 2
such setae, rarely 3, and smilar group is present on the mid dorsum of sixth abdominal
segment. Macrotubular ducts in a regular distribution, of one type about 25 µm long,
the cup smaller than diameter of setal base of largest seta; tapering gradually, the orifice
usually surrounded by a pale oval area. Microduct about 4 µm long, with ampulla, with
2 sclerotized areas swollen inner end to tube and internal collar, evenly distributed. Anal
ring with 8 setae, 120-140 µm long; anal ring situated on margin of dorsum. Cauda
present (after Williams (1985a) with modifications).
Acanthococcidae 467
Figure 164. Rhizococcus greeni (Newstead, 1898), female. After Williams (1985) with
modifications.
Ecology
Host plant: Aegilops geniculata, Agropyron intermedium, A. repens, Ammophila arenaria,
Brachypodium sp., Calamagrostis brachytricha, C. epigeios, C. canadensis var. langsdorffii, Carex sp.,
Chrysopogon gryllus, Corynephorus canescens, Dactylis sp., Deschampsia sp., Festuca sp., F. ovina,
F. pratensis, Koeleria macrantha, Lolium sp., L. perenne, Melica sp., M. turczaninowiana, Nardus
stricta, Phleum sp., Poa sp., Stipa pennata, Potentilla sp.
Distribution: Armenia, Canada, France, Greece, Hungary, Italy, Kazakhstan, Moldova,
Netherlands, Poland, Romania, Russia, Spain, Sweden, Switzerland, Turkey, Ukraine,
United Kingdom (Scotland).
Biology: It generally feeds on the leaves of poaceous plants. Eggs of this species are
laid in July and August. This species is Nearctic and a common Trans-Palearctic boreal
species.
Acanthococcidae 469
Rhizococcus guesinus (Goux, 1992) Combination nova

Eriococcus guesinus Goux, 1992: 41.
Holotype: Female. France (Hautes-Pyrénées, Artigues), on undetermined Gramineae,

12.viii.1953, by L. Goux. By original designation. Deposited in MNHN.
Lit.: Foldi, 2001: 305; Kozár, 2009: 92; Miller & Gimpel, 2000: 224 (Miller et al.,
2013).
Acanthococcus guesinus; Miller & Gimpel, 1996: 601. Change of combination.
Description
Unmounted female
Body elongate oval, 3.0 mm long, 1.2 mm wide.
Mounted female
Body elongate oval. Antennae 6 or 7 segmented. Anal lobes as long as wide, conical; with
3 enlarged setae 50–70 µm long and with a slender suranal seta.
Venter: Labium 3 segmented. Legs well developed; posterior legs larger than others.
hind coxa with long spinulae, tibiae with 5 setae (median seta present), claw with denticle,
tarsal and claw digitules capitated, longer than claw. Derm with with normal slender setae
in median areas. Macroducts 6.5 µm in diameter. Microducts absent. Quinquelocular and
trilocular pores absent.
Dorsum: Derm with pointed enlarged conical setae various sizes, up to 80 µm long,
enlarges setae form sparse longitudinal bands on mid-dorsum and on margin. On
margin of abdominal segments 3-5 spines present. Macrotubular ducts 8 µm in diameter.
Microduct not mentioned. Anal ring with 8 setae, situated on margin. Cauda not seen
(after Goux, (1992) with madifications).
Ecology
Biology: Unknown.
Comments
According to hand written remark of Goux, he was unable to find the holotype.
According to the description and drawing, this species is considered as unrecognisable
at this time. There is a typographic error in original publication. The legend of the
Planche I belongs to A. puymorensis and the drawings of A. puymorensis and A. guesinus
were replaced. According to original description and drawing this species is similar to
A. greeni (Newstead, 1898). The only difference what could be seen is the presence of
spinulae on posterior coxae, instead of pores. There is a hope to find the original slide in
Goux collection, a female specimen without right anterior leg. Goux (1992) clearly states
the absence of quinquelocular pores and microtubular ducts, which also needs attention,
because they are usually present in other species, not only in female, but even in larval
stages, too. May be he had a specimen in intermedian molting stage. For these reason we
could not include this species neither in key nor drawing.
Rhizococcus hassanicus (Danzig, 1975) (Fig. 165) Combination nova

Acanthococcus hassanicus Danzig, 1975a: 50.
Holotype: Female. Russia (Southern Primor'ye, Khasanskiy district, Kedrovaya

pad), on undetermined Poaceae, 31.vii.1961, by E. Danzig. By original designation.
Deposited in ZMAS.
Lit.: Danzig, 1980: 206; Kozár & Walter, 1985: 74; Tang & Hao, 1995: 450 (Miller et
al., 2013).
Eriococcus hassanicus; Tang & Hao, 1995: 472-473. Change of combination.

Acanthococcus hassanicus; Köhler, 1998: 377. Revived combination.
Eriococcus hassanicus; Miller & Gimpel, 2000: 226. Revived combination.
Acanthococcus hassanicus; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Adult female is oval. Ovisac is cream colored, smooth.
Mounted female
Body elongate oval, 3.0 mm long. Antennae 6 segmented. Frontal tubercle not seen.
Eyes situated on venter near margin. Anal lobes about twice as wide, conical, moderately
sclerotised; with an apical seta and dorsally with 1 outer and 2 inner enlarged setae and
ventrally with 2 slender setae, much longer than anal ring setae and a slender suranal seta.
Venter: Labium 3 segmented. Legs well developed, narrow, hind coxa with spinulae and
translucent pores. Derm with normal slender setae in median areas, marginally with a few
enlarged setae smilar to dorsal setae. Macroducts of one size; in small numbers around
margins and scattered across abdominal segments. Micruducts absent. Quinquelocular
pores numerous on abdomen, in median areas of thorax scattered and in groups around
spiracles. Cruciform pores few, in submarginal areas only of head, thorax. Anal lobes
apparently with 4 enlarged setae (Danzig, 1986).
Dorsum: Derm with pointed enlarged conical setae various sizes, pointed or blunted,
all approximately same size on thorax and head; arrangement of the setae not regular
in rows and bands, on margin 4 or 5 longer enlarged setae present. Macrotubular ducts
Acanthococcidae 471
Figure 165. Rhizococcus hassanicus (Danzig, 1975), female. After Danzig (1985) with
modifications.
in regular distribution. Microduct with ampulla, swollen inner end to tube and internal
collar, evenly distributed. Anal ring with 8 setae, as long as diameter of ring; anal ring
situated on margin of dorsum. Cauda present (after Danzig, 1975, with modifications
Ecology
Host plant: Poaceae
Acanthococcidae 473
Rhizococcus helichrysi (Goux, 1936) (Fig. 166) Redescription, Combination

nova
Eriococcus helichrysi Goux, 1936: 294.
Holotype: Female. France (Marseille), on Helichrysum stoechas, by L. Goux. By original

designation. Paratypes: one paratype on the same slide as holotype (MNHN 14475-
01). Other slides MNHN 14476-01-04, with the same data as holotype, except data
of collection: 16. v. 1934. Two slides MNHN 14477-01-02, with the same data as
holotype, except data of collection: 1. VI. 1934. Five slides MNHN 14478-01-05,
with the same data as holotype, except data of collection: 07. viii. 1934.
Lit.: Foldi, 2001: 305; 2002: 245; 2003: 149; Köhler, 1998: 377; Ouvrard & Kozár,
2009: 102 (Miller et al., 2013).
Nidularia helichrysi; Lindinger, 1943: 223. Change of combination.

Eriococcus helichrysi; Hoy, 1963: 94. Revived combination.
Acanthococcus helichrysi; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus helichrysi; Miller & Gimpel, 2000: 227. Revived combination.
Acanthococcus helichrysi; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Sac is irregular in form, white. Adult female is large and olive colored.
Mounted female
Body elongate oval, 1.57–2.20 mm long and 1.29–1.76 mm wide. Antennae 6 segmented,
length of segments: I: 25–40, II: 20–30, III: 44–70, IV: 14–18, V: 13–18 and VI: 26–30
µm long, each segment covered with a few, hair-like setae; apical segment with apical
seta 31–46 µm long; apical segment also with 3 sensory falcate setae, longest 18–21 µm
long; on the preapical segment the falcate sensory seta 13–20 µm long. Frontal tubercle
present. Eyes situated on venter near margin. Anal lobes slightly sclerotized, each with
two spinose setae along inner margin, and one spinose seta on outer margin, two slender
setae on venter much longer to those on dorsum, plus a long apical seta 160–250 µm
long.
Venter: Labium 3 segmented 80–105 µm long; stylet loop three times longer than labium.
Legs well developed; lengths of segments prothoracic legs; coxa: 34–50, trochanter:
30–46, femur: 60–96, tibia: 40–79, tarsus: 78–95and claw: 26–34 µm long; lengths of
54–79, tarsus: 77–93 and claw: 26–31; lengths of segments of metathoracic legs; coxa:
30–60, trochanter: 25–43, femur: 68–101, tibia: 58–89, tarsus: 96–106 µm long, tarsal
digitules knobbed, 34–44, claw digitules: 26–33 µm long, slightly knobbed. Metathoracic
coxae with spinulae on ventral surface and small numbers on translucent pores. Tibiae
each with 5 setae (median seta present), tarsi each with 5 setae. Claw with denticle. Length
of spiracles 23–29 µm; posterior spiracles slightly larger than anterior. Venter with a small
number of scattered, hair-like setae some slightly knobbed and with two submarginal
rows of macrospines. Quinquelocular pores, 5 µm in diameter, distributed in bands
and rows on all segments of abdomen and thorax. Macrotubular ducts about 6–7 µm
wide and 15–20 µm long, scattered. Microtubular duct present in a submarginal band.
Cruciform pores 5 µm in diameter, very few around spiracles and on head.
Dorsum: Dorsal enlarged setae all spine-like, 10–47 µm long, blunted, varying in size and
shape. Dorsal spinose setae forming rows on segments. Enlarged setae in middle part
of last segments 10–15 µm long. On margin of segments 2 enlarged setae present, the
longest 23–47 µm long. Macrotubular ducts similar to those on venter, sparse, present
throughout, 8–10 µm wide, 15–25 µm long. Microtubular ducts with 2 sclerotized areas,
each 6 µm long, scattered throughout. Anal ring 40–50 µm wide, 48–76 µm long, with
pores and with 8 hair-like setae, each 74–120 µm long. Cauda absent (Redescription is
Other stages
Immature stages described in detail by Goux (1936).
Ecology
Host plant: Helichrysum stoechas.
Biology: Unknown.
Comments
There are capitated and pointed hair-like setae on venter, importance of this character
is not clear but this character is quite diverse among some genera. The size ranges show
very great variability, what deserves attention, and needs study of further materials.
Acanthococcidae 475
Figure 166. Rhizococcus helichrysi (Goux, 1936), female. Original.

Rhizococcus henmii (Kuwana, 1931) (Fig. 167) Combination nova

Eriococcus henmii Kuwana, 1931b: 168.
Syntypes: Female. Japan (Ryukyu Islands, Amami-oshima), on Chrysanthumum sp.,

06.xi.1930, by Mr. Yoshitada Maki. Deposited in Ibaraki-ken: Insect Taxonomy
Laboratory, National Institute of Agricultural Environmental Sciences, Kannon-dai,
Yatabe, Tsukuba-shi, (Kuwana).
Lit.: Hoy, 1963: 94; Kawai, 1980: 130; Kozár & Walter, 1985: 74; Köhler, 1998: 378;
Acanthococcus henmii; Kozár & Walter, 1985: 74. Change of combination.

Eriococcus henmii; Miller & Gimpel, 2000: 228. Revived combination.
Acanthococcus henmii; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Body of adult female, dark brown, with antennae and legs pale yellow; body segmentation
distinct; dorsal spines outstanding. Sac of female is grayish white in color, closely felted
and ovate, 2.5 mm long, 1.25 mm wide. Sac of male grayish, or grayish-white, 1.5 mm
long, 0.7 mm wide.
Mounted female
Body elongate oval, 1.5 mm long, 1 mm wide. Antennae usually 6 segmented (rarely
7), length of segments: I: 45, II: 31, III: 81, IV: 21, V: 20 and VI: 32 µm long, each
segment covered with a few, hair-like setae. Eyes situated on venter near margin. Anal
lobes elongated, sclerotized, each with two spinose setae along inner margin, and one
spinose seta on outer margin, much longer to those on dorsum, plus a long apical seta.
Venter: Labium 3 segmented, large. Legs well developed; tibia and tarsus subequal in
length, and two together slightly larger than the corresponding femur and trochanter;
claw large, curved with a small subspecial denticle; both pairs of digitules long, fliform
and knobbed apically, the tarsal pair the more robust; hind pair of coxae with pores
which are large, isolated and distinct. Discodial pores fairly numerous posteriorly and
around anal ring.
Dorsum: Dorsal enlarged setae spine-like, numerous, evenly, but irregularly distributed
over whole body, slightly more numerous perhaps on cephalic area, of 2 or 3 sizes, the
smallest about half size of the largest, latter stout, sharp pointed, smaller of same shape.
Macrotubular duct scattered (after Kuwana, 1931, with modifications).
Acanthococcidae 477
Ecology
Host plant: Chrysanthemum sp.
Biology: On the leaves and stem of the host plant.
Figure 167. Rhizococcus henmii (Kuwana, 1931), female After Kuwana (1931) with
modifications.
Rhizococcus henryi (Balachowsky, 1930) (Fig. 168) Combination nova

Eriococcus henryi Balachowsky, 1930: 314.
Syntypes: Female. France (Port-Cros, Antibes and Levant), on Euphorbia pithyusa and
E. spinosa 09.ix.1929, 11.ix.1929 and 20.viii.1929. Deposited in MNHN (type no.
4957).
Lit.: Balachowsky, 1930: 184; 1932: 233; 1933: 47; Foldi, 2000: 81; 2001: 305; 2003:
149; Goux, 1989: 21; Kiritchenko, 1940: 127; Kozár & Walter, 1985: 74; Köhler, 1998:
378; Ouvrard & Kozár, 2009: 102; Reyne, 1964: 103 (Miller et al., 2013).
Nidularia henryi; Lindinger, 1933a: 116. Change of combination.

Nidularia polyphagus; Lindinger, 1933a: 116. Change of combination.
Acanthococcus henryi; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus henryi; Miller & Gimpel, 2000: 228. Revived combination.
Acanthococcus (Asiacornococcus?) henryi; Kozár, 2009: 92. Change of combination.
Eriococcus polyphagus Goux, 1931: 68.
Lectotype: (MNHN 4960-3). France (Courzieu, Rhone, Brussieu), by L. Goux, on
Scabiosa columbaria, 28.viii.1928. The slide contains 3 females, the lectotype designated
here (marked), following the drawing of Goux. Paralectotypes: 2 females on the slide
of lectotype, 4 slides, MNHN 4960-2 (in basic collection), and MNHN 4960-04-06
in Goux collection, with the same data as lectotype. MNHN 4960-1 slide absent (last
remark in the collection is: loan to Kozár VII. 82). Notes: Synonymy by Balachowsky
(1933) with R. henryi (Balachowsky, 1930) verified.
Eriococcus tucurincae Laing, 1929: 467; misidentification by Balachowsky, 1933: 47. Notes:
Boratynski (1962: 59) states that the comparison of the types of E. henryi and E. tucurincae
shows that the 2 species are quite distinct, differing in many characters including the size
of the body, size, shape and distribution of the dorsal spines, size and shape of anal
lobes, relative and absolute sizes of the legs and their component joints.
Eriococcus henryi; Balachowsky, 1933: 47. Incorrect synonymy with E. tucurincae Laing,
1929: 467; discovered by Boratynski, 1962: 59.
Eriococcus polyphagus; Balachowsky, 1933: 47. Incorrect synonymy with E. tucurincae Laing,
1929: 467; discovered by Boratynski, 1962: 59.
Description
Unmounted female
Sac is oval, 3–4 mm long, white when new, turning cream colored with age. Eggs of
insects living on Euphorbia spinosa were red.
Mounted female
Body elongate oval, body segmentation distinct. Antennae usually 7 segmented, each
segment covered with a few, hair-like setae. Frontal lobe well developed. Eyes situated
Acanthococcidae 479
Figure 168. Rhizococcus henryi (Balachowsky 1930), female After Balachowsky (1930)
with modifications.
on venter near margin. Anal lobes slightly sclerotized, each with two enlarge setae along
inner margin, and one enlarged seta on outer margin, much longer to those on dorsum,
plus a long apical seta.
Venter: Labium 3 segmented, large. Stylet loop twice longer than labium. Legs well
developed; trochanter with a long hair-like setae, tibia and tarsus almost equal in length;
claw large, curved with a small subspecial denticle; both pairs of digitules long, knobbed
apically, the tarsal pair longer than claw; hind pair of coxae with large pores, isolated and
distinct. Macrotubular duct smaller than on dorsum, scattered. Discodial pores with 5
loculi, scattered all over venter.
Dorsum: Dorsal enlarged setae spine-like, numerous, distributed in bands over whole
body, slightly more numerous perhaps on cephalic area, of 2 perhaps 3 sizes, the smallest
about half size of the largest and situated at the last abdoıminal segment, latter stout,
sharp pointed, smaller of same shape, as long as wide. Macrotubular duct scattered with
spines. Microtubular ducts not seen. Cauda not seen. Anal ring sclerotised, with 6 anal
ring setae and partly double rows of pores (after Balachowsky (1930) with modifications.
Other stages
Immature stages described in detail by Goux (1931) as R. polyphagus.
Ecology
Host plant: Dianthus carthusianorum, Euphorbia characias, E. pithyusa, E. spinosa, Genista
sagittalis, Hieracium sp., Sedum sp., Scabiosa columbaria, Teucrium scorodonia.
Biology: Unknown.
Acanthococcidae 481
Rhizococcus heteroacanthus (Balachowsky, 1927) (Fig. 169) Redescription,

Combination nova
Eriococcus heteroacanthos Balachowsky, 1927: 204.
Holotype: Female. Algeria (El Guerrah, Constantine), on Fumana glutinosa, 15.xi.1926.

By original designation. Deposited in Alger: Insectarium Jardin d'Essai, Algeria.
Notes: Balachowsky (1927) mentions a type in his original description and suggests
that it is labeled as such and is deposited in Algeria. A second slide is in MNHN (coll.
number 4943), which contains two specimens and has similar label data except that
it was collected October 26, 1926. According to Author co-type is in collection of
E.E.Green.
Lit.: Goux, 1936: 299; Kozár, 2009: 92; Kozár & Walter, 1985: 74; Köhler, 1998: 378;
Ouvrard & Kozár, 2009: 102-118; Pellizari & Kozár, 2011: 66 (Miller et al., 2013).
Nidularia heteroacanthos; Lindinger, 1933a: 116. Change of combination.

Eriococcus heteroacanthos; Hoy, 1963: 95. Revived combination.
Acanthococcus heteroacanthos; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus heteroacanthos; Miller & Gimpel, 2000: 231. Revived combination.
Acanthococcus heteroacanthos; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Adult female is oval and green 2.1–2.6 mm long.
Mounted female
segments: I: 40, II: 28, III: 86 (sometimes with unclear segmentation), IV: 20, V: 18 and
VI: 30 µm long, each segment covered with a few, hair-like setae; apical segment with
long; on the preapical segment the falcate sensory seta 13–25 µm long. Frontal tubercle
present. Eyes situated on venter near margin. Anal lobes slightly sclerotized, with two
enlarged setae along inner margin, and one enlarged seta on outer margin, longer than
setae inner side, plus a long apical seta 270 µm long.
Venter: Labium 3 segmented 116 µm long; stylet loop not seen. Legs well developed;
lengths of segments prothoracic legs: coxa 70, trochanter 40, femur 110, tibia 92, tarsus
107 and claw 34 µm long; lengths of segments of mesothoracic legs; coxa: 72, trochanter:
50, femur: 113, tibia: 95, tarsus: 108 and claw: 35; lengths of segments of metathoracic
legs; coxa: 80, trochanter: 47, femur: 110, tibia: 108, tarsus: 120 µm long, tarsal digitules
knobbed 42–45, claw digitules: 36 µm long, slightly knobbed. Metathoracic coxae with
spinulae on ventral surface and small numbers on translucent pores. Tibiae each with 4
setae (median seta absent), tarsi each with 5 setae. Claw with denticle. Length of spiracles
28 µm; posterior spiracles slightly larger than anterior. Venter with a small number of
scattered, hair-like setae and with two submarginal rows of enlarged setae. Multilocular
pores quinquelocular, 6 µm in diameter, distributed in bands and rows on all segments
of abdomen and thorax. Macrotubular ducts of two sizes, smaller ones about 4 µm
wide, scattered in middle part of body on abdomen; larger macrotubular duct situated
on marginal area as those on dorsum. Microtubular duct present in a submarginal band.
Cruciform pores 5 µm long, very few around spiracles and on head.
Dorsum: Dorsal enlarged setae all spine-like, 9–52 µm long, with blunted apex, varying
in size and shape. Dorsal enlarged setae forming rows on segments. Enlarged setae in
middle part of last segments 9–30 µm long, some of them sligytly curved. On margin of
segments 2 enlarged setae present, the longest 52 µm long. Macrotubular ducts similar
to those sparse, present throughout, 8–10 µm wide. Microtubular ducts each 6 µm long,
scattered throughout. Discodial pores absent. Anal ring 48 µm wide, 75 µm long, with
pores and with 8 hair-like setae, each 115 µm long. Cauda present (Redescription is based
on type material).
Ecology
Host plant: Fumana glutinosa.
Distribution: Algeria, Italy.
Biology: On roots.
Acanthococcidae 483
Figure 169. Rhizococcus heteroacanthus (Balachowsky, 1927), female. Original.

Rhizococcus istresianus (Goux, 1989) (Fig. 170, 171) Redescription,

Combination nova
Eriococcus istresianus Goux, 1989: 19.
Holotype: Female. France (Alpes-Maritimes, Esterel, col Notre-Dame), by L. Goux,

01.x.1985, on Cistus salviaefolius (MNHN 14497-01). Paratypes: 5 slides from the same
collection as holotype, 13 more slides from the same place as holotype, but collected
21.IX.1985 (MNHN 14496-1-13), three slides from Bouches-du-Rhone, Istres, etang
de l, Olivier, on Cistus salviaefolius, by L. Goux, 25. VI. 1933 (MNHN 14532-1-3).
Deposited in MNHN.
Lit.: Foldi, 2001: 305; Kozár et al., 2013: 56 (Miller et al., 2013).
Acanthococcus istresianus; Miller & Gimpel, 1996: 601; Change of combination.

Eriococcus istresianus; Miller & Gimpel, 2000: 243. Revived combination.
Acanthococcus istresianus; Kozár, 2009: 92; Ouvrard & Kozár, 2009: 102. Revived
combination.
Eriococcus arboisi; Goux, 1991: 46-47. (Fig. 171) Synonym nova
Holotype: France (Bouches-du-Rhône, Aix-en-Provence, Les Milles, on "Petit
Arbois"), on on Poaceae, 25/05/1953, by L. Goux (MNHN 14434). By original
designation. Deposited in MNHN.
Acanthococcus arboisi; Miller & Gimpel, 1996: 598. Change of combination.
Eriococcus arboisi; Miller & Gimpel, 2000: 131. Revived combination.
Acanthococcus arboisi; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Ovisac pyriform, strongly convex, cream-colored, up to 2 mm long, 1 mm wide.
Mounted female
VII: 38–42 µm long; apical setae of antenna 40–55 µm long. On apical segment three
sensory falcate setae are found, the longest 30–35 µm long. On the preapical segment
the falcate sensory seta 28–40 µm long. The segments of the antennae are covered with
few hair–like setae. Frontal lobes well developed. Eyes situated on venter near margin.
Anal lobes slightly sclerotized each with two enlarged setae along inner margin, and one
enlarged seta on outer margin, the posterior on inner margin narrower, but longer, all
similar in size to those on dorsal margin, apical seta 240–310 µm long.
Venter: Labium 3-segmented, 95–101 µm long; stylet loop reaching the coxae of
median legs. Legs well developed; lengths of segments of prothoracic legs; coxa: 60–68,
trochanter: 35–48, femur: 129–134, tibia: 100–109, tarsus: 110–118 and claw: 29–40 µm
Acanthococcidae 485
Figure 170. Rhizococcus istresianus (Goux, 1989), female. Original.

long; lengths of segments of mesothoracic legs; coxa: 56–65, trochanter: 35–48, femur:
100–121, tibia: 109–115, tarsus: 119–127 and claw: 29–33 µm long; lengths of segments
of metathoracic legs: coxa: 65–75, trochanter: 35–43, femur: 100–140, tibia: 115–132,
tarsus: 128–138 and claw: 32–38 µm long; tarsal digitules knobbed, 40–51 µm long,
claw digitules, 30–36 µm long, slightly knobbed; claw with denticle. Metathoracic coxae
without spinulae and with medium number (less than 20) of translucent pores; tibiae
each with 5 setae (median seta present); claw with denticle. Length of anterior spiracles
26–33 µm; posterior spiracles slightly larger than anterior. Derm with a small number of
scattered, hair-like setae and two submarginal rows of enlarged setae. Quinquelocular
pores (sometime trilocular) distributed in sparse bands and rows on all segments of
abdomen and thorax, 3–5 µm in diameter. Macrotubular ducts one size, about 6 µm wide
and 15–16 µm long, scattered. Cruciform pores 5 µm long, scattered on submargin from
head to thirth abdominal segment.
Dorsum: Dorsal enlarged setae all with blunted apex, wide and strong, all of approximately
same size, abundant over dorsal surface; 12–49 µm long, dense; enlarged setae in middle
part of last segments 12–28 µm long; on margin of segments 3 spines present, 33–49 µm
long. Macrotubular ducts sparse, present throughout, 7–9 µm wide and 19–26 µm long.
Microtubular ducts with 2 sclerotized areas, 4 µm long, scattered throughout. Anal ring
not well seen, 50–55 µm wide and 64–68 µm long, with 8 hair-like setae, 110–130 µm
long. Cauda strongly sclerotized, 40–50 µm wide and 13–17 µm long.
Ecology
Host plant: Cistus salviifolius, Helichrysum sp. and some other Asteraceae.
Distribution: France, Greece, Hungary, Turkey.
Biology: On roots.
Comments
The paratypes on slides MNHN 14532-1-3 are somewhat different from holotype by
two sizes of spines on the middle of posterior dorsal segments. R. istresianus are similar
to R. munroi Boratynski (1962), but differs from this species by having smaller number of
spines on margin and longer dorsal spines on posterior abdominal segments. R. istresianus
similar to R. desertus, but differs from this species by having blunted spines, smaller
number of spines on margin and smaller size of the dorsal spines.
A detailed study of the holotype of R. arboisi showed that this species belongs to
R. istresianus, and here R. arboisi considered as a synonym of R. istresianus. The small
differences in form and size of some spines could be a variation. This species also similar
to R. reynei, which has smaller spines on dorsum and quinquelocular pores on mid-thorax
absent. This species very similar to slide present in MNHN, named as R. gnidii, which has
somewhat longer sizes of spines on dorsum, more on margin, different sizes of spines
on anal lobes, longer subapical setae on anal lobes, and some other detailes (France: Var,
env. Agay, MNHN 9344, P. Marchal, 4. 5. 1908, on Daphne gnidium, roots, written by ink
Acanthococcidae 487
Figure 171. Rhizococcus istresianus (Goux, 1989), female, based on type material of R.
arboisi Goux, 1991. Original.
on slide glass, on the other side on a paper label Racine de Daphne gnidium - Cruice del
Estrel - P. Marchal coll. 4.5.1908). However, only one specimen deposited in MNHN
in Paris, which could be studied at this time, and even this slide is not enough good for
detailed to study. For this reason we need to find more material of these species to clarify
this questions.
Additional information: In the revision slides in the collection of PPI specimens from
Töreki, Szársomlyó, Fehérszék and Öskű (Hungary), and from Levadia (Greece) appeared
to be R. istresianus, which are a new records for these countries.
Rhizococcus kurdicus (Bodenheimer, 1943) Redescription, Status and

Combination nova
Eriococcus aceris forma kurdica Bodenheimer, 1943: 21.
Syntypes: Female. Iraq (Ruwanduz Gorge), on Platanus orientalis, 11.x.1942, by F.S.

Bodenheimer. Deposited in Bet Dagan.
Lit.: Hoy, 1963: 67; Tudor, 1982: 89 (Miller et al., 2013).
Acanthococcus aceris kurdicus; Miller & Gimpel, 1996: 598. Change of combination.
Eriococcus aceris kurdica; Miller & Gimpel, 2000: 116. Revived combination.
Acanthococcus aceris kurdicus; Kozár, 2009: 91. Revived combination.
Description
Unmounted female
Adult female is oval and green, 1.7 mm long, 1.2 mm wide.
The white cocoons were fairly common on the leaves and tender twigs of Acer sp. Mature
mobil females were collected from leaves of Platanus sp.
Mounted female
Body elongate oval. Antennae 7 segmented, length of segments: I: 48, II: 36, III: 72, IV:
55, V: 24, VI: 19 and VII: 38 µm long, each segment covered with a few, hair-like setae;
apical segment with apical seta 50 µm long; apical segment also with 3 sensory falcate
setae, longest 34 µm long; on the preapical segment the falcate sensory seta 19–38 µm
long. Frontal lobes and tubercle present. Eyes situated on venter near margin. Anal lobes
sclerotized, short, conical, with two enlarged setae along inner margin, and one enlarged
seta on outer margin, longer than setae inner side, 140 µm long.
Venter: Labium 3 segmented 120 µm long; with hair-like setae, two pairs on the basal
segment, one of them about three times longer than other, stylet loop surpassing the
second coxae. Legs well developed; lengths of segments prothoracic legs: coxa: 108,
trochanter 60, femur 216, tibia 168, tarsus 144 and claw 36 µm long; lengths of segments
of mesothoracic legs: coxa 113, trochanter 62, femur 198, tibia 168, tarsus 150 and claw
Acanthococcidae 489
32 µm long; lengths of segments of metathoracic legs: coxa 120, trochanter 60, femur
204, tibia 178, tarsus 140, claw 36 µm long, tarsal digitules knobbed 56–58, claw digitules
36–38 µm long, slightly knobbed. Metathoracic coxae with spinulae on ventral surface
and small numbers on translucent pores. Tibiae each with 5 setae (median seta present),
tarsi each with 5 setae. Claw with denticle. Length of spiracles 41 µm; posterior spiracles
slightly larger than anterior. Venter with a small number of scattered, hair-like setae and
with two (three-four on thorax and head) submarginal rows of enlarged setae. Multilocular
pores quinqueloculars (rarely triloculars), 6 µm in diameter, distributed in bands and rows
on all segments of abdomen and thorax. Macrotubular ducts of two sizes, shorter ones
about 6 µm wide, 18 µm wide scattered in middle part of body on abdomen; larger
macrotubular duct situated on marginal area as those on dorsum. Microtubular duct
present in a submarginal band. Cruciform pores 4 µm long, numerous on margin.
Dorsum: Dorsal enlarged setae all spine-like, 45–68 µm long, some with truncated apex,
others blunted. Dorsal enlarged setae forming rows and bands on segments. Enlarged
setae in middle part of last segments 45–62 µm long. On margin of segments 2 enlarged
setae present, the longest 68 µm long. Macrotubular ducts similar to those sparse, present
throughout, 8–10 µm wide, 24 µm long. Microtubular ducts each 4 µm long, scattered
throughout. Anal ring, with pores and with 8 hair-like setae 0.60 µm long (Redescription
is based on type material).
Ecology
Host plant: Platanus orientalis.
Distribution: Iraq.
Comments
Slide-mounted adult female differs from Acanthococcus aceris aceris by having in all specimens
from both localities the third antennal segment longer that the fourth segment. According
to short description the status can not be solved clearly, however several detailes (like very
short anal ring setae, and anal lobe setae, by the short antennae and legs, by five setae on
tibia, by presence of females on the leaves of host plants may indicate that this species
not related to the A. aceris species and transferred to Rhizococcus genus. But it is clear that
this species needs further studies.
Rhizococcus lactucae (Borchsenius, 1949) (Fig. 172) Redescription,

Combination nova
Acanthococcus lactucae Borchsenius, 1949: 339.
Lectotype: Female. Tajikistan (Melnikovo, near Kanibadan, near railway station),

on Lactuca sp. leaves, 15.ix.1944, by N. Borchsenius. By subsequent designation of
Danzig, 1996: 521. Deposited in ZMAS (type no. 3-44). Notes: One paralectotype on
same slide as lectotype.
Lit.: Danzig, 1975: 45; 1996: 575; Hoy, 1963: 98 (Miller et al., 2013).
Nidularia lactucae; Lindinger, 1957: 543. Change of combination.

Eriococcus lactucae; Hoy, 1963: 98. Change of combination.
Acanthococcus lactucae; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus lactucae; Tang & Hao, 1995: 452. Revived combination.
Acanthococcus lactucae; Köhler, 1998: 379. Revived combination.
Eriococcus lactucae; Miller & Gimpel, 2000: 249. Revived combination.
Acanthococcus lactucae; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Adult female is egg-shaped, 3.5 mm long, 2.0 mm wide, and olive in color.
Ovisac pyriform, strongly convex, cream-colored, up to 4.5 mm long, 2.5 mm wide.
Mounted female
Body elongate oval. 3.5 mm long, 1.9 mm wide. Antennae 7 segmented, length of
segments: I: 38–43, II: 36, III: 60–67, IV: 45–53, V: 24–29, VI: 24 and VII: 36–41 µm
long; apical setae of antenna 49–53 µm long. On apical segment three sensory falcate
setae are found, the longest 24–26 µm long. On the preapical segment the falcate sensory
seta 17–31 µm long. The segments of the antennae are covered with few hair–like setae.
Frontal lobes and tubercles present. Eyes situated on venter near margin. Anal lobes
outer margin latter shorter than others, all similar in size to those on dorsal margin, apical
seta 360 µm long.
Venter: Labium 3 segmented, 120–125 µm long; basal segment with almost the same
size of setae, stylet loop three times longer than the labium. Legs well developed; lengths
of segments of prothoracic legs; coxa 84–96, trochanter: 60–72, femur: 180–192, tibia:
144–156, tarsus: 144 and claw: 36–39 µm long; lengths of segments of mesothoracic
legs; coxa: 96–103, trochanter: 58–68, femur: 172–180, tibia: 151–156, tarsus: 151
and claw: 38–39 µm long; lengths of segments of metathoracic legs: coxa: 101–110,
trochanter: 65–79, femur: 185–190, tibia: 168–173, tarsus: 158–170 and claw: 38–41 µm
long; tarsal digitules knobbed, 48–58 µm long, claw digitules, 36–43 µm long, slightly
Acanthococcidae 491
Figure 172. Rhizococcus lactucae (Borchsenius, 1949), female. Original.

knobbed; claw with denticle at near apex. Metathoracic coxae with spinulae and medium
number (less than 20) of translucent pores (a few on edge of femur); tibiae each with 5
setae (median seta present); claw with denticle. Length of anterior spiracles 36–38 µm;
posterior spiracles slightly larger than anterior. Derm with a small number of scattered,
hair-like setae and submarginal sparse bands of enlarged setae; in group on each segment.
Quinquelocular pores distributed in sparse bands and rows on all segments of abdomen,
thorax and head (a few), 3–5 µm in diameter. Macrotubular ducts of two sizes, larger
ones as those on dorsum, on body margin; smaller ones about 3 µm wide and 21 µm long,
scattered. Cruciform pores 5 µm long, few scattered in a submarginal band from head to
second abdominal segment.
Dorsum: Dorsal enlarged setae all with blunted apex, wide and strong, varying in size
and shape; 29–72 µm long, dense; on margin of segments 2 spines present, 72 µm long.
Dorsal derm with microtrichia. Macrotubular ducts sparse, present throughout, 7 µm
wide and 24 µm long. Microtubular ducts each 5 µm long, scattered throughout. Anal
ring not well seen, 62–73 µm wide and 96 µm long, with 8 hair-like setae, 124–156 µm
long. Cauda present (Redescription based on type material).
Ecology
Host plant: Lactuca sp.
Biology: On the lower surface of leaves and shoots.
Acanthococcidae 493
Rhizococcus marginalis (Borchsenius, 1949) (Fig. 173) Redescription,

Combination nova
Acanthococcus marginalis Borchsenius, 1949: 336.
Lectotype: Female. Armenia (Megri), on Kochia prostrata leaves (xerophilous

Chenopodiaceae), 26.v.1947, by N. Borchsenius. By subsequent designation Danzig,
1996: 521. Deposited in ZMAS (type no. 80-48).
Lit.: Danzig, 1996: 575; Hoy, 1963: 101; Tao, 1999: 32; Ter-Grigorian, 1983: 877;
Wang, 2001: 209 (Miller et al., 2013).
Nidularia marginalis; Lindinger, 1957: 543. Change of combination.

Eriococcus marginalis; Hoy, 1963: 101. Change of combination.
Acanthococcus marginalis; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus marginalis; Tang & Hao, 1995: 452. Revived combination.
Acanthococcus marginalis; Köhler, 1998: 380. Revived combination.
Eriococcus marginalis; Miller & Gimpel, 2000: 262. Revived combination.
Acanthococcus marginalis; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Adult female broadly oval.
Mounted female
Body elongate oval. 3.1 mm long, 2.8 mm wide. Antennae 7 segmented, length of
segments: I: 41, II: 34, III: 43, IV: 48, V: 29, VI: 26 and VII: 43 µm long; apical setae
of antenna 41 µm long. On apical segment three sensory falcate setae are found, the
longest 20 µm long. On the preapical segment the falcate sensory seta 14–24 µm long.
The segments of the antenna are covered with few hair–like setae. Frontal lobes and
tubercles present. Eyes situated on venter near margin. Anal lobes sclerotized each with
two enlarged setae along inner margin, and one enlarged seta on outer margin, all similar
in size to those on dorsal margin, apical seta 125 µm long subapical and suranal setae
equal long, 95 m, hair-like.
Venter: Labium 3 segmented, 125 µm long; basal segment with two pair of differently
sized hair-like setae, stylet loop three times longer than the labium. Legs well developed;
lengths of segments of prothoracic legs; coxa: 79, trochanter: 60, femur: 149, tibia:
125, tarsus: 120 and claw: 41 µm long; lengths of segments of mesothoracic legs; coxa:
79, trochanter: 53, femur: 134, tibia: 137, tarsus: 122 and claw: 46 µm long; lengths of
segments of metathoracic legs; coxa: 91, trochanter: 67, femur: 151, tibia: 146, tarsus: 139
and claw: 41 µm long; tarsal digitules knobbed, 51–53 µm long, claw digitules, 36 µm long,
slightly knobbed; claw with denticle at near apex. Metathoracic coxae with spinulae and
medium number (less than 20) of translucent pores (a few on edge of femur); tibiae each
with 5 setae (median seta present); claw with denticle. Length of anterior spiracles 31 µm;
posterior spiracles slightly larger than anterior. Derm with a small number of scattered,
hair-like setae, preanal setae 80 µm long, and with submarginal band and a marginal
row of enlarged setae. Quinquelocular pores (some of them trilocular) distributed in
sparse bands and rows on all segments of abdomen, thorax and head (a few), 4–5 µm
in diameter. Macrotubular ducts of two sizes, larger ones as those on dorsum, on body
margin; smaller ones about 4 µm wide and 24 µm long, scattered. Cruciform pores absent.
Dorsum: Dorsal enlarged setae all with sharp pointed apex, wide and strong, varying
in size and shape; 24–53 µm long, on margin of segments 2 or 3 spines present, 36–
72 µm long, other dorsal setae in band on abdominal segments and thorax and head.
Macrotubular ducts, present throughout, 6 µm wide and 24 µm long. Microtubular ducts
each 5 µm long, scattered throughout. Anal ring, 72 µm wide and 72 µm long, with 8 hair-
like setae, 120 µm long. Cauda present (Redescription is based on type material).
Ecology
Host plant: Kochia prostrata.
Distribution: Armenia, China.
Acanthococcidae 495
Figure 173. Rhizococcus marginalis (Borchsenius, 1949), female. Original.

Rhizococcus matesovae (Miller & Gimpel, 1996) (Fig. 174) Redescription,

Combination nova
Acanthococcus multispinosus Matesova, 1976: 24.
Holotype: Female. Kazakhstan (West Kazakhstan, near Zhetybay, 5 km. S. Aral-Sor

Lake), 25.vi.1971, by G.Matesova. By original designation. Deposited in ZMAS (type
no. 3402). Notes: There is also 1 female paratype in ZMAS with the type data: West
Kazakhstan, near Karaulakeldy, 02.vii.1969, Matesova. There are other paratypes in
AAKA.
Lit.: Danzig, 2006: 203; Kozár & Walter, 1985: 74; Miller & Gimpel, 1996: 602; Miller
& Gimpel, 1999: 214; 2000: 263; Tang & Hao, 1995: 479 (Miller et al., 2013).
Acanthococcus matesovae; Miller & Gimpel, 1996: 602. Replacement name. Notes: A.
multispinosus Matesova, what is preoccupied by A. multispinosus (Kuhlgatz, 1898).
Eriococcus matesovae; Miller & Gimpel, 1999: 214. Change of combination. Since the
publication of Miller & Gimpel (1996) A. matesovae and A. multispinosus (Kuhlgatz) have
been transferred to Eriococcus and the name E. matesovae was still justified.
Acanthococcus multispinosus; Kozár, 2009: 93; Revived combination. Notes: Because of the
transfer of A. multispinus (Kuhlgatz) to Rhizococcus genus. Now, after the transfer of A.
multispinosus Matesova to the genus Rhizococcus, the name R. matesovae still valid.
Description
Unmounted female
Color in beginning brown, latter brownish-red, and deep-purple.
Mounted female
Body elongate oval, 1.96 mm long, 0.78 mm wide. Antennae 7 segmented, 242.5 µm,
length of segments: I: 65, II: 40, III: 47.5, IV: 45, V: 22.5, VI: 25, and VII: 35 in µm long;
each segment (except third) covered with a few, strong hair-like setae; apical segment with
apical seta 42.5 µm long; apical segment also with 3 sensory falcate setae, each 25–35
falcate seta 22.5 µm long. Frontal lobe and frontal tubercle present. Eyes situated on
venter near margin. Anal lobes strongly developed, each with 3 enlarged setae, slightly
truncated, each 35–40 µm plus 7-9 microtubular ducts on dorsal surface; apical seta 165
µm; ventral hair-like subapical seta 80–95 µm long, in addition on the ventral side of the
anal lobe with three hair-like setae each 50–55 µm long.
Venter: Labium 195 µm long, 105 µm wide, median setae on apex of labium short,
12.5 µm long. Legs well developed; lengths of segments and digitules of prothoracic
legs: coxa: 75.0–87.5, trochanter: 62.5–65.0, femur: 117.5–120.0, tibia: 100, tarsus: 105
and claw: 30, trochanther + femur: 175–180, tibia + tarsus: 205.0–207.5, tarsal digitules:
42.5, claw digitules: 32.5 µm long; lengths of segments and digitules of mesothoracic
Acanthococcidae 497
Figure 174. Rhizococcus matesovae (Miller & Gimpel, 1996), female. Original.
legs: coxa: 85, trochanter: 60.0–62.5, femur: 115, tibia: 105, tarsus: 110 and claw: 30,
trochanther + femur: 180, tibia + tarsus: 210, tarsal digitules: 45 µm long, claw digitules
broken; lengths of segments and digitules of metathoracic legs: coxa: 87.5–90.0,
trochanter: 65–70, femur: 120, tibia: 115, tarsus: 110 and claw: 32.5, trochanther + femur:
180, tibia + tarsus: 230, tarsal digitules: 37.5–42.5, claw digitules: 30.0–32.5 µm long.
Tarsal and claw digitules knobbed. Metathoracic coxae each with spinulae and translucent
pores. Tibiae each with 5 setae (median seta present), tarsi each with 5 setae. Length of
spiracles 52.5–55 µm; diameter of spiracular peritreme 32.5–35.0 µm, posterior spiracles
setae, slightly blunted, each 12.5–60 µm long. Enlarged setae, situated on margin in 2
or 3 rows, each 20–23 µm long. Multilocular pores each 5–7 µm in diameter and with
3-5 loculi, distributed in sparse bands on all abdominal and thoracic segments and head
(trilocular pores generally on head and thorax and slightly smaller than quinquelocular
pores). Macrotubular ducts of one size, each 5–6 µm wide and 15–17.5 µm long,
present on scattered on abdomen, thorax and head. Cruciform pores a few, each, 3–4
µm in diameter, distributed on the submargin and median part of thorax and head and
submargin of first abdominal segments.
Dorsum: Marginal setae blunted, slightly truncated, 40–48 µm long, numerous, not
forming a marginal row and not strongly differentiated from dorsal setae; other dorsal
setae wide, 25–35 µm long, those on thorax, head and anterior abdominal segments
slightly larger than setae on posterior 2 or 3 abdominal segments; arranged in transverse
bands across each body segment, rows irregular on head. Macrotubular ducts, each 7-8
µm wide and 14–18 µm long, scattered throughout dorsum, generally in segmental bands.
Microtubular ducts, 3–4 µm long scattered over dorsum. Anal ring strongly sclerotized,
with pores on inner side, 75 µm in diameter, with 8 setae, each 105–128 µm long; anal
ring situated on margin of dorsum. Cauda present with large microtrichia as well as on
dorsal surface (Redescription is based on type material).
Ecology
Host plant: Artemisia frigida, A. maritima, A. terrae-albae, Silene brahuica.
Biology: On the roots, rarely on root crown.
Acanthococcidae 499
Rhizococcus micracanthus (Danzig, 1975) (Fig. 175) Combination nova

Acanthococcus micracanthus Danzig, 1975: 52.
Holotype: Female. Russia (Southern Primor'ye, Lazovskiy Reserve, Glazkovka), on

unidentified host, 03.viii.1969, by E. Danzig. By original designation. Deposited in
ZMAS.
Lit.: Danzig, 1980: 206; Kaydan, et al., 2005: 399; Kaydan, et al., 2007: 90; Kozár
et al., 2009: 436; Kozár et al., 2013: 56; Kozár & Walter, 1985: 74; Kwon & Han,
2003: 151; Marotta, 1993: 157; Pellizzari & Kozár, 2011: 66; Tang & Hao, 1995: 452;
Tereznikova, 1981: 32 (Miller et al., 2013).
Eriococcus micracanthus; Tang & Hao, 1995: 477-478. Change of combination.

Acanthococcus micracanthus; Köhler, 1998: 380. Revived combination.
Eriococcus micracanthus; Miller & Gimpel, 2000: 266. Revived combination.
Acanthococcus micracanthus; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Body oval, dark-red, or violet, up to 3 mm long. Ovisac dirthy-white, eggs pink.
Mounted female
Body elongate oval. Antennae 7 segmented, apical segment with 3 narrow, long sensory
falcate setae, segments V-VI with one long, sensory falcate seta. Frontal lobe and frontal
tubercle present. Eyes situated on venter near margin. Anal lobes strongly developed,
each with 3 enlarged setae, blunted, two noticeably narrow, one similar to other marginal
setae.
Venter: Labium long, median setae on apex of labium short. Legs well developed. Tarsal
and claw digitules knobbed. Metathoracic coxae each with spinulae on ventral surface
and translucent pores on coxa. Tibiae each with 5 setae (median seta present), tarsi each
with 4 setae. Derm with a sparse covering of scattered flagellate hair-like setae. Enlarged
setae, situated on submargin in 1 row. Multilocular pores with 5-7 loculi, distributed in
wide bands on all abdominal and thoracic segments and head. Macrotubular ducts of
two sizes, larger ones on the margin, smaller ones scattered on abdomen, thorax and
head. Microtubular ducts scattered on margin. Cruciform pores few, distributed on the
submargin.
Dorsum: Marginal setae blunted, enlarged setae on thorax and head conical with sides
straight and apices acute, slightly larger along body margin, anterior abdomen, thorax and
head with small setae in medial and mediolateral areas, 2 or 3 lateral setae near margin of
each abdominal segment; forming a marginal row and quite different from dorsal setae;
those on thorax, head and anterior abdominal segments larger than setae on posterior 5
or 6 abdominal segments; arranged in 2 or 3 transverse rows across each body segment,
rows irregular on head. Macrotubular ducts, each 7–8 µm wide and 14–18 µm long,
scattered throughout dorsum, generally in segmental bands. Microtubular ducts, with 2
sclerotized areas, scattered over dorsum. Anal ring strongly sclerotized, with pores on
inner side, with 8 setae, anal ring situated on margin of dorsum. Cauda present (after
Danzig (1975), with modifications and changes).
Ecology
Host plant: Dianthus sp., Foeniculum vulgare, Galium sp., Scabiosa fischeri, S. lachnophylla,
Salvia sp., Potentilla chinensis.
Distribution: Hungary, Italy, North Korea, Russia, Turkey, Ukraine.
Comment
On studied materials the number of marginal setae more than on drawing of Danzig
(1975) and the shape of setae differs.
Acanthococcidae 501
Figure 175. Rhizococcus micracanthus (Danzig, 1975), female. After Tereznikova (1981)
with modifications.
Rhizococcus minimus (Tang, 1988) (Fig. 176) Revived combination

Acanthococcus minimus Tang in Tang & Li, 1988: 71.
Syntypes: Female. China (Nei Monggol, Ningcheheng County), on Artemisia argyi,

female. Deposited in Shanxi.
Lit.: Hua, 2000: 138; Miller & Gimpel, 2000: 268; Tang & Hao, 1995: 519; Tao, 1999:
35; Wang, 2001: 225 (Miller et al., 2013).
Rhizococcus minimus; Tang & Hao, 1995: 352. Change of combination.

Eriococcus minimus; Miller & Gimpel, 1999: 214. Change of combination.
Acanthococcus minimus; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Unknown.
Mounted female
Body elongate oval. Antennae 6 segmented. Eyes situated on venter near margin. Anal
lobes developed, with 3 enlarged setae, pointed on dorsal side; on ventral side with 2
hair-like setae.
Venter: Labium small, stylet loop twice longer than labium. Legs well developed; tarsal
and claw digitules knobbed longer than claw. Metathoracic coxae with translucent pores.
Tibiae each with 5 setae (median seta present), tarsi each with 5 setae. Derm with a sparse
covering of scattered short flagellate hair-like setae. Enlarged setae, situated on submargin
in 2 or 3 rows. Quinquelocular pores distributed in sparse bands on all abdominal and
thoracic segments and head. Macrotubular ducts of two sizes, larger ones as those on
dorsum on the margin; smaller ones in row on the abdominal segment, scattered on
thorax and head. Cruciform pores few, distributed on the submargin and median part of
thorax and head and submargin of first abdominal segments.
Dorsum: Marginal enlarged setae long, blunted, forming a marginal row 2 on each
abdominal segments; other dorsal setae much smaller than marginal setae, those on
or 3 abdominal segments; arranged in 1 or 2 transverse rows across each body segment,
rows irregular on head. Macrotubular ducts, wide, scattered throughout dorsum, generally
in segmental bands. Microtubular ducts with 2 sclerotized areas scattered over dorsum.
Anal ring strongly sclerotized, with 8 setae, situated on margin of dorsum (Tang & Hao,
1995).
Acanthococcidae 503
Figure 176. Rhizococcus minimus (Tang, in Tang & Li, 1988), female. After Tang & Hao
Ecology
Host plant: Artemisia argyi.
Rhizococcus miscanthi (Takahashi, 1957) (Fig. 177) Combination nova

Eriococcus miscanthi Takahashi, 1957: 6.
Syntypes: Female. Japan (Kyushu, Tsukumi), on Miscanthus sp., 02.ix.1955, by T.

Tachikawa, female. Deposited in Sapporo.
Lit.: Hoy, 1963: 102; Kawai, 1980: 130; Köhler, 1998: 380; Kozár & Walter, 1985: 74;
Acanthococcus miscanthi; Kozár & Walter, 1985: 74. Change of combination.

Eriococcus miscanthi; Miller & Gimpel, 2000: 269. Revived combination.
Acanthococcus miscanthi; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female is yellow, somewhat brownish at mid-dorsal line, slightly covered with wax.
Mounted female
Body narrow a little ower twice as long as wide in specimens, tapering posteriorly on the
abdomen, not much convex dorsally, about 2 mm long. Antennae 7 segmented, a little
longer than space between them, the third segment the longest, a little longer than fourth;
the fourth much longer than wide. Eyes situated on venter near margin. Anal lobes little
converging, sclerotised, without teeth, with 4 dorsal setae (2 along the lateral margin and
2 shorter ones on the mesal margin) (although on Authors drawing 3 setae are shown!),
and 3 very long fine setae on the venter, 2 of which are near the base; apical seta very long
fine distally, on the basal part stouter than the anal ring setae (Takahashi, 1957).
Venter: Legs, long, hind legs reaching beyond midlength of abdomen; hind coxae longer
than wide, with many minute pores on the lateral side; middle and hind tibiae nearly as
long as the femur; tarsi nearly as long as the tibia, tarsal digitules a little longer and stouter
than the claw digitules; claw slender, with a denticle hardly discernible. Macrotubular
ducts similar but longer than ducts present along the margin and on the marginal areae
of the venter of abdomen, twice as long as wide and 5 or 6 on each side of venter of
the last abdominal segment. Quinquelocular pores present over derm, numerous on the
posterior part, but few on the anterior part about 4 or fewer on the basal part of anal lobe.
Ventral setae long, fine, in a row on each abdominal segment, 6 on abdominal segment VII,
10 abdominal segment VI and many long setae in a cluster in front of mouth part.
Acanthococcidae 505
Figure 177. Rhizococcus miscanthi (Takahashi 1957), female. After Takahashi (1957) with
modifications.
Dorsum: Enlarged dorsal setae spine-like, blunted, not curved, numerous in a marginal
band, over the whole dorsum, but absent on the last abdominal segment in front of anal
ring; those along the margin distinctly larger, slender, over twice as long as the smallest
ones; mostly 3 of these larger setae present on each abdominal segment on each side;
shorter dorsal setae rather stout, twice or thrice as long as wide; these shorter setae
present also outside of the large marginal setae, mostly 3 or 4 on each side. Macrotubular
duct short, stout, somewhat wider than long, smaller in diameter than bases of dorsal
setae, more numerous than the dorsal setae, with a broad rim and a thin filament at the
inner end, many microtubular ducts also scattered on the dorsum intermixed with the
setae and large ducts, at least some of these microtubular ducts with a pair of minute
pointed processes at the orifice. In some specimens many small granules discernible
between segment on the dorsum, which are rounded or blunt at the tip and pale. Anal
ring strongly sclerotized, with 8 setae (Takahashi, 1957 with some modifications).
Ecology
Host plant: Miscanthus sp.
Biology: Unknown.
Rhizococcus multispinatus Tang & Hao, 1995 (Fig. 178) Revived

combination
Rhizococcus multispinatus Tang & Hao, 1995: 598.
Holotype: Female. China (Ningxia), on Artemisia sp., ?.v.1991. By original designation.

Deposited in Shanxi.
Lit.: Miller & Gimpel, 2000: 272; Tang & Hao 1995: 520, 655; Tao, 1999: 35; Wang,
2001: 225 (Miller et al., 2013).
Acanthococcus multispinatus; Miller & Gimpel, 1996: 602. Change of combination.

Eriococcus multispinatus; Miller & Gimpel, 1999: 214. Change of combination.
Acanthococcus multispinatus; Kozár, 2009: 100. Revived combination.
Description
Unmounted female
Adult female is oval.
Mounted female
third and fourth segments the longest; of which third longer than fourth. Eyes situated
Acanthococcidae 507
Figure 178. Rhizococcus multispinatus Tang & Hao, 1995, female. After Tang & Hao (1995)
with modifications.
on venter near margin. Anal lobes short, conical, and sclerotised, with three dorsal
enlarged setae each same size, shorter than on abdominal margin.
Venter: Mouthparts developed, stylet loop as long as labium. Legs well developed; tarsus
scarcely longer than tibia; digitules of tarsus and claw slender, knobbed and surpassing
the tip of claw, with end expanded; a denticle present distinctly on underside of claw;
there are number of traslusent pores on hindcoxa. Tibiae each with 5 setae (median
seta present), tarsus with 5 setae. Posterior spiracles slightly larger than anterior. Derm
with a sparse covering of scattered flagellate hair-like setae. Enlarged setae, situated on
submargin in rows. Quinquelocular pores distributed in sparse bands on all abdominal
and thoracic segments and head. Macrotubular ducts of two sizes, larger ones as those
on dorsum on the margin; smaller ones in row on the abdominal segment, scattered on
thorax and head. Cruciform pores a few, distributed on the submargin and median part
of thorax and head and submargin of first abdominal segments. Microtubular ducts
scattered over margin (Tang & Hao, 1995).
Dorsum: Marginal enlarged setae blunted, or slightly truncated, long, each 40–80 µm,
forming a marginal band; 3-5 on each abdominal segments and strongly differentiated
from dorsal setae; other dorsal setae wide, 10–20 µm long, distributed in segmental
bands, the band on thorax about 5 or 6 enlarged setae wide; and on abdomen about
2-3 setae wide. Macrotubular ducts of one size scattered throughout dorsum, generally
in segmental bands. Microtubular ducts with 2 sclerotized areas scattered over dorsum.
Anal ring strongly sclerotized, with 8 setae, anal ring situated on margin of dorsum (after
Tang & Hao, (1995) with modifications).
Ecology
Host plant: Artemisia sp., Artemisia ordosica, Taraxacum mongolicum.
Acanthococcidae 509
Rhizococcus multispinosus Kuhlgatz, 1898 (Fig. 179)

Rhizococcus multispinosus Kuhlgatz, 1898: 185.
Syntypes: Female. Germany (in the Warmhause des Königlichen botanischen

Museums zu Berlin), on Austrocylindropuntia vestita (Opuntia), by Schumann. Deposited
in Berlin.
Lit.: Cockerell, 1900: 595; Fernald, 1903: 76; Hoy, 1963: 103; Kozár & Walter, 1985:
74; Köhler, 1998: 380; Lindinger, 1931: 114; Miller & Gimpel, 2000: 273 (Miller et al.,
2013).
Eriococcus multispinosus; Cockerell, 1900: 595. Change of combination.

Acanthococcus multispinosus; Miller & Gimpel, 1996: 602. Change of combination.
Eriococcus multispinosus; Miller & Gimpel, 1999: 214. Revived combination.
Rhizococcus multispinosus; Kozár, 2009: 106; Revived combination.
Eriococcus coccineus; Cockerell, 1894a: 204. This species was incorrectly treated as a junior
synonym by Lindinger (1931) and was considered a possible synonym by Cockerell
(1900) and (Hoy, 1963).
Rhizococcus multispinosus; Kuhlgatz, 1898 is the senior secondary homonym of R.
multispinosus (Matesova, 1976), which is now named R. matesovae.
Description
Unmounted female
Adult female is oval, 1.75–2.75 mm long, darkreddish-brown.
Mounted female
Body elongate oval. Antennae 7 segmented, covered with a few, strong hair-like setae;
anal lobes well developed, each with 3 enlarged setae, blunted.
Venter: Labium well developed, stylet loop reaches behind the posterior coxae. Legs
well developed; Tarsal and claw digitules capitated. Tibiae each with 5 setae (median seta
present), tarsi each with 6 setae. Derm with a sparse covering of scattered flagellate hair-
like setae. Enlarged setae, situated on submargin in rows. Macrotubular ducts of one size,
each 5–present on scattered on abdomen, thorax and head.
Dorsum: Enlarged setae conical, sides straight, apices rounded, marginal setae
conspicuously larger than others on dorsum, 3 lateral setae on margin of most abdominal
segments; forming a marginal row, strongly differentiated from small dorsal setae;
arranged in 2 or 3 transverse rows across each body segment, rows irregular on head.
Macrotubular ducts, scattered throughout dorsum. Anal ring with 8 setae, anal ring
situated on margin of dorsum (Kuhlgatz, 1898).
Other stages
Simplifyed drawing of first instar nymph and winged adult male is given.
Ecology
Host plant: Austrocylindropuntia vestita (Opuntia).
Distribution: Germany (Specimens were collected in Germany from a South American
plant. The species is therefore considered to be of South American in origin).
Biology: On the plant surface.
Figure 179. Rhizococcus multispinosus (Kuhlgatz, 1898), female), female After Kuhlgatz
Acanthococcidae 511
Rhizococcus mumtazi (Bodenheimer, 1943) (Fig. 180) Redescription,

Combination nova
Eriococcus mumtazi Bodenheimer, 1943: 22.
Syntypes: Female. Iraq (Shaklawa and Zakho), on undetermined Gramineae,

08.x.1942 and 11.x.1942. Deposited in Bet Dagan.
Lit.: Ben-Dov & Harpaz, 1986: 30; Hoy, 1963: 103; Köhler, 1998: 381 (Miller et al.,
2013).
Acanthococcus mumtazi; Kozár & Walter, 1985: 74. Change of combination.

Eriococcus mumtazi; Miller & Gimpel, 2000: 275. Revived combination.
Acanthococcus mumtazi; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Body elongate, 2 mm long, 0.8 mm wide. Ovisac is white and felty.
Mounted female
Body elongate oval. Antennae 6 segmented, length of segments: I: 36, II: 31, III: 108,
IV: 29, V: 24, and VI: 43 µm long; each segment (except third) covered with a few, strong
hair-like setae. Eyes situated on venter near margin. Anal lobes present, each with 3
enlarged setae, 40–60 µm long, slightly blunted; ventrally with long apical seta and three
hair-like seta, each 50–100 µm long.
Venter: Labium 98 µm long, slightly longer than wide, stylet loop twice longer than
labium. Legs well developed; lengths of segments prothoracic legs; coxa: 78, trochanter:
55, femur: 155, tibia: 120, tarsus. 133 and claw: 27 µm long; lengths of segments of
mesothoracic legs; coxa: 85, trochanter: 48, femur: 165, tibia: 137, tarsus: 135 and claw:
31 µm long; lengths of segments of metathoracic legs; coxa: 91, trochanter: 60, femur:
172, tibia: 155, tarsus: 143 and claw: 35 µm long, tarsal digitules knobbed 40 µm long,
claw digitules slightly knobbed 30.0–32.5 µm long. Metathoracic coxae each with spinulae
on ventral surface and translucent pores on coxa. Tibiae each with 5 setae (median seta
present), tarsi each with 5 setae. Spiracles 33 µm diameter; posterior spiracles slightly larger
than anterior. Derm with a sparse covering of scattered flagellate hair-like setae. Enlarged
setae, situated on submargin in a row. Quinquelocular pores 6 µm in diameter distributed
in sparse bands on all abdominal and thoracic segments and head. Macrotubular ducts
of two sizes, larger ones on body margin, 4 µm wide and 15 µm long, smaller ones 3 µm
wide and 16 µm long present on scattered on abdomen, thorax and head. Microtubular
ducts 4 µm long in marginal band. Cruciform pores few, 4 µm in diameter, distributed
on the submargin and median part of thorax and head and submargin of first abdominal
segments.
Dorsum: Marginal enlarged setae sharp and pointed, long, marginal setae longer than
other dorsal setae; forming a marginal row, 41–53 µm long, other dorsal setae wide,
31–35 µm long, those on thorax, head and anterior abdominal segments slightly larger
than setae on posterior abdominal segments; arranged in 2 or 3 transverse rows across
each body segment, rows irregular on head. Macrotubular ducts, each 6 µm wide and 15
µm long, scattered throughout dorsum, generally in segmental bands. Microtubular ducts,
4 µm long scattered over dorsum. Anal ring sclerotized, with a single row of pores, with 8
setae, each 100 µm long (Bodenheimer (1943), with additions from syntype female, with
kind help of Dr. Yair Ben-Dov).
Ecology
Distribution: Iraq
Figure 180. Rhizococcus mumtazi (Bodenheimer, 1943), female. After Bodenheimer

Acanthococcidae 513
Rhizococcus munroi (Boratynski, 1962) (Fig. 181) Combination nova

Acanthococcus munroi Boratynski, 1962: 56.
Holotype: Female. United Kingdom (England, Berkshire, Sunninghill, Silwood

Park, Imperial College Field Station), on Achillea millefolium. By original designation.
Deposited in BMNH (type no. 151c).
Common name: Munro's felt scale.
Lit.: Boratynki & Williams, 1964: 91; Danzig, 1975: 42; 1980: 218; Foldi, 2000 81;
2001: 305; Hodgson, 2005: 34; Hulden, 1985: 60; Kaydan et al., 2005; Kawecki, 1985:
30; Koteja, 1971: 322; Koteja & Zak-Ogaza, 1979: 674; 1981: 514; 1983: 476; Kozár et
al., 2013: 56; Kozár & Ostafitsuk, 1987: 92; Kozár & Walter, 1985: 74; Köhler, 1998:
381; Marotta, 1993: 159; Milonas et al., 2008: 143-147; Ouvrard & Kozár, 2009: 102;
Pellizzari & Kozár, 2011: 66; Tang & Hao, 1995: 451; Tereznikova, 1981: 32; Williams,
1985a: 374 (Miller et al., 2013).
Eriococcus munroi; Williams, 1985a: 374. Change of combination.

Acanthococcus munroi; Kosztarab & Kozár, 1988: 277. Revived combination.
Eriococcus munroi; Miller & Gimpel, 2000: 275. Revived combination.
Acanthococcus munroi; Kozár, 2009: 93. Revived combination.
Eriococcus tounetae; Goux, 1995: 47-48. Synonym nova
Holotype: France (Artigues, Hautes-Pyrenees), by L. Goux, 12 July 1953, under
rocks (MNHN 14524). Deposited in MNHN.
Lit.: Foldi, 2001: 305; Miller & Gimpel, 2000: 365 (Miller et al., 2013).
Acanthococcus tounetae; (Goux, 1995). Kozár, 2009: 94. Change of combination. Notes:
This species very similar to R. munroi, the only difference found is the smaller number of
setae on posterior dorsal marginal segments, because of this A. tounetae (Goux, 1995) is
here considered as synonym of R. munroi (Boratynski, 1962).
Description
Unmounted female
Body elongate, anterior end broadly rounded, orange, yellow, and dorsally convex.
Ovisac completely encloses the female and is elongate oval, broadly rounded anteriorly,
tapering posteriorly, with small transverse oval openning at posterior end; 1.8–2.5 mm
long, 1.2–1.5 mm wide, and 0.5–0.8 mm high; closely felted with short glassy spine like
threads projecting on dorsum and margin; white to cream at first and later a dirty cream
color. Male puparium elongate, almost parallel-sided with both end rounded; 1.0–1.3 mm
long, about 0.5 mm wide; white, loosely felted, with the glassy threads longer and curled;
at later stages with horizontal split at posterior end through which tips of wings and
abdominal waxy tassels of adult male protrude (Boratynski, 1962).
Mounted female
Body elongate anterior end broadly oval, tapering posteriorly, 1.5–2.1 mm long, 0.87–
1.29 mm wide. Antennae 7 (sometimes 6) segmented, 255–269 µm, length of segments:
I: 31.5–38.5, II: 31.5–38.5, III: 38.5–59.5, IV: 45.5–59.5, V: 21.0–26.5, VI: 19.0–24.5,
and VII: 31.5–42.0 in µm long; each segment (except third) covered with a few, strong
hair-like setae; apical segment with apical seta; apical segment also with 3 sensory falcate
setae; segment VI with one sensory falcate seta, segment V with one sensory falcate.
Frontal lobe present. Eyes situated on venter near margin. Anal lobes conical, each with
3 enlarged setae, shape and relative sizes of which are constant, sharply pointed, 28–56
(other the longest) µm long on dorsal surface; apical seta 231–303 µm long; ventral hair-
like subapical seta 52.5–66.5 µm long, in addition on the ventral side of the anal lobe with
three hair-like setae, each 35.0–48.5 µm long.
Venter: Labium 3 segmented; 120 µm long, shorter than clypeolabral shield, basal
segment with 2 pairs of setae; stylet loop slightly longer than labium. Legs well developed;
hind trochanter+femur 220–230 µm long, hind tibia 130–150 µm long, hind tarsus 140–
150 µm long, the tibia+tarsus always longer than trochanter+femur, claw curved, 30
µm long with denticle near apex. Posterior tibia with some pores on upper side. Hind
coxa with 23-30 conspicuous translucesnt pores and some spinulae. Tarsal digitules 44,
claw digitules 30.0 µm long; tarsal and claw digitules knobbed. Tibiae each with 5 setae
(median seta present), tarsi each with 5 setae. Spiracles with stout bar supporting small
opening; no definite groups of pores associated with spiracular opening. Derm with
normal slender setae in median areas and a few enlarged setae, each 26–74 µm long, on
margins and submargins. Discodial pores 4 µm in diameter and with 5 loculi, numerous, in
bands across abdominal segments but not reaching margins, present also in median areas
of thorax and around the spiracles. Macrotubular ducts of two sizes; a larger type, smilar
to dorsal duct, around margins and a narrower duct in transverse rows on abdominal
segments, in median and submarginal areas of thorax and head. Microtubular ducts few,
around margin only. Cruciform pores not numerous in bands across abdominal segments
but not reaching margins, present also in median areas of thorax and around spiracles.
Dorsum: Marginal enlarged setae, conical with almost straight sides and with blunt tips,
arranged in transverse bands across the segments but absent in submarginal areas; in
various sizes, 30–60 µm long, but always with one or two large setae present in any
marginal group, larger size setae forming 3 longitudinal lines on each side of anterior
abdomen. On the median areas of the fourth and fifth abdominal segments the enlarged
setae tend to be shorter but the normal enlarged setae are replaced on the median areas
of sixth and seventh abdominal segments by minute enlarged setae about 6 µm long and
rarely longer than the diameter of a setal base, there being usually 4 across the middle
of each segment. Macrotubular ducts of one size, each about 20 µm long with almost
straight sides and with cup narrower than a setal base of an enlarged seta, fairly numerous
in wide bands across the segments. Microtubular ducts with 2 sclerotized areas, about 4
µm long with inner end of tube swollen, ampulla and saucer-shaped inner collar, evenly
distributed. Anal ring oval, sclerotized, with pores and 8 setae, each seta 90–124 µm long.
Cauda present (after Williams (1985a) with modifications).
Acanthococcidae 515
Figure 181. Rhizococcus munroi (Boratynski, 1962), female. After Boratynski (1962) with
modifications.
Other stages
Male are described by Hodgson (2005). First instar nymph shown by Schmutterer (1952)
under name of R. devoniensis probably belongs to this species.
Ecology
Host plant: Achillea distans, A. millefolium, Artemisia sp., Brachypodium pinnatum, Calluna sp.,
Chrysanthemum sp., Conyza canadensis, Crepis sp., Deschampsia sp., Euphorbia sp., Festuca sp.,
F. pratensis, F. sulcata, Fragaria sp., Helianthemum sp., Hieracium sp., Hypochoeris sp., Leontodon
crispus, Minuartia sp., M. anatolica, Potentilla sp., P. megalantha, Sarcopoterium spinosum, Saussurea
sp., S. japonica, Sedum sp., Tetragonolobus sp., Teucrium sp., Thymus sp., T. quinquecostatus var.
przewalskii, Valeriana sp., V. officinalis, Veronica sp., Vicia sp.
Distribution: Bulgaria, France, Germany (?, under the name of R. devoniensis, det. by
Schmutterer), Greece, Hungary, Italy, Kazakhstan, Moldova, Poland, Romania, Russia,
Turkey, Ukraine, United Kingdom.
Biology: This species has both male and females and has two generations per year.
The crawlers of the first generation appear in April and the adults in June. The eggs are
deposited by the females at the end of June, hatch in the middle of July, and the adults
of the second generation appear in September. The eggs are deposited by the females of
the second generation in September and overwinter in ovisacs and hatch in April of the
following year. The active stages feed on leaves, stems and at root crown. For pupation
and oviposition the insects migrate to the lower parts of the host plants and settle on half
dry curled up leaves or in axils (Boratynski, 1962).
Comments
The drawings of female, and first-instar nymph shown by Schmutterer (1952: 68) under
name of R. devoniensis, seems belonging to R. munroi.
Acanthococcidae 517
Rhizococcus nedimi Kaydan Species nova (Fig. 182)
Holotype: Female. Turkey (Erzincan-Ağıl, N: 39° 58’ 342’’, E: 039° 28’ 163’’, 1633
m altitude), on Euphorbia sp., M.B. Kaydan, F. Kozár, 07.vi.2010 (KPCT: 4744).
Paratypes: 3 adult females, same data as holotype, (KPCT: 4744); 2 adult females,
Turkey, Erzurum-Pasinler-Baldizi, N: 40°02’200’’, E: 041°32’510’’, 1900 m altitude,
on Scabiosa sp., M.B. Kaydan, F. Kozár, 09.vi.2010 (KPCT: 4810). Deposited in
Kaydan’s personal collection, Turkey.
Common name: Nedim’s felt scale.
Description
Unmounted female
Adult females redish; found on rootcrown of host plant, felt-like ovisac cream-white.
Mounted female
250–290 µm, length of segments: I: 45–65, II: 37.5–45.0, III: 52.5–62.5, IV: 45.0–62.5,
V: 17.5–25.0, VI: 17.5–20.0 and VII: 30.0–42.5 µm long, each segment covered with a
few, strong hair-like setae (except third segment); apical segment with apical seta 45–49
µm long; apical segment also with 3 sensory falcate setae, each 25–32.5 µm long; segment
VI with 1 sensory falcate seta 30.5–37.5 µm long, segment V with 1 sensory falcate seta
17.5–20 µm long. Frontal lobes and frontal tubercle present. Eyes situated on venter near
margin. Anal lobes strongly developed, each with 3 enlarged setae, each 32.5–67.5 µm
plus 4–6 microtubular ducts on dorsal surface; apical seta 250–280 µm; ventral hair-like
subapical seta 70–95 µm long.
Venter: Labium 3 segmented, 135.0–152.5 µm long, 80.0–87.5 µm wide, median setae
on apex of labium short, 12.5–15.0 µm long. Legs well developed; lengths of segments
and digitules of prothoracic legs; coxa: 72.5–85, trochanter: 55.0–62.5, femur: 117.5–
137.5, tibia: 105–120, tarsus: 95–120; claw: 30–32.5, trochanther + femur: 170–195, tibia
+ tarsus: 230–235, tarsal digitules: 42.5–45; claw digitules: 32.5–35.0 µm long; lengths of
segments and digitules of mesothoracic legs; coxa: 77.5–95.0, trochanter: 55–60, femur:
125–135, tibia:110–130, tarsus: 110.0–122.5; claw: 32.5–35.0, trochanther + femur:
182.5–185.0, tibia + tarsus: 215–245, tarsal digitules: 45.0–52.5, claw digitules: 30.0–37.5
µm long; lengths of segments and digitules of metathoracic legs; coxa: 95.0–110.0,
trochanter: 62.5–70.0, femur: 135–155, tibia: 115–135, tarsus: 112.5–132.5 and claw:
31–35, trochanther + femur: 195–215, tibia + tarsus: 230–260, tarsal digitules: 50, claw
digitules: 30.0–32.5 µm long. Tarsal and claw digitules knobbed. Meso- and metathoracic
coxae each with spinulae on ventral surface and translucent pores on metathoracic coxa
(13-44). Tibiae each with 5 setae (median seta present), tarsi each with 8 setae. Length of
spiracles 55.0–62.5 µm; diameter of spiracular peritreme 27.5–32.5 µm, posterior spiracles
slightly larger than anterior. Setae on venter two kind, flagellate setae, each 37–65 µm
long, situated on medial part of the body, spine like setae, each 11–20 µm long scattered
on submedian and submarginal area. Enlarged setae, situated on submargin in a a sparse
band, each 12.5–30.0 µm long. Multilocular pores each 4.0–7.5 µm in diameter and with
3 or 5 loculi, distributed in sparse bands on all abdominal segments and scattered on
thoracic segments especially around spiracles. Macrotubular ducts of two size, smaller
macrotubular ducts each 3–4 µm wide and 15.0–27.5 µm long, present on submedian
of abdominal segments I–VI, thorax and head, larger macrotubular ducts each 5.0–10.0
µm wide and 20.0–27.5 µm long, sparse in band on last abdominal segments, and on
submarginal band on abdomen, thorax and head. Microtubular ducts sclerotized, with
oval orifice, present on all venter, 5.0–7.0 µm long; cruciform pores scattered on thorax
and hand on generally on the submedian and median part, each 5.0–6.0 µm in diameter.
Dorsum: Marginal dorsal setae spine-like each 37.5–47.5 µm forming a marginal row
and differentiated from dorsal setae, but on last abdominal segments with three (two
long and one short) truncated setae, 20.0–47.5 µm; other dorsal setae conical, spine-like,
various sizes, seta 12.5–40.0 µm long, those on thorax, head and anterior abdominal
segments larger (each 7.5–12.5 µm long) on median part of posterior 3 or 4 abdominal
segments; arranged in transverse rows across each body segment, rows irregular on head.
Macrotubular ducts, each 7–9 µm wide and 20.0–27.5 µm long, scattered throughout
oval dermal orifice, scattered over dorsum. Anal ring strongly sclerotized, with 14-17
pores on each inner side, 62.5–77.5 µm in diameter, with 8 setae, each 80–120 µm long;
anal ring situated on margin of dorsum. Cauda present.
Etymology
The new species is named after Good Nedim Salman, who helped collecting the scale
samples for this study.
Ecology
Host plant: Euphorbia sp., Scabiosa sp.
Acanthococcidae 519
Figure 182. Rhizococcus nedimi Kaydan & Kozár sp. n., female.
Rhizococcus ningxianensis (Tang & Hao, 1995) (Fig. 183) Combination

nova
Eriococcus ningxianensis Tang & Hao, 1995: 481, 594.
Holotype: Female. China (Ningxia), ?.v.1993 (1992?). By original designation.

Deposited in Shanxi.
Lit.: Tao, 1999: 32; Wang, 2001: 209 (Miller et al., 2013).
Acanthococcus ningxianensis; Miller & Gimpel, 1996: 602. Change of combination.

Eriococcus ningxianensis; Miller & Gimpel, 2000: 280. Revived combination.
Acanthococcus ningxianensis; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Unknown.
Mounted female
Body ovoid, 2.5 mm long, 1.6 mm wide. Antennae 7 segmented, length of segments:
I: 25, II: 30, III: 40, IV: 39, V: 15, VI: 15 and VI: 30 µm long; each segment (except
third) covered with a few, strong hair-like setae. Frontal tubercle present. Eyes situated on
venter near margin. Anal lobes developed, sclerotised, each with 3 enlarged setae, slightly
truncated; ventral with long apical seta and three hair-like seta.
Venter: Labium 3 segmented. Stylet loop three times longer than labium. Legs well
developed, small. Tibia shorter than tarsus; claw denticle distinct; digitules of tarsus and
claw slender, knobbed and surpassing the tip of claw; hindcoxa with translucent pores
in high numbers. Posterior spiracles slightly larger than anterior. Derm with a sparse
covering of scattered flagellate hair-like setae. Enlarged setae, situated on submargin in
row. Discodial pores with 3 or 5 loculi, pores distributed in sparse bands on all abdominal
and thoracic segments and a few on head. Macrotubular ducts of two sizes, larger ones
on submarginal area; narrower ones present on medial area. Microtubular ducts in the
marginal band. Cruciform pores few, distributed on submargin and median part of
thorax and head and submargin of first abdominal segments.
Dorsum: Marginal enlarged setae blunted, in 3 sizes; there are 2 spines on abdominal
segment VIII, segments anterior to it scattered in segmental transverse rows and
bands. Macrotubular ducts, scattered throughout dorsum, generally in segmental bands.
Microtubular ducts short, with 2 sclerotized areas, scattered over dorsum. Anal ring
sclerotized, with 8 setae (Tang & Hao, 1995).
Acanthococcidae 521
Figure 183. Rhizococcus ningxianensis Tang & Hao 1995, female. After Tang & Hao (1995)
with modifications.
Ecology
Host plant: Unknown.
Rhizococcus palustris Dziedzicka & Koteja, 1971 (Fig. 184)

Rhizococcus palustris Dzeidzicka & Koteja, 1971: 573.
Holotype: Female. Poland (Nowy Targ, Piekeilnik, in the Puscizna Wielka peat-bog),
on Eriophorum vaginatum and Carex sp., 10 and 28.viii.1963, by E. Koteja. By original
designation. Deposited in Crac Notes: Paratypes in BMNH and ZMAS.
Common name: Polish felt scale.
Lit.: Dziedzicka, 1977: 59, 71; Koteja, 1974b: 76; Koteja & Zag-Ogaza1981: 514;
Kozár & Walter, 1985: 75; Köhler, 1998 : 400; Lagowska, 2002: 243; Miller & Gimpel,
1996: 600; Koteja in Nast et al., 1990: 120; Williams, 1985a: 239 (Miller et al., 2013).
Acanthococcus palustris; Kosztarab & Kozár, 1978: 68. Change of combination.

Eriococcus podhalensis; Dziedzicka & Koteja, 1985: 31 in Kawecki, 1985: 31. Change of
combination and replacement name for Eriococcus palustris (Dzeidzicka & Koteja 1971).
Rhizococcus palustris; Kosztarab & Kozár, 1988: 304. Revived combination.
Acanthococcus dziedzickae; Miller & Gimpel, 1996: 605. Unjustified replacement name for
Rhizococcus palustris Dzeidzicka & Koteja 1971. Notes: Miller & Gimpel (1996) proposed
the replacement name Acanthococcus dziedzickae for the homonym Acanthococcus palustris
(Dodds, 1923) not realizing that Dziedzicka & Koteja (1985) had already put forth the
replacement name Eriococcus podhalensis.
Rhizococcus palustris; Tang & Hao, 1995: 250. Revived combination.
Eriococcus podhalensis; Miller & Gimpel, 2000: 303. Revived combination.
Rhizococcus palustris; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Ovisac is white or cream-colored and encloses female completely. Adult female elongate-
oval, tapering posteriorly.
Mounted female
Body elongate oval, 2.1–2.3 mm long, 1.2–1.3 mm wide. Antennae 7 segmented, 250–270
µm, length of segments: I: 40, II: 38-47, III: 47–52, IV: 42–57, V: 9–24, VI: 19 and VII:
33–38 µm long, each segment covered with a few, strong hair-like setae (except third
segment); apical segment with apical seta; apical segment also with 3 sensory falcate setae;
segment VI and V with 1 sensory falcate seta. Frontal lobes present. Eyes situated on
Acanthococcidae 523
Figure 184. Rhizococcus palustris Dziedzicka & Koteja, 1971, female. After Dziedzicka &
Koteja (1971) with modifications.
venter near margin. Anal lobes conical, partly sclerotised, 85–100 µm long, 70 µm wide,
each with 3 enlarged setae, 23–38 µm (inner side with one longer one shorter enlarged
setae) plus 2 microtubular ducts on dorsal surface; apical seta 260–280 µm long; ventral
surface with 3 hair-like setae, each 70–95 µm long and with some quinquelocular pores.
Venter: Labium 3 segmented, 100–120 µm long, 85–95 µm wide. Legs well developed;
posterior pair 570–760 µm long, anterior pair slightly shorter; hindcoxa: 95, trochanther
+ femur: 190–240, tibia: 123–150, tarsus 128–140, claw 24–33 µm long, tarsal digitules
42–50, claw digitules 28–38 µm long; tarsal and claw digitules knobbed. Metathoracic
coxae with 7-13 translucent pores. Tibiae each with 5 setae (median seta present), tarsi
each with 6 or 7 setae. Posterior spiracles slightly larger than anterior; no definite pores
associated with spiracular openning. Setae on venter unmodifiated, hair-like, 10–60 µm
long, forms groups on cephalothorax and transverse on abdomen. Enlarged setae, one to
three rows on submarginal area. Multilocular pores each 5.0–7.0 µm in diameter and with
3-7 loculi, very numerous on posterior abdominal segments, towards the anterior end
of the body becoming scattered. Macrotubular ducts of two sizes, smaller macrotubular
ducts each 5.0 µm wide and 22–25 µm long; scattered, larger macrotubular ducts each
10 µm wide and 22–25 µm long, on submarginal band on abdomen, thorax and head.
Microtubular ducts sclerotized, on margin of venter. Cruciform pores scattered on thorax
and head on generally on the submedian and median part, each 5.0–6.0 µm in diameter.
Dorsum: Marginal enlarged setae conical 15–33 µm long, on the anterior end of the body
sharply pointed, on the posteror end with blunt or truncate apex; forming one row, 2 or
3 situated on the very margin, one row dorsally; other dorsal setae conical, spine-like,
various sizes, those on thorax, head and anterior abdominal segments larger than setae on
median part of posterior abdominal segments; arranged in transverse rows across each
body segment, rows irregular on head. Macrotubular ducts, 10 µm wide and 22–25 µm
long,, scattered throughout dorsum, generally in segmental bands. Microtubular ducts,
each 2.0 µm long with oval dermal orifice, scattered over dorsum. Anal ring strongly
sclerotized, with 8 setae, each 100–150 µm long; anal ring situated on margin of dorsum
(after Dziedzicka & Koteja (1985); Kosztarab & Kozár, (1988) with modifications.
Ecology
Host plant: Carex sp., Eriophorum vaginatum, Luzula pilosa.
Distribution: Poland, Switzerland.
Biology: On the leaves. Females completed egg laying by August. Males not known.
Acanthococcidae 525
Rhizococcus puymorensis (Goux, 1992) (Fig. 185) Redescription,

Combination nova
Eriococcus puymorensis Goux, 1992: 42.
Holotype: Female. France (Pyrenees-Orientales, Col de Puymorens, vers 1700 m,

Ariège), 14.viii.1953, by L. Goux (MNHN 14505). Deposited in MNHN.
Common name: Goux’s felt scale.
Lit.: Foldi, 2001: 305; Ouvrard & Kozár, 2009: 102 (Miller et al., 2013).
Acanthococcus puymorensis; Miller & Gimpel, 1996: 603. Change of combination.

Eriococcus puymorensis; Miller & Gimpel, 2000: 310. Revived combination.
Acanthococcus puymorensis; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult females redish; found at root crown of host plant, ovisac cream-white
Mounted female
segments: I: 55, II: 46, III: 110, IV: 26, V: 27 and VI: 38 µm long; on apical segment
three sensory falcate setae are found; the segments of the antenna are covered with few
hair-like setae. Frontal lobes present. Eyes visible, situated on venter. Anal lobes slightly
sclerotized each with two spinose setae along inner margin, and one spinose seta on
outer margin, similar in size to those on dorsal abdominal margin, apical setae each 175
µm long.
Venter: Labium 3 segmented, 110 µm long; stylet loop reaching the coxae of median
legs. Legs well developed; lengths of segments and digitules of prothoracic legs; coxa:
62, trochanter: 62, femur: 115, tibia: 84, tarsus: 125 and claw: 31, tarsal digitules: 52, claw
digitules: 34 µm long; lengths of segments and digitules of mesothoracic legs; coxa: 67,
trochanter: 62, femur: 113, tibia: 96, tarsus: 134 and claw: 31, tarsal digitules: 52, claw
digitules: 35 µm long; lengths of segments and digitules of metathoracic legs; coxa: 71,
trochanter: 66, femur: 119, tibia: 100, tarsus: 144 and claw: 34, tarsal digitules: 54, claw
digitules: 36 µm long. Tarsal and claw digitules knobbed. Meso- and metathoracic coxae
each with spinulae on ventral surface. Tibiae each with 5 setae (median seta present).
The diameter of anterior spiracles 38 µm, posterior spiracles slightly larger than anterior.
Venter with a small number of scattered, hair-like setae, the median pair of setae very
long, 70 µm, as long as the width of the segment, and with a marginal and submarginal
row of enlarged setae. Multilocular pores each 4.0–5.0 µm in diameter and with 3, 5 or
7 loculi, distributed in sparse bands and rows on all segments of abdomen and thorax.
Macrotubular ducts about 4 µm wide and 20 µm long, scattered all body. Microtubular
ducts present in a submarginal band, 6 µm long. Cruciform pores 4 µm long, very few
scattered on submarginal band, and on middle of head, 4 µm in diameter.
Dorsum: Enlarged setae spine-like, on margin 43–50 µm long, varying in size and
shape, blunted. Other dorsal enlarged setae very few, forming sparse rows; in middle
part of posterior segments 7–8 µm long, on thorax they are somewhat longer, 10–34
µm. On margin of abdominal segments usually 2 longer spines present. Macrotubular
ducts similar to those on venter, sparse, present throughout, 6 µm wide, 16 µm long.
Microtubular ducts with bifid orifice, each 6 µm long, scattered throughout. Anal ring
conspicuous, 55 µm wide and 72 µm long, with 8 hair-like setae, 101 µm long. Cauda not
Ecology
Host plant: Undetermined.
Comments
Here we provide a new drawing based on the holotype. According to the slide and
description under the name of A. guesinus (Goux, 1992, Planche II), was belong to R.
puymorensis. Female on the slide similar to A. thymi (Schrank, 1801), but differs in several
details, such as longer hair-like setae on abdominal venter, presence of spinulae and
absence of pores on posterior coxae, presence of frontal lobes, and higher number
of longer spines on mid dorsum. The significance of these differences will be better
understood if more material will be found for examination.
Acanthococcidae 527
Figure 185. Rhizococcus puymorensis (Goux, 1992), female. Original.

Rhizococcus reynei (Schmutterer, 1952) (Figs. 186, 187 a, b, 188)

Redescription, Combination nova
Eriococcus reynei Schmutterer, 1952: 414.
Lectotype: Female. Germany (Franconia Basin, Bavaria, Alterlangen, on Thymus

serpyllum, 15. VI.1950. Designated here (Kozár’s coll. 7664). Deposited in DEI.
Common name: Schmutterer’s felt scale.
Lit.: Foldi, 2001: 305; Hoy, 1963: 112; Kaydan, et al., 2005: 400; Kozár, 1986: 172;
Kozár et al., 2013: 56; Köhler, 1998: 385; Ouvrard & Kozár, 2009: 102 (Miller et al.,
2013).
Eriococcus reynei; Lindinger, 1957: 548. incorrect synonymy with R. devoniensis

Acanthococcus reynei; Schmutterer, 1980: 50. Change of combination.
Eriococcus reynei; Lindinger, 1957: 548. incorrect synonymy with A. thymi .
Eriococcus reynei; Miller & Gimpel, 2000: 314. Revived combination.
Acanthococcus reynei; Kozár, 2009: 93. Revived combination.
Eriococcus tavignani; Goux, 1991: 46. (Fig. 188) Synonym nova.
Holotype: Female. France (Haute-Corse, Corte, Tavignano), on Euphorbia characias,
23/08/1931, by L. Goux (MNHN 14516-01). Notes: Paratypes: 8 slides from the
same collection as holotype, some of them contains first and second instar nymph.
Deposited in MNHN.
Acanthococcus tavignani; Miller & Gimpel, 1996: 604. Change of combination.
Eriococcus tavignani; Miller & Gimpel, 2000: 353. Revived combination.
Acanthococcus tavignani; Kozár, 2009: 94. Revived combination.
Rhizococcus tavignani; Pellizzari & Kozár, 2011: 66. Change of combination.
Description
Unmounted female
Unknown.
Mounted female
segments: I: 36, II: 34, III: 43, IV: 38, V: 24, VI: 22 and VII: 31 µm long; each segment
covered with a few, strong hair-like setae (except third segment); apical segment with
long; segment VI with 1 sensory falcate seta broken, segment V with 1 sensory falcate
seta 12 µm long. Frontal lobes present. Eyes visible, situated on venter. Anal lobes slightly
outer margin, similar in size to those on dorsal abdominal margin, apical seta broken.
Venter: Labium 3 segmented, 89 µm long. Stylet loop almost reaching posterior coxae.
Legs well developed; length of segments and digitules of prothoracic legs: 65, trochanter:
Acanthococcidae 529
Figure 186. Rhizococcus reynei (Schmutterer, 1952), female, first instar on top left.
Original.
46, femur: 108, tibia: 96, tarsus: 103, claw: 26, tarsal digitules: 36, claw digitules: 26 µm
long; length of segments and digitules of mesothoracic legs; coxa: 65, trochanter: 46,
femur: 113, tibia: 101, tarsus: 115, claw: 29, tarsal digitules: 34, claw digitules: 26 µm
long; length of segments and digitules of metathoracic legs; coxa: 74, trochanter: 48,
femur: 122, tibia: 110, tarsus: 127, claw: 31, tarsal digitules: 38, claw digitules: 29 µm
long. Tarsal and claw digitules knobbed, slightly longer than claw; claw with denticle at
apex. Meso- and metathoracic coxae each without spinulae on ventral surface, translucent
pores on dorsal surface. Tibiae each with 5 setae (median seta present). The diameter
of anterior spiracles 28 µm, posterior spiracles slightly larger than anterior. Venter with
scattered, hair-like setae and with a marginal and submarginal row of enlarged setae.
Quinquelocular pores each 5.0 µm in diameter, distributed in sparse bands and rows on
all segments of abdomen and thorax. Macrotubular ducts of two sizes, larger ones on
marginal area, about 4.0 µm wide, narrower ones about 7.0 µm wide, scattered all body.
Microtubular duct not seen. Cruciform pores very few, scattered on submarginal band,
and on middle of head, each 5.0 µm in diameter.
Dorsum: Enlarged setae spine-like, on margin 34–41 µm long, varying in size and shape,
blunted, with two spines on margin of segments. Other dorsal enlarged setae, forming
sparse rows; in middle part of posterior segments 19 µm long, on thorax they are
somewhat longer; submarginal band without enlarged setae; on margin of abdominal
segments usually 3 longer enlarged setae present. Macrotubular ducts similar to those
on venter, sparse, present throughout, 7.0 µm wide. Microtubular ducts with 6.0 µm
long, scattered throughout. Discodial pores absent. Anal ring, 48 µm wide. Cauda present
(Redescription is based on lectotype).
Other stages
First instar nymph (Fig. 186, top left)

Body oval. Antennae 6 segmented, segments with a few hair-like setae; apical segment
also with 3 sensory falcate setae, two preapical segments also with 1 sensory falcate seta.
Frontal lobe absent, frontal tubercle present. Eyes situated on venter near margin.
apical segment with 5 pairs of equal length hair-like setae, and 1 pair short apical setae.
Stylet loop long, reaching behind posterior coxae. Legs normal: tarsal and claw digitules
slightly knobbed, longer than claw. Claw with a denticle. Legs with a few hair-like
setae; tibia with 5 setae. Multilocular pores with 3-5 loculi, mostly with 5 loculi; in two
longitudinal rows on venter. Microtubular ducts absent on venter. Cruciform pores; a
few in marginal area of thorax. A few hair-like setae present on submedian venter, short
spine-like setae on submargin. Suranal setae hair-like.
Dorsum: Enlarged setae blunted, conical; in a row on margin, one on margin of each
segments, and a group on head margin, shorter setae in four longitudinal rows on
dorsum. Microtubular ducts scattered, in four longitudinal rows. Anal ring situated on
margin of dorsum; oval, sclerotized, with one row of pores and with 6 setae. Anal lobes
Acanthococcidae 531
well developed, membranouos, with 3 blunted conical spines, ventrally with two flagellate
setae; and a long apical seta. Cauda wide, plate-like.
Other stages
Second instar female nymph (Fig. 187 a)

Body oval, Antennae 6 segmented, segments with a few hair-like setae; apical segment
Stylet loop not seen. Legs normal: tarsal and claw digitules slightly knobbed, longer than
claw. Claw with a denticle. Legs with a few hair-like setae; tibia with 5 setae. Median and
posterior coxae with spinulae. Multilocular pores with 3 or 5 loculi, mostly with 5 loculi;
scattered al over venter. Microtubular ducts absent on venter. Cruciform pores; a few in
marginal area of thorax. A few hair-like setae present on submedian venter, short spine-
like setae on submargin. Suranal setae hair-like.
Figure 187. Rhizococcus reynei (Schmutterer, 1952), second instar, a female, b male.
Original.
Dorsum: Enlarged setae blunted, conical; in a row on margin, two on margin of each
segments, and a group on head margin, shorter setae in 6-8 longitudinal rows on dorsum.
Microtubular ducts scattered. Anal ring situated on margin of venter; oval, sclerotized,
with one row of pores and with 6 setae. Anal lobes well developed, membranouos, with
3 blunted conical spines, ventrally with two flagellate setae; and a long apical seta. Cauda
wide, plate-like.
Second-instar male nymph (Fig. 187 b)

Body oval, Antennae 7 segmented, segments with a few hair-like setae; apical segment
Stylet loop not seen. Legs normal: tarsal and claw digitules slightly knobbed, longer than
claw. Claw with a denticle. Legs with a few hair-like setae; tibia with 5 setae. Median and
posterior coxae with spinulae. Multilocular pores with 3 or 5 loculi, mostly with 5 loculi;
scattered al over venter. Macrotubular ducts scattered all over venter. Microtubular ducts
absent. Cruciform pores; a few in marginal area of thorax and abdomen. A few hair-like
setae present on submedian venter, short spine-like setae on submargin. Suranal setae
hair-like.
Dorsum: Enlarged setae blunted, conical; in a row on margin, two on margin of each
segment, and a group on head margin, shorter setae in 6-8 longitudinal rows on dorsum.
Macrotubular and microtubular ducts scattered all over dorsum. Anal ring situated on
margin of venter; oval, sclerotized, with one row of pores and with 6 setae. Anal lobes
well developed, membranouos, with 3 blunted conical spines, ventrally with two flagellate
setae; and a long apical seta. Cauda wide, plate-like.
Ecology
Host plant: Euphorbia characias, Thymus serpyllum.
Distribution: Bulgaria, China, France, Germany, Greece, Hungary, Italy, Iran, Moldova,
Turkey.
Comments
Here we consider R. tavignani as a synonym of R. reynei, as we could not find enough
differences at this stage in order to treat them as separate species. However, the difference
in host plants (Euphorbia sp. and Thymus sp.) deserves more attention.
Acanthococcidae 533
Figure 188. Rhizococcus reynei (=tavignani) (Schmutterer, 1952), female. Original.

Rhizococcus rugosus (Wang, 1982) (Fig. 189) Revived combination

Eriococcus rugosus Wang, 1982a: 441.
Holotype: Female. China (Zhejiang Province, Yuhang Xian), on Phyllostachys pubescens,

28.iii.1981, by Xu Tian-shen. By original designation. Deposited in Beijing.
Common name: Wang’s felt scale.
Lit.: Fang et al., 2001; Hua, 2000: 138; Kozár et al., 2013: 56; Tang & Hao, 1995: 520;
Tao, 1999: 35; Wang, 2001: 225. Wang, et al., 1998: 81, 186 (Miller et al., 2013).
Rhizococcus rugosus; Kozár & Walter, 1985: 75. Change of combination.

Acanthococcus wangi; Miller & Gimpel, 1996: 605. Homonym of Eriococcus rugosus Froggatt
1933; discovered by Miller & Gimpel, 1996: 605. Change of combination and replacement
name for Eriococcus rugosus Wang 1982a.
Rhizococcus rugosus; Köhler, 1998: 401. Revived combination.
Eriococcus wangi; Miller & Gimpel, 2000: 375. Change of combination.
Acanthococcus rugosus; Kozár, 2009: 101. Reestablishment of Acanthococcus wangi Miller &
Gimpel, 1996: 605. Revived combination.
Description
Unmounted female
Adult female is broadly oval.
Mounted female
Body elongate oval, 3.5 mm long. Antennae short, 6 segmented. Eyes visible, situated on
venter. Anal lobes small, strongly sclerotized with two enlarged setae along inner margin,
and one enlarged seta on outer margin, bearing small sclerotised bar with one enlarged
setae.
Venter: Labium 3 segmented. Legs well developed; short, claw with a well-developed
denticle. The spiracles strongly sclerotised, associated with 12-16 quinquelocular pores at
opening. Venter with small number of scattered hair-like setae and with a marginal and
submarginal row of enlarged setae. Quinquelocular pores, distributed in sparse bands
and rows on all segments of abdomen and a few on thorax. Macrotubular ducts of one
size scattered over entire body. Cruciform pores and microtubular ducts not mentioned.
Dorsum: Enlarged setae spine-like, sharply pointed, on margin distinctly larger than other
dorsal setae; marginal row on the abdominal segments II-VII carries 2-4 setae. Other
dorsal enlarged setae much shorter, forming sparse rows. Macrotubular ducts similar to
those on venter, sparse, present throughout. Microtubular ducts absent. Anal ring with 8
long setae. Cauda well developed (Wang (1982a) with modifications).
Acanthococcidae 535
Figure 189. Rhizococcus rugosus (Wang, 1982), female. After Wang (1982) with
modifications.
Ecology
Host plant: Artemisia sp., Phyllostachys pubescens.
Distribution: China, Hungary.
Rhizococcus sachalinensis (Siraiwa, 1939) (Fig. 190) Combination nova

Eriococcus sachalinensis Siraiwa, 1939: 65.
Syntype: Female. Russia (Saghalien (Sakhalin) Island, Hoye, on undetermined grass,

?.viii.1935, by H. Siraiwa. Type designation unknown. Notes: Siraiwa types are
apparently lost (Ben-Dov, 1994).
Lit.: Borchsenius, 1949: 59; Köhler, 1998: 382; Kozár & Walter, 1985: 74; Miller &
Gimpel, 1996: 604 (Miller et al., 2013).
Acanthococcus sachalinensis; Borchsenius, 1949: 351. Change of combination.

Eriococcus sachalinensis; Hoy, 1963: 114. Revived combination.
Acanthococcus sachalinensis; Danzig, 1980: 207. Revived combination.
Eriococcus sachalinensis; Miller & Gimpel, 2000: 323. Revived combination.
Acanthococcus sachalinensis; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Sac of female is elongate, 2.6 mm long, 1.4 mm wide, moderately convex, closely felted,
white or greyish, but changing to ochreous or pale straw with age. Adult female pale
greenish yellow.
Mounted female
Body elongate oval, 2.0–2.4 mm long, 1.0–1.4 wide. Antennae short, 6 or 7 segmented; I:
42, II: 40, III: 110, IV: 29, V: 27 and VI: 39 µm long. Eyes visible, situated on venter. Anal
lobe small, sclerotized with two enlarged setae along inner margin, and one enlarged seta
on outer margin, almost equal long.
Venter: Labium large, 3 segmented. Legs well developed; long, claw with well-dveloped
denticle, tarsal and claw digitules knobbed, longer than claw. Posterior coxae with large
pores, tibia with 5 setae, tarsus longer than tibia. Venter with scattered, in small number
hair-like setae and with a marginal and submarginal row of enlarged setae. Quinquelocular
pores, distributed in sparse bands and rows on all segments of abdomen. Macrotubular
ducts of one size, scattered all body.
Dorsum: Enlarged setae spine-like apices acute, on margin distinctly larger than other
dorsal setae; marginal row on the abdominal segment II–VII carries two longer setae
and 2-4 shorter setae. Other dorsal enlarged setae forming 3 longitudinal lines and
Acanthococcidae 537
sparse transverse rows. Macrotubular ducts similar to those on venter, sparse, present
throughout. Microtubular ducts not mentioned. Anal ring, with 8 long setae. Cauda
present (after Siraiwa (1939), with modifications).
Ecology
Comments
Danzig (1986) states that R. sachalinensis is probably a synonym of R. greeni. According to
the drawing of Siraiwa (1939) the shape, size and the number of enlarged setae differs.
For a final decision on their identity, a study based on new material would be necessary.
Figure 190. Rhizococcus sachalinensis (Siraiwa 1939), female. After Shiraiva (1939) with
modifications.
Rhizococcus saxatilis (Kiritchenko, 1940) (Fig. 191) Combination nova

Eriococcus saxatilis Kiritchenko, 1940: 126.
Lectotype: Ukraine (Crimea, Kekeneis, Opolzmevoe, Limeni), on Teucrium chamaedrys.

By subsequent designation Danzig, 1996: 522. Deposited in ZMAS. Notes: Three
paralectotype females on same slide as lectotype and 10 additional paralectotypes on
two slides in ZMAS.
Lit.: Borchsenius, 1937: 184; Danzig, 1996: 575; Hadzibejli, 1983: 269; Köhler, 1998:
393; Lindinger, 1957: 544; Matesova, 1957: 169; Matesova, 1968: 113; Tang & Hao,
1995: 452; Tereznikova, 1967: 475; Tereznikova, 1968: 49; Tereznikova, 1975: 29;
Tereznikova, 1981: 38; Ter-Grigorian, 1983: 880.
Acanthococcus saxatilis; Borchsenius, 1949: 52. Change of combination.

Nidularia saxatilis; Lindinger, 1957: 544. Change of combination.
Eriococcus saxatilis; Hoy, 1963: 115. Revived combination.
Acanthococcus saxatilis; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus saxatilis; Miller & Gimpel, 2000: 329. Revived combination.
Acanthococcus saxatilis; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Ovisac is very dense, pale ochreous, relatively large, 2.3 long, 1.8 wide. Body greenish
grey. Adult female is covered with a fine layer of white mealy secretion, 1.8 long, 0.8 wide
(Kiritchenko, 1940).
Mounted female
Body elongate oval. Antennae short, 7 segmented, I: 33.2, II: 33.5, III: 46.3, IV: 34.5,
V: 20.8, VI: 18.1 and VII: 38.1 µm long. Eyes visible, situated on venter. Frontal lobes
present. Anal lobes small, strongly sclerotized with two enlarged setae along inner
margin, and one enlarged seta on outer margin, 45–60 µm long, apical setae 300 µm long,
subapical setae 75–95, suranal setae 95 µm long.
Venter: Labium 3 segmented. Stylet loop reaching median coxae. Legs well developed;
femur: 155, tibia: 125, tarsus: 128 µm long, claw with strong denticle, tarsal and claw
digitules longer than claw. Median and posterior coxae with spinulae, and posterior coxae
with large pores. The spiracles strongly sclerotised, associated with 6-10 quinquelocular
pores at openning. Venter with scattered, in small number hair-like setae, up to 60 µm
long and with a submarginal row of enlarged setae. Multilocular pores with 5-7 loculi,
distributed in sparse bands and rows on all segments of abdomen and a few on thorax.
Macrotubular ducts of one size, 22 µm long and 8 µm wide scattered all body. Cruciform
pores in a marginal row.
Acanthococcidae 539
Figure 191. Rhizococcus saxatilis (Kiritchenko, 1940), female. After Tereznikova (1981)
with modifications.
Dorsum: Enlarged setae spine-like, blunted, 48–62 µm long, on margin distinctly larger
than other dorsal setae; marginal band on abdominal segment II-VII carries on margin
3 or 4 setae. Other dorsal enlarged setae, forming sparse rows. Macro- and microtubular
ducts sparse, present throughout. Anal ring with partly double rows of pores, with 8,
each 110 µm long setae. Cauda present (after Kiritchenko (1941); Borchsenius (1949)
with modifications).
Ecology
Host plant: Ajuga sp., Artemisia frigida, A. marschalliana, A. vulgaris, Centaurea sibirica,
Dianthus capitatus, Euphorbia sp., Fragaria sp., Mentha sp., Plantago sp., Hylotelephium telephium,
Potentilla anserina, Teucrium chamaedrys, Ziziphora clinopodioides.
Distribution: Armenia, Azerbaijan, Kazakhstan, Turkey, Ukraine.
Biology: On the roots and root crowns.
Rhizococcus saxidesertus (Borchsenius, 1949) (Fig. 192) Redescription,

Combination nova
Acanthococcus saxidesertus Borchsenius, 1949: 343.
Lectotype: Female. Tajikistan (Gissar Ridge, near Ziddy, on stones), 17.vii.1944, by

N. Borchsenius. By subsequent designation Danzig, 1996: 522. Deposited in ZMAS
(type no. 4-44). Notes: 2 female paralectotypes on 1 slide with same data as lectotype.
Original description gives host as Asplenium ceterach.
Lit.: Bazarov, 1962: 63; 1963: 67; Borchsenius, 1950: 120; Köhler, 1998: 383; Matesova,
Nidularia saxidesertus; Lindinger, 1957: 543. Change of combination.

Eriococcus saxidesertus; Hoy, 1963: 115. Change of combination.
Acanthococcus saxidesertus; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus saxidesertus; Miller & Gimpel, 2000: 330. Revived combination.
Acanthococcus saxidesertus; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female is egg-shaped, 2.2 mm long, 1.2 mm wide. Ovisac is grayish, compact,
entirely enclosing the body.
Mounted female
segments: I: 48–56, II: 36–43, III: 72–80, IV: 60, V: 29, VI: 29 and VII: 48 µm long; each
Acanthococcidae 541
Figure 192. Rhizococcus saxidesertus (Borchsenius, 1949), female. Original.

segment covered with a few, strong hair-like setae (except third segment); apical segment
with apical seta 39 µm long; apical segment also with 3 sensory falcate setae, longest 33
falcate seta 20 µm long. Frontal lobes and frontal tubercle present. Eyes visible, situated
on venter. Anal lobes slightly sclerotized each with two enlarged setae along inner margin,
and one enlarged seta on outer margin, similar in size to those on dorsal abdominal
margin, 30–40 µm long, apical seta 271–300 µm long.
Venter: Labium 3 segmented, 101 µm long; stylet loop two-three times longer than
labium. Legs well developed; lengths of segments and digitules of prothoracic legs;
coxa: 72–86, trochanter: 55–62, femur: 173–190, tibia: 154–180, tarsus: 163, claw: 34,
tarsal digitules 50, claw digitules: 38 µm long; lengths of segments and digitules of
tarsus: 157–161, claw: 36–40 µm long, tarsal digitules broken, claw digitules broken on
studied material; lengths of segments and digitules of metathoracic legs; coxa: 89–101,
trochanter: 60, femur: 173–192, tibia 169–177, tarsus 160–168, claw: 36–38, tarsal
digitules: 48, claw digitules: 42 µm long. Tarsal and claw digitules knobbed, longer than
claw; claw with denticle at apex. Meso- and metathoracic coxae each without spinulae on
ventral surface, translucent pores on dorsal surface of posterior coxae. Tibiae each with
5 setae (median seta present). The diameter of anterior spiracles 29–31 µm, posterior
spiracles slightly larger than anterior. Venter with hair-like setae, scattered; setae on the
middle of abdominal segment the longest and with a marginal and submarginal row of
enlarged setae. Quinquelocular pores 5.0 µm in diameter, distributed in sparse bands
and rows on all segments of abdomen and thorax, trilocular pores 5.0 µm in diameter, a
few on thorax and head. Macrotubular ducts of two sizes, larger ones of the same size
as those on dorsum in a band on marginal area, narrower ones about 4 µm wide, 22 µm
long, scattered over entire body. Microtubular ducts on submarginal area. Cruciform
pores absent.
Dorsum: Enlarged setae spine-like, slightly blunted, on margin 46–53 µm long, varying
in size and shape. Other dorsal enlarged setae, some slightly curved, forming sparse
rows; in middle part of posterior segments 15–24 µm long, on thorax they are somewhat
longer; submarginal band without enlarged setae; on margin of abdominal segments
usually 3 longer enlarged setae present. Macrotubular ducts sparse, present throughout, 7
µm wide, 22 µm long. Microtubular ducts with 3.0 µm long, scattered throughout. Anal
ring, 55 µm wide, 72–86 µm long with eight setae, each 110–113 µm long. Cauda present
Ecology
Host plant: Asplenium ceterach.
Distribution: China (?), Iran, Tajikistan, Turkey.
Acanthococcidae 543
Rhizococcus siakwanensis (Borchsenius, 1960) (Fig. 193) Combination

nova
Acanthococcus siakwanensis Borchsenius, 1960c: 916.
Holotype: Female. China (Yunnan, Siakwan), on unidentified Compositae, 17.iv.1957,

by N. Borchsenius. By original designation. Deposited in ZMAS.
Lit.: Ali, 1970: 77; Hoy, 1963: 115; Hua, 2000: 137; Tao, 1999: 33; Wang, 1980: 115;
2001: 208; Yang, 1982: 105.
Eriococcus siakwanensis; Hoy, 1963: 115. Change of combination.

Acanthococcus siakwanensis; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus siakwanensis; Tang & Hao, 1995: 451. Revived combination.
Acanthococcus siakwanensis; Köhler, 1998: 383. Revived combination.
Eriococcus siakwanensis; Miller & Gimpel, 2000: 332. Revived combination.
Acanthococcus siakwanensis; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female is oval, about 2 mm long.
Mounted female
Body elongate oval, 3.5 mm long. Antennae short, 6 or 7 segmented. Frontal lobes
present. Eye visible, situated on venter. Anal lobes small, strongly sclerotized with three
enlarged setae along inner margin, and one enlarged seta on outer margin; apical setae
twice longer than anal ring setae, cauda present.
Venter: Labium 3 segmented, basal segment with a pair of setae one is three times
longer than other, apical segment with long hair-like setae, stylet loop reaches median
coxae. Legs well developed; short, tarsus longer than tibia, claw with strong denticle,
tarsal and claw digitules knobbed, longer than claw. Posterior coxae with spinulae and
large pores. The spiracles strongly sclerotised, associated with 10–14 quinquelocular
pores at openning. Venter with scattered, in small number strong hair-like setae, some
of them ae capitated, and with a submarginal row of enlarged setae. Quinquelocular
and trilocular pores, distributed in bands and rows on all segments of abdomen and on
thorax. Macrotubular ducts of two sizes, scattered over entire body. Cruciform pores
absent.
Dorsum: Enlarged setae spine-like, blunted, in transverse rows and bands, on the margin
of abdominal segments with 3 to 5 setae. Macro- and microtubular ducts similar to those
on venter, sparse, present throughout. Anal ring with 8 long setae. Cauda present (after
Borchsenius (1960c), with modifications).
Ecology
Host plant: Anaphalis sp., Carpesium abrotanoides, Carpesium sp., Dendranthema sp., Pistacia
weinmannifolia.
Biology: On the roots and root crown.
Acanthococcidae 545
Figure 193. Rhizococcus siakwanensis (Borchsenius, 1960), female. After Borchsenius

Rhizococcus sojae (Kuwana, 1917) (Fig. 194) Combination nova

Eriococcus sojae Kuwana, 1917: 136.
Holotype: Female. Japan: Honshu, Okayama-ken, on Glycine soja, Fall 1915. Syntypes,
female, type designation unknown. Type depository: Ibaraki-ken: Insect Taxonomy
Laboratory, National Institute of Agricultural Environmental Sciences, Kannon-dai,
Yatabe, Tsukuba-shi, (Kuwana), Japan.
Lit.: Hoy, 1963: 116; Ishihara and Uwagawa, 1950: 187; Kawai, 1980: 130; Kwon
& Han, 2003: 151; Takahashi, 1957: 7; Tao, 1999: 33; Wang, 2001: 209 (Miller et al.,
2013).
Acanthococcus sojae; Kozár & Walter, 1985: 74. Change of combination.

Eriococcus sojae; Tang & Hao, 1995: 416, 595. Revived combination.
Acanthococcus sojae; Köhler, 1998: 383. Revived combination.
Eriococcus sojae; Miller & Gimpel, 2000: 336. Revived combination.
Acanthococcus sojae; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female is enclosed in the grayish white, cottony sac. Female body is elliptical, dark
purplish red. Eggs are elliptical and flesh colored (Kuwana, 1917).
Mounted female
Body elongate oval. Antennae short, 7 segmented. Eyes visible, situated on venter. Anal
lobes small, strongly sclerotized with two enlarged setae along inner margin, and one
enlarged seta on outer margin.
Venter: Labium 3 segmented. Legs well developed; short, claw with strong denticle. Tarsal
and claw digitules knobbed, longer than claw. Posterior coxae with pores. The spiracles
strongly sclerotised, associated with 12-16 quinquelocular pores at openning. Venter with
scattered, in small number of hair-like setae and with a marginal and submarginal row of
enlarged setae. Quinquelocular pores distributed in wide bands and rows on all segments
of abdomen and on thorax. Macrotubular ducts of one size scattered all body. Cruciform
pores scattered on submargin.
Dorsum: Enlarged setae spine-like with pointed apices, on margin distinctly larger than
other dorsal setae; marginal row on the abdominal segment II-VII bears 2 or 3 enlarged
setae. Other dorsal enlarged setae, forming sparse rows. Macrotubular ducts wider than
those on venter, with microtubular ducts present throughout. Anal ring with 8 long setae.
Cauda not shown (after of Tang & Hao, (1995) with modification).
Acanthococcidae 547
Figure 194. Rhizococcus sojae (Kuwana, 1917), female. After Tang & Hao (1995) with
modifications.
Ecology
Host plant: Calystegia hederacea, Cephalanoplos segetum, Glycine max, G. soja.
Distribution: China, Japan, South Korea.
Biology: It is assumed that the species overwinters as eggs which hatch once a year. First
instars attack plants in late July, and then reach the sac formation stage in the middle of
October, laying eggs in the same season. The damage caused by the scale can be heavy,
resulting in death (Kuwana, 1917).
Rhizococcus spiniferus (Borchsenius, 1949) (Fig. 195) Redescription,

Combination nova
Acanthococcus spiniferus Borchsenius, 1949: 338.
Lectotype: Female. Tajikistan (near Ayvadzh), on grass leaves, by N. Borchsenius.

By subsequent designation Danzig, 1996: 522. Deposited in ZMAS (type no. 7-44).
Notes: One paralectotype female on separate slide with same data in ZMAS (Danzig,
1996). According to original description on Aeluropus littoralis.
Lit.: Babaev, 1980: 57; Borchsenius, 1950: 119; Tereznikova, 1977: 571 (Miller et al.,
2013).
Eriococcus spiniferus; Hoy, 1963: 117. Change of combination.

Acanthococcus spiniferus; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus spiniferus; Tang & Hao, 1995: 451. Revived combination.
Acanthococcus spiniferus; Köhler, 1998: 384. Revived combination.
Eriococcus spiniferus; Miller & Gimpel, 2000: 338. Revived combination.
Acanthococcus spiniferus; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female is elongate, 2.2 mm long. Sac is elongate oval, up to 3 mm long, felted,
compact, grayish, entirely covering the body.
Mounted female
covered with a few, strong hair-like setae (except third segment); apical segment with
long; segment VI with 1 sensory falcate seta 24 µm, segment V with 1 sensory falcate
seta 17 µm long. Frontal lobes not seen. Eyes visible, situated on venter. Anal lobes
slightly sclerotized each with two enlarged setae along inner margin, and one enlarged
Acanthococcidae 549
Figure 195. Rhizococcus spiniferus (Borchsenius, 1949), female. Original.

seta on outer margin (a bit larger those on inner side), similar in size to those on dorsal
abdominal margin, apical seta broken.
Venter: Labium 3 segmented, 94 µm long, basal segment with two pairs of setae almost
equal long, median setae on apical segment very short. Legs well developed; lengths of
segments and digitules of prothoracic legs; coxa: 72, trochanter: 48, femur: 156, tibia:
132, tarsus: 120 and claw: 36, tarsal digitules: 51, claw digitules: 17 µm long; lengths
of segments and digitules of mesothoracic legs; coxa: 90, trochanter: 62, femur: 144,
tibia: 144, tarsus: 132 and claw: 41 µm long, tarsal digitules: 53, claw digitules: 17 µm
long; lengths of segments and digitules of metathoracic legs; coxa: 96, trochanter: 62,
femur:156, tibia: 156, tarsus: 149 and claw: 44, tarsal digitules: 48, claw digitules: 16 µm
long. Tarsal digitules knobbed, longer than claw; claw digitules setalike shorter than
claw; claw with denticle at apex. Meso- and metathoracic coxae each without spinulae
on ventral surface, translucent pores on dorsal surface. Tibiae each with 5 setae (median
seta present). Posterior spiracles slightly larger than anterior. Venter with scattered, hair-
like setae and with a marginal and submarginal row of enlarged setae, on submargin
areae of abdomen, enlarged setae and a few hair-like setae together in groups. Discodial
pores each 4.0–5.0 µm in diameter, with 3-5 loculi; distributed in very sparse bands and
rows on all segments of abdomen and thorax. Macrotubular ducts of two sizes, larger
ones on marginal area, about 7 µm wide, narrower ones about 4 µm wide, 24 µm long, in
small numbers scattered over entire body. Microtubular ducts not seen. Cruciform pores
absent.
blunted. Other dorsal enlarged setae, forming sparse transverse bands; larger size forming
3 longitudinal lines on each side of abdomen, in middle part of posterior segments, 24
µm long, on thorax they are somewhat longer; on margin of abdominal segments usually
4 enlarged setae present. Macrotubular ducts sparse, present throughout body, 7 µm
wide, 22 µm long. Microtubular ducts with 4 µm long, scattered throughout. Anal ring 63
µm wide, 74 µm long, with a row of pores, and 8 setae. Cauda not seen (Redescription is
Ecology
Host plant: Aeluropus littoralis.
Distribution: Tajikistan
Acanthococcidae 551
Rhizococcus subterraneus (Borchsenius, 1949) (Fig. 196) Redescription,

Combination nova
Acanthococcus subterraneus Borchsenius, 1949: 349.
Lectotype: Female. Uzbekistan (near Dzhizak), on Artemisia sp. roots, 21.v.1940, by,
N. Borchsenius. By subsequent designation Danzig, 1996: 522. Deposited in ZMAS
(type no. 9-44). Notes: Ten paralectotypes on 9 slides from Armenia: Megri, on roots
of Artemisia sp., 24-26.v.1947, by N. Borchsenius, numbered 134-48, 187-47, 195-47,
205-47 (Danzig, 1996).
Common name: Mugworth felt scale.
Lit.: Babaeva, 1980: 57; Bazarov, 1971: 90; Borchsenius, 1950: 121; Kosztarab &
Kozár, 1978:75; Matesova, 1971: 26; Ter-Grigorian, 1983: 880 (Miller et al., 2013).
Nidularia subterraneus; Lindinger, 1957: 543. Change of combination.

Eriococcus subterraneus; Hoy, 1963: 118. Change of combination.
Acanthococcus subterraneus; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus subterraneus; Tang & Hao, 1995: 452. Revived combination.
Acanthococcus subterraneus; Köhler, 1998: 384. Revived combination.
Eriococcus subterraneus; Miller & Gimpel, 2000: 351. Revived combination.
Acanthococcus subterraneus; Kozár, 2009: 93. Revived combination.
Description
Unmounted female
Adult female body broad, egg-shaped, dark olive in color, 3 mm long, 2 mm wide.
Mounted female
segments: I: 31, II: 29, III: 72, IV: 22, V: 22, and VI: 29 µm long; each segment covered
with a few, strong hair-like setae; apical segment with apical seta 43 µm long; apical
segment also with 3 sensory falcate setae, longest 29 µm long; segment VI with 1 sensory
falcate seta 24 µm, segment V with 1 sensory falcate seta 12 µm long. Frontal lobes absent,
frontal tubercle present. Eyes visible, situated on venter. Anal lobes slightly sclerotized
each with two enlarged setae along inner margin, and one enlarged seta on outer margin,
similar in size to those on dorsal abdominal margin, apical seta 268 µm long.
Venter: Labium 3 segmented, 120 µm long. Legs well developed; lengths of segments
and digitules of prothoracic legs; coxa: 53, trochanter: 41, femur: 113, tibia: 84, tarsus: 96
and claw: 26, tarsal digitules broken, claw digitules 26 µm long; lengths of segments and
digitules of mesothoracic legs: coxa: 50, trochanter: 38, femur: 101, tibia: 84, tarsus: 96
and claw: 41, tarsal digitules: 38, claw digitules broken; lengths of segments and digitules
of metathoracic legs; coxa: 50, trochanter: 48, femur: 113, tibia: 96, tarsus: 106 and claw:
26, tarsal digitules: 31, claw digitules: 26 µm long. Tarsal and claw digitules knobbed; claw
with denticle at apex. Meso- and metathoracic coxae each without spinulae on ventral
surface, translucent pores on dorsal surface of posterior coxae. Tibiae each with 5 setae
(median seta present). Spiracle 29 µm in diameter, posterior spiracles slightly larger
than anterior. Venter with scattered, hair-like setae and with a marginal row of enlarged
setae, two groups enlarged setae on first abdominal segment and between spiracles on
submarginal area. Discodial pores each 5.0 µm in diameter, with 3-5-8 loculi, distributed
in very sparse bands and rows on all segments of abdomen and thorax, a few associated
with spiracles. Macrotubular ducts of two sizes, larger ones on marginal area, about 7 µm
wide, narrower ones about 5 µm wide, 24 µm long, in small number scattered all body.
Microtubular ducts on submarginal area of body. Cruciform pores few, scattered on
thorax and head.
blunted. Other dorsal enlarged setae, forming sparse rows; in middle part of posterior
segments 12–20 µm long, some of them slightly curved, on thorax they are somewhat
longer; on margin of abdominal segments 2 enlarged setae present. Macrotubular ducts
sparse, present throughout all body, 8 µm wide, 28 µm long. Microtubular ducts with 5
µm long scattered throughout. Anal ring 55 µm wide, 53 µm long, with eight setae, each
134 µm long. Cauda not seen (Redescription is based on type material).
Ecology
Host plant: Artemisia sp., Salicornia sp.
Distribution: Armenia, Spain, Uzbekistan.
Biology: On the roots and root crown.
Comments
Specimens from Spain (Aquadulae (Almeria) from Salicornia sp., (No. 7796 in PPI) Leg: J.
G. Menor were identified as R. subterranea, a new record for the fauna of Spain.
Acanthococcidae 553
Figure 196. Rhizococcus subterraneus (Borchsenius, 1949), female. Original.

Rhizococcus targassonensis (Goux, 1993) (Fig. 197) Redescription

Eriococcus targassonensis Goux, 1993: 67.
Holotype: Female. France (Tergassone, Pyrénées), on undetermined Compositae,

17.viii.1953, by L. Goux (MNHN 14515-01). Notes: Paratypes: 5 slides are with the
same data as holotype (MNHN 14515-02-06). Deposited in MNHN.
Lit.: Foldi, 2001: 305; Miller & Gimpel, 2000: 353 (Miller et al., 2013).
Acanthococcus targassonensis; Miller & Gimpel, 1996: 604. Change of combination.

Rhizococcus targassonensis; Kozár, 2009: 94; Ouvrard & Kozár, 2009: 102. Change of
combination.
Description
Unmounted female
Body oval, 1.8 mm long, and 1 mm wide.
Mounted female
VII: 30–43 µm long; each segment covered with a few, strong hair-like setae (except
3 sensory falcate setae, longest 30–32 µm long; segment VI with 1 sensory falcate seta
20–29 µm, segment V with 1 sensory falcate seta 16 µm long. Frontal lobes present. Eyes
visible, situated on venter. Anal lobes slightly sclerotized each with two enlarged setae
along inner margin, and one enlarged seta on outer margin, each 46–70 µm long, similar
in size to those on dorsal abdominal margin, apical seta 240–290 µm long.
Venter: Labium 3 segmented, 105–115 µm long, stylet loop reaching behind median
coxae. Legs well developed; lengths of segments and digitules of prothoracic legs; coxa:
70–89, trochanter: 55–66, femur: 115–145, tibia: 95–110, tarsus: 118–130; claw: 30–37,
tarsal digitules: 44–48, claw digitules: 28–32 µm long; lengths of segments and digitules
of mesothoracic legs; coxa: 60–82, trochanter: 45–70, femur: 126–140, tibia: 115–125,
tarsus: 128–136; claw: 30–34, tarsal digitules: 45–48, claw digitules: 30–32 µm long;
femur: 148–151, tibia: 105–130, tarsus: 136–140; claw: 28–32, tarsal digitules: 45–51, claw
digitules: 28–32 µm long. Tarsal and claw digitules knobbed, slightly longer than claw;
claw with denticle at apex. Metathoracic coxae and femur with large translucent pores
on dorsal surface, metathoracic coxae without spinula. Tibiae each with 5 setae (median
seta present). Spiracle 31–36 µm in diameter, with a group of quinquelocular pores
around anterior spiracles, posterior spiracles slightly larger than anterior. Venter with
scattered, hair-like setae and with a marginal band and a submarginal row of enlarged
setae. Quinquelocular pores each 5.0–6.0 µm in diameter, distributed in very sparse bands
Acanthococcidae 555
Figure 197. Rhizococcus targassonensis (Goux,1993), female. Original.

and rows on all segments of abdomen and thorax. Macrotubular ducts of two sizes,
larger ones on marginal area, about 7–10 µm wide, 21–25 µm long, narrower ones about
4–7 µm wide, 14–15 µm long, scattered all body. Microtubular ducts scattered on body.
Cruciform pores few, scattered on thorax and first abdominal segment on submarginal
area.
blunted, on margin of abdominal segments 2 enlarged setae present; other dorsal enlarged
setae, forming sparse rows, 10–18 µm long; on thorax they are somewhat longer, setae
in middle part of posterior segments comperatevely small and spinelike, 10–13 µm long.
Macrotubular ducts sparse, present throughout all body, 8–10 µm wide, 21–25 µm long.
Microtubular ducts with 4–5 µm long scattered throughout. Anal ring 58–64 µm wide,
61–82 µm long, with a row of pores, and with 8 setae, each 110–130 µm long. Cauda not
Ecology
Host plant: Asteraceae.
Biology: Unknown.
Rhizococcus teucriicolus (Bodenheimer, 1943) (Fig. 198) Combination

nova
Eriococcus teucriicolus Bodenheimer, 1943: 24.
Syntype: Female. Iraq (Addaye), on Teucrium polium, 10.10.1942. Deposited in Bet

Dagan.
Lit.: Ben-Dov & Harpaz, 1986: 30; Hoy, 1963: 118; Köhler, 1998: 384 (Miller et al.,
2013).
Acanthococcus teucriicolus; Kozár & Walter, 1985: 74. Change of combination.

Eriococcus teucriicolus; Miller & Gimpel, 2000: 356. Revived combination.
Acanthococcus teucriicolus; Kozár, 2009: 94. Revived combination.
Description
Unmounted female
Sac is ovoid, felted, white in color, 6 mm long, 4 mm wide. Body of adult female is ovoid.
Mounted female
Acanthococcidae 557
Figure 198. Rhizococcus teucriicolus (Bodenheimer, 1943), female. After Bodenheimer

covered with a few, strong hair-like setae. Frontal lobes present. Eyes visible, situated on
venter. Anal lobes short, conical, with two enlarged setae along inner margin, and one
enlarged seta on outer margin (enlarged setae 30–60 µm long); apical seta 320 µm long.
Venter: Labium 3 segmented, much longer than width. Stylet loop as long as labium.
Legs well developed; hind femur: 180 µm long, 160 µm long, claw: 20 µm long. Metacoxa
without translucent pores. Femur distinctly, tibia slightly swollen from base to apex, tarsus
slightly tapering; claw slender, slightly curved, with denticle; tarsal and claw digitules
knobbed, slightly longer than claw. Tibiae each with 5 setae (median seta present).
Spiracle, small, slender, dumb bell shaped, posterior spiracles slightly larger than anterior.
Venter with 1 or 2 rows of hair-like setae, 40–70 µm long, marginal row of enlarged setae
not shown. Quinquelocular pores in bands on abdominal segment and scattered a few
on thorax and head.
Dorsum: Enlarged setae stout and pointed, 4-8 irregularly strong enlarged setae on head
and thorax, enlarged setae on other segments various sizes, in wide bands, setae on
margin ca. 50 µm long, on dorsum ca. 40 µm long. Macrotubular ducts scattered over
body. Anal ring almost circular, pore bands in partly in double rows, with 8 strong setae,
each 60–100 µm long. Cauda not seen (after Bodenheimer (1943).
Ecology
Host plant: Teucrium polium.
Distribution: Iraq.
Biology: On twigs and branches, rather common.
Acanthococcidae 559
Rhizococcus thaleri (Szita & Kozár, 2011) (Fig. 199) Combination nova
Acanthococcus thaleri Szita & Kozár, 2011 (in Szita et al., 2011: 37).
Holotype: Female. Austria (Siebenstein, near Molln vill., Kirchdorf an der Krems distr.),
in ?.?.1934, on Erica carnea, by E.E. Green. By original designation. (type no. 180). Notes:
Holotype collected at N: 47º19', E: 14º12', ca. 1100 m altitude. Paratypes on a separate
slide from Austria, Gloggnitz town on Erica carnea by E.E. Green. Deposited in BMNH.
Lit.: Szita et al., 2011: 37 (Miller et al., 2013).
Description
Unmounted female
Unknown.
Mounted female
length of segments: I: 30–31, II: 17–25, III: 55–70, IV: 14–15, V: 14–17 and VI: 24–30
µm long; each segment covered with a few, strong hair-like setae; apical segment with
apical seta 28–31µm long; apical segment also with 3 sensory falcate setae, longest 30–32
µm long; segment V with 1 sensory falcate seta 12–14 µm, segment IV with 1 sensory
falcate seta 14 µm long. Frontal tubercle and frontal lobe present. Eyes situated on venter
near margin. Anal lobes well developed, each with three enlarged setae (different from
dorsal enlarged setae) on dorsal surface, outer setae stronger than those on inner side;
anal lobe seta 139–160 µm long.
Venter: Labium 3 segmented, 86–91 µm long, basal segment not well developed, but
with two setae present on each side; median setae on apex of labium needle-like. Legs
well developed; lengths of segments and digitules of prothoracic legs; coxa: 70–89,
trochanter: 43–45, femur: 89–104, tibia: 70–80, tarsus: 84–96; claw: 24–29, tarsal digitules:
45, claw digitules: 30 µm long; lengths of segments and digitules of mesothoracic legs;
coxa: 48–50, trochanter: 46–55, femur: 84–105, tibia: 77–84, tarsus: 82–108, claw: 26,
tarsal digitules broken, claw digitules: 31 µm long; lengths of segments and digitules of
metathoracic legs; coxa: 50–58, trochanter: 48–55, femur: 89–110, tibia: 79–90, tarsus:
89–102, claw: 29–30, tarsal digitules: 47–48, claw digitules: 34 µm long. Tarsal and claw
digitules knobbed, slightly longer than claw; claw with denticle at apex. Median and
posterior coxae with spinulae on anterior surface; posterior coxae and femur with small
pores on posterior surface. Tibiae each with 5 setae (median seta present). Diameter of
anterior spiracles 31–38 µm, posterior spiracles slightly larger than anterior. Derm with
a few hair-like setae; with a submarginal band of enlarged setae, and a row of truncated
conical spines seen on margin, with a band of macrotubular ducts 7 µm wide 18 µm long
in marginal position. Multilocular pores each 4–6 µm in diameter and with 3-9 loculi,
mostly 5, distributed in sparse rows on all abdominal segments, on thoracic segments
mainly on thoracic segment III. Macrotubular ducts of two sizes, larger ones on marginal
area, about 7–10 µm wide, 21–25 µm long, narrower ones about 4–7 µm wide, 14–15
µm long, scattered all body. Smaller macrotubular ducts 4 µm wide and 18 µm long
scattered throughout venter, but more abundant submarginally. Microtubular ducts
sparse submarginally and marginally. Cruciform pores sparse on submargin, 4 µm in size.
Dorsum: Dorsal setae truncated conical spines, 19–29 µm long, at its base ca. two-third
times wide as long, in sparse rows across all segments. Macrotubular ducts 7 µm wide and
19–29 µm long, associated with truncated conical spines. Microtubular ducts 4 µm long
scattered. Anal ring strongly sclerotized, with partly double rows of pores; slightly oval,
53–62 µm wide, 62–72 µm long, with 4 pairs of 74–86 µm long setae; anal ring situated
on margin of dorsum, posterior part without pores. Cauda not seen (after Szita & Kozár
(2011)).
Ecology
Host plant: Erica carnea.
Distribution: Austria.
Biology: Unknown.
Acanthococcidae 561
Figure 199. Rhizococcus thaleri (Szita & Kozár, 2011), female. After Szita et al. (2011).
Rhizococcus thymelaeae (Newstead, 1897) (Fig. 200) Redescription,

Combination nova
Eriococcus thymelaeae Newstead, 1897a: 102.
Lectotype: Female. Algeria (Constantine, on the slopes of Mansourah, near Depot

des fourrages of the Chasseurs d'Afrique), on Thymelaea hirsuta, 28.x.1895, by A.E.
Eaton. Designated here. Deposited in BMNH.
Lit.: Balachowky, 1927: 188-189; 1935: 265; Bodenheimer, 1935: 251; Cockerell,
1899a: 391; Fernald, 1903: 79; Gómez-Menor Ortega, 1937: 345, 432; 1965: 113;
Hodgson & Miller, 2010: 99; Hodgson & Trencheva, 2008: 31; Kiritchenko, 1940:
127; Köhler, 1998: 384; Lindinger, 1912: 324; 1933: 117; Martin-Mateo, 1985: 93;
Ouvrard & Kozár, 2009: 102. Trabut, 1911: 53; Vieira et al., 1983: 136 (Miller et al.,
2013).
Nidularia thymelaeae; Lindinger, 1935: 135. Change of combination.

Eriococcus thymelaeae; Hoy, 1963: 119. Revived combination.
Acanthococcus thymelaeae; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus thymelaeae; Miller & Gimpel, 2000: 359. Revived combination.
Acanthococcus thymelaeae; Kozár, 2009: 94. Revived combination.
Eriococcus tucurincae madeirensis; Balachowsky, 1939. Synonym nova
Eriococcus tucurincae madeirensis; Balachowsky, 1939: 267.
Syntypes: Female. Portugal (Madeira Islands, Paulo de Serra), on Thymus caespititius,
?/08/1936, by Balachowsky; Désertas Islands, Grande Désertas, on Silene gallica,
?/08/1936, by A. Balachowsky (type no. 4956). Notes: There are three slides with
seven syntype specimens from Paulo de Serra and seven slides with twenty-five
syntype specimens from Grande Désertas. The slide mounted specimens indicate
that the host from Grande Désertas is Silene maritima. A second MNHN number is
4958.
Nidularia henryi; Lindinger, 1943: 223. Incorrect synonymy; discovered by Hoy, 1963: 101.
Eriococcus madeirensis; Balachowsky, 1946: 215. Change of status.
Acanthococcus madeirensis; Kozár & Walter, 1985: 74. Change of combination.
Eriococcus madeirensis; Miller & Gimpel, 2000: 260-261. Revived combination.
Acanthococcus madeirensis; Kozár, 2009: 92. Revived combination.
Description
Unmounted female
Female sac is short, ovate, irregular in form, 3 mm long, 2 mm wide. Adult female is
elongate oval.
Acanthococcidae 563
Mounted female
VII: 38–41 µm long; each segment covered with a few, strong hair-like setae (except
3 sensory falcate setae, longest 28–29 µm long; segment VI with 1 sensory falcate seta
20–34 µm, segment V with 1 sensory falcate seta 17–20 µm long. Frontal tubercle and
frontal lobe present. Eyes situated on venter near margin. Anal lobes well developed,
each with three enlarged setae on dorsal surface, outer setae stronger than those two on
inner side; enlarged setae similar to other dorsal setae; anal lobe seta 310–353 µm long.
Venter: Labium 3 segmented, 100–115 µm long, basal segment with two setae present
on each side. Stylet loop reaching behind median coxae. Legs well developed; lengths of
segments and digitules of prothoracic legs; coxa: 79–95, trochanter: 55–58, femur: 144–
148, tibia: 125–128, tarsus: 124–134, claw: 39, tarsal digitules: 48, claw digitules: 32 µm
long; lengths of segments and digitules of mesothoracic legs: coxa: 72, trochanter: 60,
femur: 137, tibia: 129, tarsus: 130 claw: 40, tarsal digitules: 58, claw digitules: 35 µm long;
lengths of segments and digitules of metathoracic legs; coxa: 86, trochanter: 60, femur:
149, tibia: 134, tarsus: 139, claw: 41, tarsal digitules: 50, claw digitules: 37 µm long. Tarsal
and claw digitules knobbed, slightly longer than claw; claw with denticle at apex. Median
and posterior coxae with spinulae on anterior surface; posterior coxae and femur with
small pores on posterior surface. Tibiae each with 5 setae (median seta present). Diameter
of anterior spiracles 35–36 µm, posterior spiracles slightly larger than anterior. Derm
with a few hair-like setae, some of them capitated; and with a submarginal and marginal
rows of enlarged setae with a band of macrotubular ducts. Multilocular pores each 6–7
µm in diameter and with 3-5 loculi, mostly with 5 loculi, distributed in bands and rows on
all abdominal segments, scattered on thorax and head. Macrotubular ducts of two sizes,
larger ones on marginal area, about 8.0 µm wide, 14.0 µm long, narrower ones about 4–7
µm wide, 14–15 µm long, scattered all body. Microtubular ducts sparse submarginally and
marginally. Cruciform pores sparse on submargin and margin, 4 µm wide.
Dorsum: Marginal dorsal enlarged setae conical spine-like, blunted, 42–51 µm long, each
abdominal segment with 3 or 4 enlarged setae on margin, other dorsal enlarged setae
various sizes, 32–47 µm long, in sparse rows and bands across all segments; setae on the
middle part of thorax somehow larger than others dorsal setae. Macrotubular ducts 4–6
µm wide and 17–25 µm long, sitiuated among enlarged setae. Microtubular ducts 5 µm
long, scattered. Anal ring strongly sclerotized, oval, with partly double rows of pores;
40–48 µm wide, 55–85 µm long, with 4 pairs of long setae, 115–125 µm long. Cauda
present (after Hodgson & Trencheva, 2008, with modifications).
Other stages
Nymphs:
Second instar female nymph described by Hodgson & Trencheva (2008). All spines on the
dorsum are much shorter, than marginal ones, macrotubular ducts absent, microtubular
ducts present on dorsum, some trilocular and quinquelocular pores and cruciform pores
present on venter.
Ecology
Host plant: Poterium spinosum, Quercus coccifera, Ruta montana, Silene gallica, Thymelaea hirsuta,
Thymelaea virgata var. broussonetii, Thymus caespititius.
Distribution: Algeria, Cyprus Island, Egypt, Greece, Israel, Morocco, Portugal (Madeira),
Spain, Tunisia.
Comments
Because of all studied characters number and size categories of syntype materials were
very similar to the lectotype of R. thymelaeae, here we consider R. madeirensis as synonym
of the R. thymelaeae.
Acanthococcidae 565
Figure 200. Rhizococcus thymelaeae (Newstead, 1897), female. Original.

Rhizococcus timidus (Hulden, 1985) (Fig. 201) Combination nova

Eriococcus timidus Hulden, 1985: 59.
Holotype: Female. Finland (Kuhmo, Timonniemi), on Erica tetralix, 08.vii.1965, by Y.

Mäkinen. By original designation. Deposited in Helsinki.
Lit.: Gertsson, 2001: 125; Köhler, 1998: 385 (Miller et al., 2013).
Acanthococcus timidus; Miller & Gimpel, 1996: 604. Change of combination.

Eriococcus timidus; Miller & Gimpel, 2000: 362. Revived combination.
Acanthococcus timidus; Kozár, 2009: 94. Revived combination.
Description
Unmounted female
Adult female brown, cylindrical, 2.2 mm long, 0.8 mm wide. Ovisac white, felt-like,
cylindrical in shape, 2.7 mm long, 1.2 mm wide (Hulden, 1985).
Mounted female
Body elongate oval. Antennae 6 segmented, length about 190 µm. Anal lobes well
developed, each with three enlarged setae on dorsal surface, enlarged setae similar to
other dorsal setae; anal lobe seta 220 µm long.
Venter: Legs well developed; lengths of tarsus 144–145 µm long, index of length/
width of femur about 2.5. Tarsal and claw digitules knobbed, slightly longer than claw.
Tibiae each with 5 setae (median seta present). Derm with a few hair-like setae; a row
of enlarged setae seen on margin with some macrotubular ducts in marginal position.
Multilocular pores not shown.
Dorsum: Marginal dorsal enlarged setae conical spine-like, apices acute or slightly
rounded, setae of 1 general size, abundant over dorsum 20–40 µm long, each abdominal
segment with 3 or 4 enlarged setae, other dorsal enlarged setae in sparse rows and bands
across all segments. Macrotubular ducts sitiuated among enlarged setae. Dorsal derm
with some granulation. Anal ring with 6 (?) setae, each 120 µm long (Hulden, 1985).
Ecology
Host plant: Erica tetralix
Distribution: Finland.
Biology: On underside of the leaves.
Comments
The description is very short and the drawing is not clear, and not easy to interpret.
Hulden (1985) disccuss the similarity with A. ericae, R. devoniensis and R. marginalis. The
difference from first two species cited by Hulden (1985) could be accepted. Actually
Acanthococcidae 567
there are some similarities with R. marginalis, but differs by their host plant (Kochia sp.) and
geographic distribution (Armenia). The question on systematic position of this species
needs further material and needs more study.
Figure 201. Rhizococcus timidus (Hulden, 1985), female. After Hulden (1985) with
modifications.
Rhizococcus turkmenicus (Archangelskaya, 1930) (Fig. 202) Combination

nova
Eriococcus turkmenicus Archangelskaya, 1930: 77.
Syntype: Female. Turkmenistan (Kushka), on Artemisia sp. branches, ?.vi.1927, by M.

Umnov. Notes: Type material lost.
Lit.: Archangelkya, 1931: 69; 1937: 128, 139; Borchsenius, 1949: 53; Danzig, 1974:
70; 1982: 147; Köhler, 1998: 385; Lindinger, 1957: 544; Myartseva, 1981: 99; Tang &
Hao, 1995: 451 (Miller et al., 2013).
Acanthococcus turkmenicus; Borchsenius, 1949: 53. Change of combination.

Nidularia turkmenicus; Lindinger, 1957: 544. Change of combination.
Eriococcus turkmenicus; Hoy, 1963: 123. Revived combination.
Acanthococcus turkmenicus; Kozár & Walter, 1985: 74. Revived combination.
Eriococcus turkmenicus; Miller & Gimpel, 2000: 369. Revived combination.
Acanthococcus turkmenicus; Kozár, 2009: 94. Revived combination.
Description
Unmounted female
Adult female body is egg-shaped, light-yellow color, legs and antennae are light-brown,
2.5 mm long, 1.4 mm wide.
Mounted female
Body elongate oval. Antennae 7 segmented, length of segments: I: 41, II: 30, III: 49, IV:
37, V: 24, VI: 19 and VII: 38 µm long; each segment covered with a few, strong hair-like
setae (except third segment); Anal lobes strongly sclerotised, each with three strong hair-
like setae on dorsal surface 60 µm long, anal lobe seta 255 µm long, subapical setae 150
µm long.
Venter: Labium wide. Stylet loop reaching median coxae. Legs well developed, strong;
femur: 170, tibia: 125, tarsus: 135, claw digitules much shorter than claw, blunted,
posterior coxae and femur with small pores, claw with large denticle. Derm with a few
strong hair-like setae; and a band of enlarged setae on submargin, 13–38 µm long, preanal
setae 80 µm long, with a band of macrotubular ducts in marginal position, and scattered
elsewhere, 6 µm wide and 22 µm long. Multilocular pores distributed in rows and bands
on abdominal segments, scattered on thorax and head. Microtubular ducts and cruciform
pores not mentioned.
Dorsum: Marginal dorsal enlarged setae conical spine-like, apices acute abundant in wide
bands over surface; 27–60 µm long, abdominal segments with about 5 enlarged setae, 55–
60 µm long, Macrotubular ducts 11 µm wide and 38 µm long, sitiuated among enlarged
setae in bands and rows. Anal ring strongly sclerotized, with partly double rows of pores;
Acanthococcidae 569
slightly oval, with 4 pairs of long setae, 135–150 µm long (after Archangelskaya (1931);
Borchsenius (1949) with modifications).
Ecology
Biology: On the roots and branches.
Comments
There is some contradiction between the description of Archangelskaya (1930) and
Borchsenius (1949), concerning the presence of multilocular pores on dorsum. According
to Archangelskaya multilocular pores are present (as mentioned in the Russian and
English text of the author), Borchsenius (1949) mentioned their presence clearly only
on venter, and did not pay attention to this part of the text by Archangelskaya (1930).
Macrotubular ducts were also not mentioned by Archangelkaya (1930).
Figure 202. Rhizococcus turkmenicus (Archangelskaya, 1930), female. After Archangelskaya

Rhizococcus variabilis (Goux, 1940) (Fig. 203) Redescription, Combination

nova
Eriococcus variabilis Goux, 1940: 62.
Holotype: Female. France (Marseilles), on Cynodon dactylon, ?.vi.1934, by L. Goux

(type no. 977/b). Deposited in MNHN. Notes:. Paratypes: 23 slide with the same
data as holotype (MNHN 14527-02-24), and additionally 1 slide in MNHN 8756,
Marseille, St-Tronc (B.d.R), by L. Goux, 19. XI. 1939, on Cynodon dactylon.
Lit.: Foldi, 2001: 305; Kaydan, et al., 2005: 400; Kaydan, et al., 2007: 90; Miller &
Gimpel, 1996: 605; 2000: 374; Ouvrard & Kozár, 2009: 102 (Miller et al., 2013).
Eriococcus greeni; Goux (1948: 69) synonymy with R. greeni.

Acanthococcus variabilis; Danzig & Kozár, 1974: 10. Reestablishment and change of
combination.
Eriococcus variabilis; Goux, 1989: 21. Revived combination.
Acanthococcus variabilis; Kozár, 2009: 94. Revived combination.
Description
Unmounted female
Sac is elongate, closely felted, whitish in color, but rapidly becoming beige and even
ochraceous. Adult female is yellowish olive, elongate.
Mounted female
length of segments: I: 34–43, II: 27–31, III: 80–92, IV: 19–21, V: 17–28, and VI: 29–38
µm long; each segment covered with a few, strong hair-like setae; apical segment with
apical seta 34–38 µm long; apical segment also with 3 sensory falcate setae, longest 22–26
µm long; segment V with 1 sensory falcate seta 24–32 µm, segment IV with 1 sensory
falcate seta 14 µm long. Frontal tubercle and frontal lobe not seen. Eyes situated on
venter near margin. Anal lobes well developed, each with three enlarged setae on dorsal
surface, outer setae stronger (60–67 µm long) than those two on inner side (32–46 µm
long); enlarged setae similar to other dorsal setae; anal lobe setae each 241–335 µm long.
Venter: Labium 3 segmented, 77–90 µm long, basal segment with two setae present on
each side. Stylet loop as long as labium. Legs well developed; lengths of segments and
digitules of prothoracic legs; coxa: 47–55, trochanter: 44–48, femur: 118–137, tibia: 88–
103, tarsus: 108–115, claw: 24–35, tarsal digitules: 39–45, claw digitules: 31–40 µm long;
lengths of segments and digitules of mesothoracic legs; coxa: 52–62, trochanter: 35–48,
femur: 124–142, tibia: 98–127, tarsus: 124–158, claw: 24–26, tarsal digitules: 41–45, claw
digitules: 31–35 µm long; lengths of segments and digitules of metathoracic legs; coxa:
62–71, trochanter: 40–48, femur: 124–142, tibia: 103–127, tarsus: 124–158, claw: 26–29,
tarsal digitules: 41–46, claw digitules: 40 µm long. Tarsal and claw digitules knobbed,
Acanthococcidae 571
Figure 203. Rhizococcus variabilis (Goux, 1940), female. Original.

slightly longer than claw; claw with denticle at apex. Median and posterior coxae without
spinulae; posterior coxae with some small pores on posterior surface. Tibiae each with 5
setae (median seta present). Diameter of anterior spiracles 28–31 µm, posterior spiracles
slightly larger than anterior. Derm with a few short hair-like setae; a row of enlarged
setae seen on margin with a band of macrotubular ducts in marginal position, and
additional sparse enlarged setae band on submarginal area. Multilocular pores each 4–6
µm in diameter and with 3-5 loculi, mostly 5, distributed in sparse rows on all abdominal
segments, scattered on thorax and head. Macrotubular ducts of two sizes, larger ones
on marginal area, narrower ones about 3–5 µm wide, 16–18 µm long, scattered all body.
Microtubular ducts sparse submarginally and marginally. Cruciform pores in a sparse
band on submargin, 5 µm in size.
Dorsum: Marginal dorsal enlarged setae conical spine-like, apices acute, 53–67 µm long,
marginal setae slightly larger than others on dorsum, setae forming 3 longitudinal lines on
each side of abdomen; each abdominal segment with 3 or 4 enlarged setae, other dorsal
enlarged setae various sizes, 11–34 µm long, in sparse rows across all segments; setae on
the middle part of thorax larger than others dorsal setae. Macrotubular ducts 9–11 µm
wide and 20–22 µm long, situated among enlarged setae in bands and rows. Microtubular
ducts 4–5 µm long, with 2 sclerotized areas, scattered. Anal ring strongly sclerotized, with
partly double rows of pores; slightly oval, 53–57 µm wide, 58–71 µm long, with 4 pairs
of long setae, 98–115 µm long. Cauda not seen (Redescription is based on type material).
Ecology
Distribution: France, Turkey.
Biology: Sac formation has been observed in June and November and thus has two
generations a year (Goux, 1940).
Comments
Goux (1948) considered E. variabilis to be a junior synonym of E. greeni, however in 1989
he treated it again as valid (Goux, 1989), his latest decision is accepted by us.
Acanthococcidae 573
Rhizococcus zernae (Tereznikova, 1977) (Fig. 204) Combination nova

Acanthococcus zernae Tereznikova, 1977: 571.
Holotype: Female. Ukraine (Crimea, Belogorsky District), on Zerna cappadocica, by

E. Tereznikova. By original designation. Deposited in Kiev: Notes: 1 paratype with
same data in ZMAS.
Lit.: Kaydan et al., 2007: 90; Kozár et al., 2013: 56; Kozár & Walter, 1985: 74; Köhler,
1998: 386; Miller & Gimpel, 2000: 377; Tereznikova, 1981: 43; Ülgentürk et al., 2003:
Eriococcus zernae; Miller & Gimpel, 1999: 215. Change of combination.

Acanthococcus zernae; Kozár, 2009: 94. Revived combination.
Description
Unmounted female
Adult female body is egg-shaped. Eggsac with wooly waxy needles.
Mounted female
Body elongate oval, 2–3 mm long, 1.2 mm wide. Antenna 7 segmented. Frontal tubercle
present. Eyes situated on venter near margin. Anal lobes well developed, each with three
enlarged setae on dorsal surface, about equal sized, similar to other marginal setae.
Venter: Labium well developed, stylet loop short not reaches the median legs. Legs well
developed; narrow. Tarsal and claw digitules knobbed, slightly longer than claw; claw with
denticle at apex, posterior coxae with small pores on posterior surface. Tibiae each with 5
setae (median seta present). Derm with a few hair-like setae; a row of enlarged setae seen
on margin with a band of macrotubular ducts in marginal position. Multilocular pores
each 6–7 µm in diameter and with 3-7 loculi, mostly quinqueloculars, distributed in sparse
rows on all abdominal segments, scattered on thorax and head. Macrotubular ducts of
two sizes, larger ones on marginal area, narrower ones scattered all body. Microtubular
ducts not shown. Cruciform pores sparse on margin.
Dorsum: Marginal dorsal enlarged setae conical spine-like, pointed, marginal setae
conspicuously larger than other setae on dorsum of abdomen, medial and mediolateral
setae on head approximately same size as marginal setae, medio and mediolateral setae on
thorax and abdomen smaller than marginal setae, of various sizes, in sparse rows across
all segments; each abdominal segment with 3 or 4 enlarged setae. Macrotubular ducts
situated among enlarged setae in bands and rows. Microtubular ducts scattered. Anal
ring with partly double rows of pores. Cauda not seen (after Tereznikova (1977), with
modifications based on paratype).
Other stages
Fist instar nymphs

Described by Tereznikova (1977). All spines on the dorsum much shorter than marginal
ones, antennae 6 segmented, stylet loop reaching posterior coxae, macrotubular ducts
absent, microtubular ducts and discoidal pores not shown.
Second instar male (?) nymphs

Described by Tereznikova (1977). All spines on the dorsum are much shorter, than
marginal ones, antennae six segmented, macrotubular ducts present in two rows on
dorsum, microtubular ducts not shown, some discoidal pores present on venter.
Ecology
Host plant: Artemisia vulgaris, Agropyron repens, Triticum turanicum, Zerna cappadocica.
Distribution: Hungary, Turkey, Ukraine.
Acanthococcidae 575
Figure 204. Rhizococcus zernae (Tereznikova, 1977), female, first instar on top right. After
Tereznikova (1981) with modifications.
Rhizococcus zygophylli (Archangelskaya, 1931) (Fig. 205)

Eriococcus zygophylli Archangelskaya, 1931: 72.
Lectotype: Female. No locality on slide: on Zygophyllum fabago, ?.vi.1928, by A.

Archangelskaya. By subsequent designation Danzig, 1996: 522. Notes: Three
paralectotype females on same slide as lectotype. 1 paralectotype with label Middle
Asia, on Zygophyllum sp., ?.vi.1928 (Danzig, 1996). Deposited in ZMAS.
Common name: Syrian bean caper felt scale.
Lit.: Borchsenius, 1950: 123; Danzig, 1962a: 841; 1972b: 341; 1996: 575; Fetykó, et
al., 2010: 296; Lashin, 1956: 114; Tang & Hao, 1995: 519; Tao, 1999: 35; Wang, 2001:
225 (Miller et al., 2013).
Rhizococcus zygophylli; Borchsenius, 1949: 362. Change of combination.

Eriococcus zygophylli; Hoy, 1963: 124. Revived combination.
Rhizococcus zygophylli; Kozár & Walter, 1985: 75. Revived combination.
Acanthococcus zygophylli; Miller & Gimpel, 1996: 605. Change of combination.
Rhizococcus zygophylli; Köhler, 1998: 401. Revived combination.
Eriococcus zygophylli; Miller & Gimpel, 2000: 378. Revived combination.
Rhizococcus zygophylli; Kozár, 2009: 106. Revived combination.
Description
Unmounted female
Adult female oval, brown, 3 mm long, 2 mm wide. Sac is white, up to 4 mm long, 2.75
mm wide (Borchsenius, 1949). Male second instar before pupation enclosed in elongate
white felted sac with an aperture on its hind end. The male sac is comparatively more
flattened than thatthe female. Male imago bright red, pilose, with short stylus and 2 long
white waxy filaments twice as long as the body. Antennae long and hairy, wings colorless
(Archangelskya, 1931).
Mounted female
Body elongate oval. Antennae 7 segmented, length of segments: I: 51, II: 46.9, III: 77.1,
IV: 64.2, V: 30.4, VI: 27.7 and VII: 47.7 µm long; each segment covered with a few, strong
hair-like setae (except third segment). Eyes situated on venter near margin. Anal lobes
wide, each with three enlarged setae on dorsal surface, 50 µm long, outer setae stronger
than those two on inner side; enlarged setae similar to other marginal setae; anal lobe seta
300, subapical setae 100 and others 60 µm long.
Venter: Labium 3 segmented. Stylet loops almost reaching median coxae. Legs well
developed; femur 225, tibia 180, tarsus 200 µm long. Tarsal and claw digitules knobbed,
slightly longer than claw; claw with denticle at apex, posterior coxae with small pores
on posterior surface. Tibiae each with 5 setae (median seta present). Derm with a few
hair-like setae, preanal setae 60 µm long; a row of enlarged setae seen on margin and
Acanthococcidae 577
Figure 205. Rhizococcus zygophylli (Archangelskaya, 1930), female. After Tang & Hao
submargin, with a band of macrotubular and microtubular ducts in marginal position.

Multilocular pores quinqueloculars, distributed in bands on all abdominal segments,
scattered on thorax and head. Macrotubular ducts of one size, about 8 µm wide, 24 µm
long, scattered over body. Cruciform pores not mentioned.
Dorsum: Marginal dorsal enlarged setae conical spine-like, pointed, some curved, 40–60
µm long, each abdominal segment with 2 enlarged setae, other dorsal enlarged setae
various sizes, 10–14 µm long, in numerous rows across all segments. Macrotubular
ducts 11 µm wide and 24 µm long, situated among enlarged setae in bands and rows.
Microtubular ducts 5 µm long, scattered. Anal ring with partly double rows of pores;
with 4 pairs of long setae, 130 µm long. Cauda not shown (Borchsenius (1949); Danzig
(1962a), with modifications).
Ecology
Host plant: Artemisia sp., Euphorbia sp., Alhagi maurorum, Dodartia orientalis, Zygophyllum
fabago.
Distribution: China, Mongolia, Romania, Turkmenistan, Uzbekistan.
Biology: This species has 2 generations per year, found in the vicinity of Samarkand
and Old Bokhara on roots of Zygophyllum fabago, Artemisia sp. and Dodartia orientalis, in
1927 and 1928 its principal food plant was Z. fabago. Newly hatched first instars spread
around on the ground, are sometimes are found attached to other plants, but in very
small quantities (Archangelskya, 1931).
Acanthococcidae 579
Genus:
Uhleria Cooke, 1881
Type sp.: Eriococcus araucariae Maskell, by monotypy. Synonymy by Morrison
& Morrison, 1966: 200. Notes: This name was first used by Cooke (1881) in the
combination "Uhleria araucariae, Comstock". This is the first use of the generic
name Uhleria and therefore must be attributed to Cooke. Apparently, Cooke was in
contact with Comstock, but Comstock changed his mind about the use of Uhleria
and treated it as a senior synonym of the armored scale genus Fiorinia. Comstock
(1883) presented an invalid nomenclatural scenario making Uhleria a valid armored
scale name and included three species currently considered to be in the genus Fiorinia.
Thus, Comstock's description of Uhleria is a junior homonym of Uhleria Cooke.
Uhleria Cooke is here considered a junior subjective synonym of Eriococcus. Even
though there was no formal description of a generic name, according to Article: 12
(b) (6) of the ICZN, Uhleria was validly described by Cooke by indication.
Lit.: Cooke, 1881: 41; Fernald, 1903: 71; Hoy, 1963: 132; Kozár, 2009: 113; Miller &
Miller, 1992:9; Miller & Gimpel, 2000: 384; Morrison & Morrison, 1966: 201 (Miller
et al., 2013).
Description
Adult female elongate-oval, narrowed posteriorly, with anal lobes conical and normally
heavily sclerotized, antennae 7 segmented; frontal lobes present, labium 3 segmented,
with well developed segments and a weakly developed basal segment with two pairs
of hair-like setae; legs well-developed, midcoxae and hindcoxae often with spinulae on
anterior surfaces, translucent sensory pores present, inner side of hind tibia with only
four setae, claw usually with a denticle; tarsal and claw digitules longer than claw, spiracles
often with a few associated disc pores; discoidal pores usually quinquelocular, cruciform
pores on prosomal venter in a marginal band; macrotubular ducts of two sizes on venter;
larger ones generally situated on the margin of venter, and form longitudinal marginal
band or row; or in transverse sparse bands on dorsum, smaller macrotubular ducts
situated on the middle of abdominal segments and submargin on venter; microtubular
ducts slender with special, bitubular orificies, scattered or form transverse rows or bands
on dorsum; marginal enlarged conical, slightly blunted or truncuted, dorsal setae small,
generally as one third of the enlarged marginal setae and truncated or blunted, where they
form transverse bands or rows; hair-like setae on venter only; anal ring well developed,
sclerotized with partly double rows of pores and 8 anal ring setae, latter often as long as
apical seta on anal lobes; each anal lobe with a long apical seta and usually with 3 elongate
dorsal conical setae, at least with 2 ventral hair-like setae also present; suranal setae hair-
like, cauda conspicuous.
Distribution
Unknown origin, probably Neotropic and Australian, in the Palaearctic Region with
probably 2 introduced species.
Biology
Feed on plants from Araucariaceae and Myrtaceae.
Comments
The structure of the microtubular ducts of the females and the dorsal spines of first
instar nymphs are quite different from the common and species rich genera in the
Palaearctic Region as Acanthococcus and Rhizococcus.
Key to species
1. – On the abdominal margin with 2 long enlarged setae................................U. mariannae

– On the abdominal margin with 2 long and 1 small enlarged setae........... U. araucariae
Uhleria araucariae (Maskell, 1879) (Fig. 206) Revived combination

Eriococcus araucariae Maskell, 1879: 218.
Syntype: Female. New Zealand (South Island, Governor's Bay, near Christchurch),
1879. By original designation. Deposited in NZAC and USNM.
Common name: Araucaria Scale, Felted Pine Coccid, Norfolk Is. Pine Eriococcin.
araucaria mealybug, Norfolk Island pine scale.
Lit.: Afifi, 1968: 167; Balachowsky, 1939: 267; Ben-Dov, 1985a: 187; Borchsenius,
1949: 361; Danzig, 1971a: 822; De Lotto, 1967: 117; Fernald, 1903: 71; Ferris, 1955:
95; Foldi, 2001: 305, 307; Froggatt, 1900: 101; Fuller, 1901: 107; Gill, 1993: 156;
Gómez-Menor Ortega, 1937: 350; Hodgson & Miller, 2002: 193; Goux, 1944: 137;
Green, 1922: 347; Hodgson & Miller, 2010: 99; Köhler, 1998: 396; Kondo et al., 2006:
34; Kozár et al., 2009: 1; Maskel, 1887: 93; Mestre Novoa, 2011: Miller & Gimpel,
2000: 384; Miller & González, 1975: 138; Miller & Miller, 1992:9; Ouvrard & Kozár,
2009: 102; Pellizzari & Germain, 2010a: 478; Pellizzari & Kozár, 2011: 66; Tang &
Hao, 1995: 520, 654; Williams, 2007: 1352; Williams & Watson, 1990: 216 (Miller et
al., 2013).
Uhleria araucariae; Cooke, 1881, 41. Change of combination.

Rhizococcus araucariae; Comstock, 1881a: 339. Change of combination.
Nidularia araucariae; Lindinger, 1933a: 108. Change of combination.
Eriococcus araucariae; Hoy, 1962: 31; 132. Revived combination.
Acanthococcus araucariae; Tereznikova, 1981: 19. Change of combination.
Acanthococcidae 581
Eriococcus araucariae araucariae; Miller & Gimpel, 2000: 384. Revived combination and
change of status.
Eriococcus araucariae; Miller & Gimpel, 2000: 384. Revived combination.
Uhleria? araucariae; Kozár, 2009: 113. Revived combination.
Acanthococcus araucariae; Hodgson & Miller, 2010: 99. Revived combination.
Eriococcus araucariae var. minor; Maskell, 1895a: 21.
Syntypes: Female. Australia (New South Wales), Manly, on Kunzea capitata. Deposited
in NZAC and USNM.
Lit.: Fernald, 1903: 71; Hoy, 1963: 71; Miller & Gimpel, 2000: 130; Williams, 2007:
1353 (Miller et al., 2013).
Eriococcus araucariae minor; Hoy, 1963: 71. Change of status.
Acanthococcus araucariae minor; Miller & Gimpel, 1996: 598. Change of combination.
Uhleria? araucariae minor; Kozár, 2009: 107. Change of combination.
Description
Unmounted female
Newly mature adult female brownish-yellow with 1 pair of purple stripes on sublateral
area of dorsum; old females purple. Eggs are yellow. Ovisac felted, oval, white and
covering body.
Mounted female
Body elongate oval, 1.86–2.61 mm long, 0.80–1.40 mm wide. Antennae 7 (rarely 6)
segmented, apical segment with apical seta and 3 sensory falcate setae. Frontal lobes
present. Eyes situated on venter near margin. Anal lobes large protruding, with entire
lobe area heavily sclerotised; each lobes with 3 enlarged setae on dorsal surface, all nearly
equal in size, with 6–10 microtubular ducts; each lobe ventrally with 4 slender body setae
and 0-9 sessile pores.
Venter: Labium 3 segmented, with 8 pairs of hair-like setae. Stylet loop long, reaching
the abdominal segment IV. Legs well developed; hind coxa dorsally with 48-84 pores,
ventral surface with 92–117; hind femur dorsally with 0-8 pores, absent ventrally; hind
2 pairs of tibia with 4 setae, front tibia with 5; tarsus and tibia nearly equal (hind tibia/
tarsus ratio 0.95–1.00); claw with very small denticle near tip. Setae lanceolate body setae
abnormally elongate (longest setae on abdominal segment VII from 59–84 µm long, on
segment II from 94–147 µm long), medial setae apically acute. Enlarged setae absent.
Multilocular pores of two kinds. Quinquelocular pores present on abdomen, absent in
medial areas of segments IV through III, present on thorax, rare or absent on head;
trilocular pores abnormally abundant, almost completely replacing quinquelocular pores
on anterior abdominal segments, thorax, and head. Cruciform pores small, difficult to
distinguish, if present, scattered along body margin. Macrotubular ducts of 2 sizes; larger
size same as on dorsum, restricted to body margin; smaller size with flattened “cup”,
present in medial or sublateral areas of abdominal segments VII through III, most
numerous on posterior abdominal segments Microtubular ducts absent.
Dorsum: Enlarged setae of 2 sizes; larger one present along entire body margin, with two
or three present on margin of each abdominal segment; smaller size restricted to medial
and sublateral areas. Largest lateral setae 28–32 µm long, largest cup shape setae 6–7 µm
long; on abdominal segments VII through II longest lateral setae 4.3–4.8 times longer
than longest medial and sublateral setae. Lateral setae slender, with blunt apices; medial
setae small, with blunt apices; setal rings thin. Enlarged setae in moderate numbers with
no longitudinal pattern. Macrotubular ducts scattered on dorsum. Microtubular ducts
long (8–10 µm long), with area farthest from dermal orifice lightly sclerotised and weakly
divided into two parts, apical portion blunt or rounded, from 1/4–1/2 of remaining
sclerotised portion; total sclerotised area approximately equal to unsclerotised area;
dermal orifice large, heavily sclerotised, with 2 protruding tubes, numerous over dorsum.
Anal ring usually with 4 pairs of setae. Cauda present (after Miller & Miller, 1992, with
modifications).
Other stages
First instar and adult male described by Leonardi (1920) (Fig. 206 on top right).
Ecology
Host plant: Araucaria sp., A. angustifolia, A. araucata, A. bidwillii, A. columnaris, A.
cunninghamii, A. heterophylla?, A. hunsteinii, A. orientalis?, Cupressus sp.(?), Cycas sp.(?), Eleusine
indica (?), Juniperus sp.(?), Kunzea capitata (?), Pinus pinea (?).
Distribution: Kenya, Mauritius, South Africa, Zimbabwe, Australia, Cook Islands,
Fiji; New Caledonia, New Zealand, Papua New Guinea, Vanuatu, Mexico, USA,
Argentina, Brazil, Chile, Costa Rica, Cuba, Czech Republic (new? record in 2011 from
Milena Breziková), Guatemala, Nicaragua, Panama, Puerto Rico & Vieques Island,
Uruguay, Venezuela, India, Malaysia, Philippines, Sri Lanka, Algeria, Croatia, Czech
Republic, Egypt, France, Germany, Greece, Iran, Israel, Italy, Malta, Monaco, Morocco,
Netherlands, Portugal, Portugal (Azores, Madeira), Russia, Spain, Spain (Canary Islands),
Turkey, United Kingdom, United Kingdom (Bermuda), USA (Hawaiian Islands) former
Yugoslavia.
Biology: On the lower surface of leaves, twigs, and in twig axils. Pest. Adult males are
active in August. The collection data indicate two generations per year. In the United
States, it occurs out-of-doors only in the warmer parts of the country. The species has
a worldwide distribution wherever Araucaria is grown. The large amounts of honeydew
produced by the insect provides a medium for sooty molds which often blacken the host
foliage.
A widespread ornamental pest, U. araucariae has been carried all over the world, but is
most likely of Australian and Nearctic origin.
Comments
The species well known and widely distributed on different Araucaria species, other host
plant records needs verification.
Acanthococcidae 583
Figure 206. Uhleria araucariae (Maskell, 1879), female. After Miller & Miller (1993).
Uhleria mariannae (Pellizzari, 2010) (Fig. 207) Combination nova

Acanthococcus mariannae Pellizzari in Pellizzari & Germain, 2010b: 52.
Holotype: Female. Italy (Genova), on Leptospermum scoparium, 8.viii.2004, collector

unknown. Slide No. 1128/1. By original designation. Deposited in DEAE and
paratypes deposited in MNHN, and at Montferrier-sur-Lez, France at Laboratoire
National de la Protection des Végétaux, France.
Lit.: Pellizzari & Kozár, 2011: 66 (Miller et al., 2013).
Uhleria ? nr. araucariae sp. n. France (Corsica, Port. It.); Kozár, 2009: 107. Notes: The
studied slides originates from the type series from which some were sent by Authors to
F. Kozár for study, and the drawing is based on them.
Description
Unmounted female
Adult female elongate, oval, grey-brown; abdomen with distinct segmentation. Females
settled on upper surface of leaves or on twigs of host plant. Before egg-laying, the adult
females become enclosed in a felted, white, lightly convex eggsac, open only at anal end.
Male test oval, white, on under surface of leaves. Adult males winged.
Mounted female
Body elongate oval, 1.44–1.96 mm long, 0.48–0.96 mm wide. Post-reproductive female
2.0–2.35 mm long, 1.14–1.2 mm wide. Total length of each antenna 140–160 µm. Scape
with 3 setae, 2nd segment with 2 setae and 1 sensory pore, segment III without setae,
segment IV with 2 setae, segment V with 1 fleshy seta, segment VI with 1 fleshy seta + 2
setose setae, segment VII with 3 fleshy setae + 7 flagellate or hair-like setae. Frontal lobes
present. Eyes situated on venter near margin. Anal lobes protruding, slightly sclerotised,
each 60–74 µm long, each lobe dorsally with 3 enlarged setae and 3 or 4 microtubular
ducts; ventrally with 2 hair-like setae. Apical seta 110–136 µm long.
Venter: Labium 3 segmented, with 2 pairs of almost equal long hair-setae on unsclerotised
basal segment, one pair on middle segment and 5 pairs on apical segment. Stylet loop
slightly exceeding level of second coxae. Legs well developed; hind coxa and femur with
translucent pores and spinulae, median coxae with spinulae; claw with small denticle.
Tarsal digitules knobbed; claw digitules longer than claw, knobbed. Measurements of
metathoracic leg; coxa: 40–46 µm long; trochanter + femur: 106–136 µm long; tibia:
50–68 µm long; tarsus: 84–96 µm long; claw 18 µm long. Tibia normaly with four setae.
Body setae: ventral setae hair-like, with 2 on median part of each abdominal segment,
more numerous near coxae on thorax and on head; enlarged setae situated submargin in
longiditual row. Minute hair-like setae sparse on abdominal segments. Multiloclular pores
quinquelocular and 3–4 µm wide, numerous, forming transverse bands on posterior
abdominal segments, becoming less abundant anteriorly and sparse on head and on
Acanthococcidae 585
Figure 207. Uhleria mariannae (Pelizzari & Germain, 2010), female, original. First instar on
top right, after Pellizzari & Germain (2010) with modifications.
medial and submarginal parts of thorax; also with 2-6 laterad to each spiracle opening.
Cruciform pores few, each 3–4 µm long, present on submargin of thorax and head.
Macrotubular ducts smaller than dorsal ducts, 18 µm long, 6 µm wide, mainly distributed
along body margin and submarginally. A few very small macrotubular ducts, each 11–16
µm long and 2.5 µm wide, with a sclerotised ring 4 µm in diameter and a very long inner
filament, intermingled with quinquelocular pores on posterior abdominal segments.
Microtrichia present on abdominal segments.
Dorsum: Enlarged setae small, 5.0–6.5 µm long and 3–4 µm wide at base, distributed in
an irregular row of 7-12 setae across each abdominal segment; also sparse on thorax and
head. Dorsal macrotubular ducts large, about 20 µm long and 10 µm wide, with a well-
developed sclerotised rim and inner ductule; numbering 10-20 across each abdominal
segment, becoming less abundant posteriorly, and sparse on head and thorax. Microducts
numerous, scattered, 6.5 µm long. Anal ring with 4 pairs of setae and with an outer
row of cone-shaped pores. Cauda present, triangular in shape, with an irregular margin.
Margin and dorsal surface reticulatated with microtrichia (after Pellizzari & Germain
(2010b), with modifications).
Other stages
First instar nymph

with strong falcate, sensory setae as in adult female. Frontal tubercle present. Dorsum
with strong, short almost triangle-shaped spine-setae in four median and a marginal,
longitudinal rows. The marginal row of setae consist of one strong blunted spine on
last abdominal segments. Microtubular ducts in a marginal row on abdomen and two
on midthorax. Venter with transverse rows of four small hair-like median setae on
each abdominal segment; and a row of small submarginal setae. Two cruciform pores
present on thorax margin of venter. With one pair of trilocular pores on each abdominal
segment, and one pair near each spiracle, plus some on frons, and one pair in middle-line
of thorax. Stylet loop reaching the median coxae. Anal lobe with two conical setae and
with a long apical setae. Anal ring normal, with 6 hair-like setae (Pellizzari & Germain,
2010b).
Ecology
Host plant: Leptospermum scoparium.
Distribution: France (Corsica), Italy, (It seems almost certain that A. mariannae was
accidentally introduced to Europe on Leptospermum sp., and the native region remains
unknown.
Biology: The young adult females and the ovipositing females, enclosed in their white
felted eggsac, settle on the under surface of leaves, on the axil of leaves and twigs, and
along the thin twigs of Leptospermum scoparium. The collection data for A. marianne suggest
that this species could develop several overlapping generations.
Acanthococcidae 587
Fossil genera and species
Genus:
Balticococcus Koteja, 1988
Type sp.: Balticococcus oblicus Koteja, 1988, 512, by monotypy and original designation.
Notes: We can see no basis for separating this genus from Eriococcus and note that
Koteja (1988a) writes "I am quite aware that the described forms may belong to some
recent genera or may even be synonymous with some recent species." (Miller et al.,
2013).
Lit.: Koteja, 1988a: 512; 2000b: 207; Miller & Gimpel, 2000: 72 (Miller et al., 2013).
Description
First-instar nymph with: dorsal enlarged setae across abdominal segments; about 5 setae
on apical antennal segment (Koteja, 1988b). The description of this genus is based on
the first instar only.
Host plant
Unknown. Found in Baltic Amber.
Distribution
Denmark, Poland.
Balticococcus oblicus Koteja, 1988

Balticococcus oblicus Koteja, 1988b: 512-514.
Holotype: Immature. Denmark (Jutland, Baltic amber), 03/05/1952, by C.V.

Henningsen., by original designation. Deposited in ZMUC.
Lit.: Koteja, 2000b: 207; Kozár, 2009: 96; Miller & Gimpel, 2000: 72 (Miller et al.,
2013).
Description
First instar in amber with: enlarged setae conical, slightly curved, sides slightly concave,
apices acute, all setae of approximately same size, mediolateral setae present from head
to segment 7, medial setae present on head to segment 1, 1 lateral seta on margin of each
abdominal segment; anal lobes each apparently with 3 enlarged setae (Koteja, 1988b).
Very similar to many first instars of contemporary species of Eriococcus s. l.
Ecology
Host plant: Unknown. Found in Baltic Amber.
Distribution: Denmark.
Comments
Very similar to many first instars of contemporary species of of Acanthococcidae (like
species in Acanthococcus).
Balticococcus spinosus Koteja, 1988

Balticococcus spinosus Koteja, 1988b: 514-517.
Holotype: Immature. Poland (Baltic amber, near Gdansk). by original designation.

Deposited in Warsaw: Museum of Earth, Polish Academy of Sciences, Poland; type
no. 15493.
Lit.: Koteja, 1998: 194; 2000b: 207; Kozár, 2009: 96; Miller & Gimpel, 2000: 72-73.
Description
First instar in amber with: enlarged setae conical, slightly curved or straight, sides slightly
concave or straight, apices slightly rounded, all setae of approximately same general size
except decreasing anteriorly and mediolateral setae on segments 7 and 6 largest, setae
forming 3 complete longitudinal lines on each side of body, 1 lateral seta on margin of
each abdominal segment; anal lobes each with 3 enlarged setae (Koteja, 1988b).
Ecology
Distribution: Poland.
Comments
Very similar to many first instars of contemporary species of Acanthococcidae (like
species in Hujinlicoccus genus).
Acanthococcidae 589
Genus:
Gedanicoccus Koteja, 1988
Types.: Gedanicoccus gracilis Koteja, 1988, by monotypy and original designation. Notes:
We can see no basis for separating this genus from Eriococcus and note that Koteja
(1988a) wrote "I am quite aware that the described forms may belong to some recent
genera or may even be synonymous with some recent species."
Lit.: Koteja, 1988b: 506-507; 2000b: 207; Miller & Gimpel, 2000: 383 (Miller et al.,
2013).
Description
Generic characteristics that distinguish this genus from all other eriococcids are: the
possession of dorsal medial plate; claw without denticle; dorsal setae inconspicuous or
absent; marginal setae enlarged (Koteja, 1988b). The description of this genus is based
on the first instar only.
Host plant
Distribution
Poland.
Gedanicoccus gracilis Koteja 1988

Gedanicoccus gracilis Koteja, 1988c: 507-512.
Holotype: Immature. Poland (Baltic Amber, Baltic Shore, near Gdansk), by T.

Giecewicz. by original designation. Deposited in Warsaw: Museum of Earth, Polish
Academy of Sciences, Poland; type no. 15494.
Lit.: Koteja, 1988b: 507-512; 2000b: 207; Kozár, 2009: 102; Miller & Gimpel, 2000:
383-384 (Miller et al., 2013).
Description
Body elongate, the presence of stellate hairs indicates a spring hatching. First instar in
amber with: enlarged setae conical, sides slightly convex, apices slightly rounded, enlarged
setae confined to marginal area with 1 lateral seta on margin of each abdominal segment
(Koteja, 1988b).
Ecology
Comments
species in Rhizococcus genus).
Genus:
Jutlandicoccus Koteja, 1988
Type s.: Jutlandicoccus perfectus Koteja, 1988b: 517-518. by monotypy and original
designation. Notes: We can see no basis for separating this genus from Eriococcus s.l.
and note that Koteja (1988b) writes "I am quite aware that the described forms may
belong to some recent genera or may even be synonymous with some recent species."
Lit.: Koteja, 1988b: 517-518; 2000b: 207; Miller & Gimpel, 2000: 388 (Miller et al.,
2013).
Description
Body is broadly oval, anal lobes tuberculiform, apically rounded with nodulate surface;
dorsal medial plate present. Generic characteristics that distinguish this genus from all
other eriococcids are: the possession of dorsal medial plate; tibia shorter than tarsus;
claw without denticle; dorsal setae hair-like; marginal setae enlarged (Koteja, 1988b). The
description of this genus is based on the first instar only.
Host plant
Distribution
Denmark.
Acanthococcidae 591
Jutlandicoccus pauper Koteja, 1988

Jutlandicoccus pauper Koteja, 1988b: 518-520.
Holotype: Immature. Denmark (Jutland, Baltic amber), 01/11/1964. by original

designation. Deposited in Copenhagen: Zoological Museum, University of
Copenhagen, Department of Entomology, Denmark. Illust.
Lit.: Koteja, 1988b: 518; 2000b: 207; Kozár, 2009: 102; Miller & Gimpel, 2000: 388.
Description
First instar in amber with: enlarged setae conical, slightly curved, sides slightly
concave, apices acute, marginal setae and setae in mediolateral areas of head and
thorax conspicuously longer that other dorsal setae, mediolateral setae on abdomen
and metathorax smaller, decreasing in size posteriorly, 1 lateral seta on margin of each
abdominal segment except on segment 7 which has 2 (Koteja, 1988b).
Ecology
Comments
species in Rhizococcus genus).
Jutlandicoccus perfectus Koteja, 1988

Jutlandicoccus perfectus Koteja, 1988b: 520-523.
Holotype: Immature. Denmark (Jutland, Baltic amber), 09/09/1974, by C.V.

Henningsen., by original designation. Deposited in Copenhagen: Zoological Museum,
University of Copenhagen, Department of Entomology, Denmark.
Lit.: Koteja, 1988b: 520; 2000b: 207; Kozár, 2009: 102; Miller & Gimpel, 2000: 389
Description
First instar in amber with: enlarged setae conical, slightly curved, sides slightly concave,
apices acute, marginal setae slightly longer that other dorsal setae, mediolateral setae
present on thorax, medial setae present on head to segment 7, slightly decreasing in size
posteriorly, 1 lateral seta on margin of each abdominal segment except on segment 7
which has 2; anal lobes each apparently with 2 enlarged setae (Koteja, 1988b).
Ecology
Comments
species in Acanthococcus genus).
Genus:
Kuenowicoccus Koteja, 1988
Type sp.: Kuenowicoccus pietrzeniukae Koteja, 1988: 182, by monotypy and original
designation.
Lit.: Koteja, 1988a: 405; 2000b: 208; Kozár, 2009: 103; Miller & Gimpel, 2000: 389
Description
Generic characteristics that distinguish this genus from all other eriococcids are: the
possession of 10 setae on the scape of the antenna and the shape of the penial sheath
which evenly tapers to a broad truncate tip (Koteja, 1988a). The description of this genus
is based on the male only.
Host plant
Unknown. Found on Baltic amber.
Distribution
Poland.
Acanthococcidae 593
Kuenowicoccus pietrzeniukae Koteja, 1988

Kuenowicoccus pietrzeniukae Koteja, 1988a: 405.
Holotype: Male. Poland (Gdansk, in amber), 10/11/1986. by original designation.

Deposited in ZMHB; type no. 393.
Lit.: Koteja, 1988a: 406; 2000b: 208; Kozár, 2009: 103; Miller & Gimpel, 2000: 389
Description
See under generic description.
Ecology
Host plant: Unknown. Found in Baltic amber.
Biology: Koteja (1988a) considers this species to be extinct.
Family:
CRYPTOCOCCIDAE
Kosztarab, 1967
Type genus: Cryptococcus Douglas, 1890.
Common name: Bark-crevice scale.
Lit.: Danzig, 1980: 234; Howell & Williams, 1976:187; Koteja, 1974b: 50; Kosztarab
& Kozár, 1978: 82; Kozár & Konczné Benedicty, 2008: 247; Morrison & Morrison,
1966: 50; Schmutterer, 2008: 84; Tang & Hao, 1995: 426; Tereznikova, 1975; Williams,
1976: 187 (Miller et al., 2013).
Dactylopiidae (in part) Ferris, 1955:69.

Eriococcidae (in part) Hoy, 1963:53.
Cryptococcidae Kosztarab, 1968:12.
Description
Female oval, derm sclerotized at full maturity. Body elongate to broadly oval, usually
membranous; antennae 1-6 segmented; clypeolabral shield pentagonal, clypeal setae
absent, labral ridge very short, marginal ridges strong; labium short and broad, 3
segmented, with 8 pairs of setae; legs if present, well developed, tarsus terminating in one
claw, claw with denticle, or without, and with a pair of claw digitules, trochanter with one
pair of campaniform sensillae (pores) on both sides, ostioles and circuli often present;
spiracles surrounded by a sclerotized frame, with a group of associated quinquelocular
pores, normally a cluster pore plate just below each posterior spiracle; anal ring heavily
sclerotized, with 4-6 short setae or hairs, with or without pores, surrounded by large setae;
no cruciform pores; simple disc pores in dorsal submargin, normally trilocular pores on
dorsum, quinquelocular pores on both derms, seldom quadrilocular pores scattered near
spiracles; tubular ducts on both body surfaces, often of two types. Anal lobes often
developed; anal ring usually dorsal, with 2 rows of pores and 6 setae.
Host plant
Woody plants from Aceraceae, Fagaceae, Oleaceae, Rosaceae, Salicaceae, Tiliaceae.
Distribution
Holarctic, with an exception, C. nudatus Brittin, known only in New Zealand. In Palaearctic
Region 6 species in two genera.
Cryptococcidae 595
Biology
Adult females in bark crevices of deciduous trees, covered with fine, white, waxy
secretion or felt-like ovisac. Apparently with one yearly generation. Adult males present
in Pseudoehermes fraxini only.
Key to genera
1. – Legs developed, hind coxae with translucent pores; no clusters of pore plates
posterior of hind spiracles; antennae 6 segmented................................... Pseudochermes
– Legs absent or reduced to unsegmented stubs; with clusters of pore plates posterior
of each hind spiracle; antennae 1-5-segmented............................................Cryptococcus
Genus:
Cryptococcus Douglas, 1890
Type sp.: Coccus fagi Baerensprung (= Cryptococcus fagisuga Lindinger), by monotypy.
Notes: The type species Coccus fagi Baerensprung, 1849 is a junior primary homonym
of Coccus fagi Sulzer, 1776. This genus was placed in a separate family (Cryptococcidae)
by Kosztarab (1968), but the characters given as diagnostic of the family are typical
of other legless adult female eriococcids.
Lit.: Balachowsky, 1942: 42; Boratynski & Williams, 1964: 91; Borchsenius, 1937: 40;
1949: 31; Cockerell, 1894b: 1050; Danzig, 1964: 632; 1986: 271; 1988: 707; Douglas,
1890: 155; Ferris, 1955 Gill, 1993: 153; Green, 1915: 180; Henderson 2007b: 3-4 Hoy,
1963: 53; Kosztarab, 1968: 12; 1996: 225; Kosztarab & Hale, 1968: 11; Kosztarab
& Kozár, 1988: 269; Koteja, 1974a: 269; 1974b: 78; Kozár, 2009: 114; Kozár &
Drozdják, 1998: 167; Kozár & Walter, 1985: 75; Lindinger, 1937: 183; Miller, 1985:
101; 2005: 491; Miller & Gimpel, 2000: 83; Miller & Miller, 1993: 68; Morrison &
Morrison, 1966: 50; Newstead, 1903: 214; Schmutterer, 1952: 417; 2008: 84; Šulc,
1895: 13; Tang & Hao, 1995: 427; Tereznikova, 1981: 13; 1982: 35; Williams, 1985a:
352; Wise, 1977: 96; Zahradník, 1959: 538; Xie, 1998: 101 (Miller et al., 2013).
Description
Adult females covered with fine, white, waxy secretion. Female oval, 0.5–1.0 mm;
antenna normally 1 or 2, rarely 4 or 5 segmented; labium 3 segmented, basal segment
without setae; legs absent, rarely reduced to unsegmented stubs; spiracles with sclerotized
frame and a few associated quinquelocular pores; with a cluster pore plate below each
hind spiracle; anal ring with 4 or 5 setae, rarely with pores; anal ring forming a broad
plate surround by setae; macrotubular ducts present, except in one species; microtubular
with simple pores, without cruciform pores; tubular ducts, normally two types on both
surfaces.
Host plant
Woody plants from Aceraceae, Fagaceae, Rosaceae, Tiliaceae.
Distribution
Worldwide with 5 species; 4 species known from the Palearctic.
Biology
Live under bark flakes or in crevices of bark. One yearly generation.
Key to species
1. – Anal ring with 4 thick setae................................................................................ C. fagisuga

– Anal ring with 6 thin setae.................................................................................................2
2. – Legs absent, 10 setae surrounding anal ring........................................................C .aceris
– Legs reduced to tubercles, less then 10 setae surrounding anal ring...........................3
3. – Antennae 6 segmented, on Ulmus spp................................................................... C. ulmi
– Antennae 4 segmented, on Tilia spp...........................................................C. integricornis
Cryptococcidae 597
Cryptococcus aceris Borchsenius, 1937 (Fig. 208) Redescription

Cryptococcus aceris Borchsenius, 1937: 62.
lectotype: Female. Azerbaijan (Abkhazia, Gagra District, Bzyb Gorge), on Acer sp.,
1931. By subsequent designation Danzig, 1996: 521. Deposited in ZMAS. Notes:
There are 5 female paralectotypes on the same slide as the lectotype.
Common name: Maple Bark Scale.
Lit.: Borchsenius, 1939: 43; 1949: 48; Cook & Gullan, 2004: 444; Danzig, 1971b:
1415; 1985: 147; 1996: 574; Ferris, 1955: 83; Hadzibejli, 1950: 256, 257, 263; Hoy,
1963: 54; Kosztarab & Hale 1968: 11; Kosztarab & Kozár, 1988: 269; Kozár, 2009:
96; Kozár & Drozdják, 1998: 167; Kozár & Walter, 1985: 75; Lagowska, 2002: 243;
Longo et al., 1999: 145; Miller & Gimpel, 2000: 84; Schmutterer, 1980: 50; Tang &
Hao, 1995: 427; Tereznikova, 1959: 448; Ter-Grigorian, 1962: 132; Williams, 1985a:
352; Zahradník, 1972: 405 (Miller et al., 2013).
Description
Unmounted female
Adult female hemispherical and yellow, completely enclosed within white sac
(Borchsenius, 1949). Kosztarab & Kozár (1988) state that the female is covered by a
white waxy secretion.
Mounted female
Body hemispherical oval, ca. 0.9 mm. Antennae reduced, 4 segmented, size of each
segment as follows: I: 10–12, II: 5–7, III: 4–5 and IV: 10–12 µm long, each segment
covered with a few, strong setae (except second segment); apical segment with 4 sensory
falcate setae, the longest 7–9 µm long. Frontal tubercle and frontal lobes not seen. Eyes
situated on venter near margin. Anal lobes not developed, wide, each with 2 enlarged
setae, 18–25 µm long, on dorsal surface.
Venter: Labium 3 segmented, 38–41 µm long; on basal segment seta not seen, stylet loop
extending to abdominal sternites VII. Legs absent. The diameter of anterior spiracles 7–8
µm; each spiracle with 6-9 associated quinquelocular pores, 4 µm in diameter. Setae short,
conical. Quinquelocular pores few, scattered on submarginal area, a few scattered mid
abdominal segments, each 4 µm in diameter. Macrotubular ducts of two sizes, narrower
ones situated around labium, each about 3 µm wide and 9 µm long, larger ones 5–7 µm
wide and 10 µm long on margin. Microtubular ducts each 3 µm long, scattered through.
Dorsum: Dorsal setae few, short, conical, each spine 3–4 µm long. Macrotubular ducts
present throughout in sparse rows, 5–7 µm wide and 10 µm long. Microtubular ducts 3
µm long, scattered throughout. Discodial pores absent. Anal ring almost circular, 25–30
µm wide, 22–26 µm long, with 5–7 pores and 6 thin, 10–12 µm long setae; anal ring
surrounded by 10 short setae (Redescription is based on type material).
Ecology
Host plant: Acer sp., A. pseudoplatanus, Pyrus sp., Tilia sp.
Distribution: Azerbaijan, China, Germany, Hungary, Russia, Ukraine.
Biology: Feed on trunk of trees, biology poorly known. In South Germany it infested
the trunk of Acer pseudoplatanus (Schmutterer, 1955).
Cryptococcidae 599
Figure 208. Cryptococcus aceris Borchsenius, 1937, female. Original.

Cryptococcus fagisuga Lindinger, 1936 (Fig. 209)

Coccus fagi Baerensprung, 1849: 174.
Type material: Germany (Berlin). Described: female. Homonym of Coccus fagi Sulzer
1776; discovered by Lindinger, 1936: 444. Notes: Type material has probably been
lost (Williams, 1985a).
Common name: Beech coccus; beech eriococcin; beech scale; felted beech coccus.
Lit.: Balachowsky, 1937b: 6; Bareth & van Liefferinge, 1976: 209; Bodenheimer, 1935:
271; 1953: 134; Boratynski & Williams, 1964: 91; Borchsenius: 1949: 371; Cockerell,
1896: 324; Danzig, 1985: 111; Fernald, 1903: 120; Ferris, 1955: 83; Foldi, 2001: 305;
Gertsson, 1997: 113, 114; Green, 1928a: 9; Hadzibejli, 1983: 270; Hoy, 1963: 54;
Jansen, 2001: 200; Kawecki, 1985: 33; Kaydan et al., 2007: 90; Kiritchenko, 1935: 1;
Kosztarab, 1996: 262; Kosztarab & Hale 1968: 11; Kosztarab & Kozár, 1988: 270;
Koteja, 2000a: 242; Kozár, 2009: 96; Kozár & Drozdják, 1998: 167; Kozár, et al.,
1996: 65; Kozár, et al., 1994: 154; Kozár & Kosztarab, 1982: 204; Kozár & Walter,
1985: 75; Kozarzhevskya, 1986: 306; Lindinger, 1958: 367; Masten Milek & Simala,
2008: 105; Miller, 1985: 101; 1991: 101; 2005: 491; Miller & Gimpel, 2000: 85; Miller
& Miller, 1993: 68; Newstead, 1903: 215; Ossiannilsson, 1951: 4; Ouvrard & Kozár,
2009: 101; Pellizzrai & Kozár, 2011: 66; Reh, 1904: 34; Reyne, 1961: 138; Schmutterer,
1952: 378; Šulc, 1895: 13; Swaine & Hutchings, 1926: 47; Tang & Hao 1995: 427,
641; Tereznikova, 1981: 46; 1982: 36; Ter-Grigorian, 1983: 882; Walker, 1852: 1086;
Wünn, 1925c: 438; Zahradník, 1959: 538; 1972: 403 (Miller et al., 2013).
Coccus fagi; Walker, 1852: 1086.

Lectotype: England. Lectotype female, by subsequent designation Williams, 1985a:
353-356. Synonymy by Lindinger, 1936: 444. Homonym. Notes: Coccus fagi Walker
is a junior homonym of Coccus fagi Sulzer, 1776 and Coccus fagi Baerensprung, 1849.
Pseudococcus fagi; Douglas, 1886: 152. Change of combination.
Cryptococcus fagi; Douglas, 1890: 155. Change of combination.
Eriococcus fagi; Perrier, 1926: 122. Described: female. Change of combination.
Cryptococcus fagisuga; Lindinger, 1936: 444. Replacement name for Coccus fagi Baerensprung
1849.
Kermes fagi; Lindinger, 1957: 549. Change of combination. Notes: Lindinger (1957) meant
to reiterate the placement of fagi in Cryptococcus and he cites Guérin (1818) as the creator
of the Kermes fagi combination. The date of this is probably wrong, and no publication
by Guérin mentioning the combination can be found. Therefore, at least for the time
being, this combination will be listed as by Lindinger (1957) even though it is not what
he intended.
Cryptococcidae 601
Description
Unmounted female
Adult female, in life, globular, bright to lemon yellow, completely covered in a filamentous
white ovisac.
Mounted female
Body rotund, membranous, nodulose, largest about 0.5–1.0 mm long, often wider than
long. Antennae reduced, 1or 3 segmented.
Venter: Labium 3 segmented, 48 µm long; often wider than long, shoter than clypeolabral
shield, basal segment without setae. Stylet loop extends to abdominal sternites VII. Legs
absent, but position of each third leg represented by a small flap often containing a
single setae. Spiracles often triangular, sclerotised. Setae about same size as dorsal setae
except on anal lobe segment where the suranal longer setae, and one or two near margins,
are larger and conical, about 10 µm long and usually a pair of slightly longer setae near
anterolateral concerns of anal ring, probably the anal lobe setae. Quinquelocular pores
about same diameter as the cup of large macroduct, in more or less single rows on
posterior abdominal segments, there being also one or two near each spiracular opening
and one or two between clypeolabral shield and labium. Macrotubular ducts of two sizes;
a large type, about 8 µm long, with the cup wider than a large setal base; a small type,
not much longer than microtubular duct, arranged singly with shallow cup. Microtubular
ducts about 6 µm long, sparse, present in median areas as well as margins (Williams,
1985a).
Dorsum: Dorsal setae pointed and stiff; 5 µm long, in more or less single rows across
the segments. Macrotubular ducts in single rows across the segments, about 8 µm long,
with the cup wider than a dorsal setal base. Microtubular ducts in moderate numbers,
each about 6 µm long, in the form of a double tube tapering towards double orifice, each
tobe with inner end swollen into an ampulla with a filament between. Discodial pores
absent. Anal ring at apex of body, sclerotised, almost quadrate, with 4 short setae, each
12 µm long, occupying corners, 6 or 8 setae surrounding anal ring a few minute pores on
posterior margin (after Williams (1985a)).
Other stages
Description of first and second instars and adult female by Williams (1985a).
Ecology
Host plant: Fagus grandifolia, F. orientalis, F. sylvatica, Pinus sylvestris(?). Notes: This unusual
record is from crevices of Pinus sylvestris bark at Lászlótanya, Nagymilic Mtn., North
Hungary, 13 Aug. 1980 (1 female), coll. M. Kosztarab and F. Kozár (Slide No. 1278); also
2 females on 2 slides, from same host and locality, by same collectors on 7 Aug. 1975;
deposited in the collections of the Plant Protection Institute (PPI) (Slide No. 534a),
Budapest, and the Virginia Polytechnic Institute and State University, Blacksburg (Slide
No. 534b). The adults somewhat resemble those of C. aceris. When an adequate sample
is available for study of this population (if not an accidental infestation from the nearby
Fagus inhabiting typical C. fagi population) might turn out to represent a new species or
subspecies.
Distribution: Armenia, U.S.A, Austria, Belgium, Bulgaria, Canada, Croatia, former
Czechoslovakia, Denmark, France, Georgia, Germany, Hungary, Iran, Italy, Netherlands,
Poland, Romania, Russia, Slovenia, Sweden, Switzerland, Turkey, Ukraine, United
Kingdom, former Yugoslavia.
Biology: The general pattern of life cycle is as follows. Yellow eggs are laid in the spring
or early summer within the ovisac. Eggs hatch in late summer and early fall. Crawlers
are quite mobile and may move some distance before settling on the bark. The first
instar is usually the overwintering stage. During the winter, yellow first instar nymphs
change in body form from oval to rotund. Molting occurs during the spring and the short
lived apodous second instar is produced. Adult females occur throughout the spring and
summer. Males have never been collected (Miller & Miller, 1993). In adult females of
C. fagisuga having restricted access of oxygen to the spiracles are located in deep cavities
of scleotised walls which probably contain air reserves (Podsiadlo, 2006). Occasionally
becomes a pest. This scale aids in the transmission of beech bark disease, caused by
Nectria coccinea var. faginata and N. galligena can cause extensive damage to the native beech
trees.
Cryptococcidae 603
Figure 209. Cryptococcus fagisuga Lindinger, 1936, female, with first instar on top right
Williams (1985) with modifications.
Cryptococcus integricornis Danzig, 1971 (Fig. 210)

Cryptococcus integricornis Danzig, 1971b: 1415.
Holotype: Female. Russia (Primorsky Kray, Suchan, Partizansk, District, Tigrovoy),

on Tilia amurensis, 29.vi.1963, E. Danzig. By original designation. Deposited in ZMAS
(type no. 181-64). Notes: 1 female paratype on same slide as holotype. 2 female
paratypes on 1 slide with same data. 2 paratypes on 1 slide Russia (Primorsky Kray,
Suputinsky, Ussuriysky Reserve), on Tilia amurensis, 12/07/1969, E. Danzig; all in
ZMAS.
Lit.: Bareth & van Liefferinge, 1976: 210; Danzig, 1977: 44; Danzig, 1980: 235;
Kozár, 2009: 96; Kozár & Drozdják, 1998: 168; Kozár & Walter, 1985: 75; Miller &
Gimpel, 2000: 89; Tang & Hao, 1995: 427; Williams, 1985a: 352 (Miller et al., 2013).
Description
Unmounted female
Adult female pink, body hemispherical, 1 mm long. Nymphs and females grow in small
white silky wax sacks.
Mounted female
Body rotund, membranous, nodulose, largest about 0.5–1.0 mm long, often wider than
long (Fig. 210). Antenna with about 4 segments.
Venter: Labium appearing two segmented, often wider than long, shorter than
clypeolabral shield, basal segment without setae, stylet loop extending to abdominal
sternites VII, with 3 pairs of legs; first two pairs only a sclerotized spine, the posterior
a sclerotized plate with reticulation. Anal ring with pores and 6 short hair-like setae.
Spiracles often triangular, sclerotised. Setae about same size as dorsal setae, marginal
row of setae absent. Quinquelocular pores about same diameter as the cup of large
macroduct, in a submarginal row on venter where there are found in groups near each
spiracular opening. Macrotubular ducts of one size. Microtubular ducts sparse, present
on margin.
Dorsum: Dorsal setae pointed and stiff; in more or less single rows across the segments.
Macrotubular ducts in single rows across the segments. Microtubular ducts in moderate
numbers in the form of a tube towards simple orifice, few quinquelocular pores in a
submarginal band on abdomen. Anal ring at apex of body, sclerotised, elongated, with 6
short setae occupying anterior margin, a few minute pores are on posterior margin (after
Danzig 1971b).
Cryptococcidae 605
Figure 210. Cryptococcus integricornis Danzig, 1971, female. After Danzig (1971) with
modifications.
Ecology
Host plant: Tilia amurensis.
Distribution: China, Russia.
Biology: Colonies observed in late June and early July consisted of nymphs and adult
females (Danzig, 1986).
Cryptococcus ulmi Tang & Hao, 1995 (Fig. 211)

Cryptococcus ulmi Tang & Hao, 1995: 429.
Holotype: Female. China (Shanxi, Dingxiang County), on Ulmus pumila, 21.v.1990.

By original designation. Deposited IESC.
Lit Kozár, 2009: 96; Miller & Gimpel, 2000: 90; Tao, 1999: 31; Wu, 2000: 251; Xie,
1998: 101 (Miller et al., 2013).
Description
Unmounted female
Adult female body is nearly circular and white (Tang & Hao, 1995).
Mounted female
Body nearly circular, 1.0 mm long, 0.9 mm wide. Antennae 6 segmented, with sensory
setae on third, fourth and apical segments. Anal lobe indistinct, with one short seta.
Venter: Labium 3 segmented. Legs reduced to tubercles, but claws exist; each of fore
and middle legs bearing many setae around base, each hind leg with a group of pores.
A few setae on the body margin. Quinquelocular pores numerous, with a group around
mouthparts, 6–15 pores near each spiracle, arranged in a longitudinal band along body
margin and distributed on abdominal segments. Cup shape pores on the body margin.
Dorsum: Macrotubular ducts, long scattered throughout. Cup shape ducts large and
scattered, Microtubular ducts few, distributed over entire surface. Discodial pores absent.
Anal ring circular with 6 short setae, and associated with 4 spines and 2 long setae nearby,
anal tube short (after Tang & Hao (1995), Wu, (2000).
Ecology
Host plant: Ulmus sp., U. pumila.
Biology: Unknown.
Cryptococcidae 607
Figure 211. Cryptococcus ulmi Tang & Hao, 1995, female, after Tang & Hao (1995) with
modifications. First instar on top right, after Wu (2000) with modifications.
Genus:
Pseudochermes Nitsche, 1895
Type sp.: Chermes fraxini Kaltenbach, 1860: 259. By original designation.
Lit.: Afifi, 1968: 193; Boratynski & Williams, 1964: 92; Borchsenius, 1949: 43; Danzig,
1980: 231; 1986: 269; Fernald, 1903: 114; Green, 1922: 20; Hoy, 1963: 187; Kaydan
& Kozár, 2008; Kosztarab & Kozár, 1978: 67; 1988: 272; Koteja, 1974a: 295; 1974b:
78; Kozár, 2009: 111; Kozár & Konczné Benedicty, 2008b: 247; Kozár & Walter,
1985: 75; Köhler, 1998: 394; Lindinger, 1933b: 31; 1937: 179; Miller & Gimpel, 2000:
443; Morrison & Morrison, 1966: 164; Newstead, 1903: 209; Schmutterer, 1952:
418; Schmutterer, 2008: 84; Tereznikova, 1981: 13; Williams, 1985a: 385; 2007: 1357;
Zahradník, 1959: 539 (Miller et al., 2013).
Apterococcus Newstead, 1898: 97.

Type sp.: Ripersia fraxini Newstead, by monotypy and original designation. Synonymy
by Lindinger, 1937: 179, 185.
Description
Ovisac felt-like, oval, cream colored, covers female completely. Adult female ovoid, with
body segmentation well visible. Antennae 6 segmented. Frontal tubercle present. Labium
3 segmented, basal segment with a pair of short setae, total number of setae on labium 8
pairs. Eyes present. Stylet loop much longer than body. Quinquelocular pores present on
both surfaces in small numbers on venter. Legs well developed, each tibia about as long
as tarsus, mid-coxae and hindcoxae with spinulae on anterior surfaces, hindcoxae with
some translucent pores on posterior surfaces, claw without denticle. Only 1-3 discodial
pores associated with each spiracle; disc pores, predominantly quinquelocular pores,
found on both surfaces. Macrotubular ducts present, of 2 sizes, main ducts with terminal
gland flower-shaped. A smaller type of duct present with small filamentous ductule.
Cruciform pores absent. Hair-like setae on venter only. Venter of each anal lobe with 1
apical seta. Enlarged setae spinelike on venter present around margins, present in rows
on all segments on dorsum. Anal lobes undeveloped, each with two spine-like setae. Anal
ring with 6 setae, each as long as the ring diameter and with few pores present in a single
row. Cauda absent. No pores or setae anteriorly, conical setae slender, not enlarged, they
form transverse rows on dorsal segments.
First instar nymph with 6 segmented antennae, with microtubular ducts. Second instar
nymph with 6 segmented antennae (after Williams (1985a)).
Host plant
Woody plants from Oleaceae, Rosaceae, Rubiaceae, Salicaceae.
Cryptococcidae 609
Distribution
Widely distributed in the Palearctics, with 2 species.
Biology
Live under bark flakes or in crevices of bark. One yearly generation.
Key to species
1. – Dorsal abdominal segments with about 20-30 macrotubular ducts on the first
segments, Translucent pores in number of 26-53, large, situated on both surfaces
of coxae, found on Fraxinus spp. and other deciduous trees.......................... P. fraxini
– Dorsal abdominal segments with about 8–12 macrotubular ducts on the first
segments, translucent pores 14-16 in number, small, situated mostly on posterior
surface, on Coffea spp......................................................................................... P. williamsi
Pseudochermes fraxini (Kaltenbach, 1860) (Fig. 212)

Chermes fraxini Kaltenbach, 1860: 259.
Syntypes: Female. Germany. Notes: Williams (1985a) states that the type material,
which was held in Germany, is probably lost.
Common name: Ash bark scale, ash coccus; ash scale; felted ash coccus; felted ash
scale.
Lit.: Afifi, 1968: 8; Balachowsky, 1935: 265; Borchsenius, 1937: 62; 1939: 43; 1949:
48; Cook & Gullan, 2004: 444; Danzig, 1971b: 1415; 1985: 147; 1996: 574; Gertsson,
1997: 113; Hadzibejli, 1950: 262; Hoy, 1963: 54; Kiritchenko, 1931: 312; Kosztarab
& Kozár, 1988: 269; Koteja, 1974a: 249; Koteja & Zak-Ogaza, 1983: 478; Kozár,
2009: 96; Kozár & Drozdják, 1998: 167; Kozár & Kosztarab, 1982: 204; Kozár &
Walter, 1985: 75; Kuwana, 1923b: 57; Longo et al. 1999: 145; Malumphy at al., 2009:
120-147; Miller & Gimpel, 2000: 84; Schmutterer, 1980: 50; Tang & Hao, 1995: 427;
Tereznikova, 1959e: 448; Ter-Grigorian, 1962: 132; Williams, 1985a: 352; Wünn,
1926: 48; Zahradník, 1959: 539 (Miller et al., 2013).
Eriococcus fraxini; Newstead, 1891: 165-166.

Lectotype: England (Cheshire, Ince), on Fraxinus excelsior, ?.viii.1890, by R. Newstead.
Female, by subsequent designation Williams, 1985a: 386. Deposited in BMNH.
Synonymy by Judeich & Nitsche, 1895: 1249. Notes: In addition to the lectotype,
there are nine paralectotypes on the same slide in the BMNH.
Ripersia fraxini; Newstead, 1892: 147. Change of combination.
Coccus fraxini; Judeich & Nitsche, 1895: 1247. Change of combination.
Pseudochermes fraxini; Nitsche, 1895: 1247-1249. Change of combination.
Fonscolombia fraxini; Cockerell, 1899b: 264. Change of combination.

Apterococcus fraxini; Newstead, 1903: 210-213. Change of combination.
Pseudochermes fraxini; Lindinger, 1937: 194. Revived combination.
Description
Unmounted female
Adult female bright red, or orange-red, more or less globular. Sac of adult female is more
or less spherical, white and closely felted. A single sac contains 2 or more individuals.
Second stage female pale red or yellowish-red, form short ovate, partially covered with a
thin cottony secretion. Male apterous bright orange-red or yellowish-red, last 6 segments
of the abdomen greenish-yellow, eyes black. Pupae usually the same color as the male,
more or less ovate and narrowed posteriorly. First instars elongate-ovate, pale crimson-
yellow, mottled with bright crimson, the latter color being also the color of the eyes. Eggs
pink when first laid, but become mottled with reddish-crimson (Newstead, 1903).
Mounted female
Young adult female oval, 0.9 mm long, 0.9 mm wide, but mature adult female sometimes
almost circular, membranous, posterior end rounded. Antenna 6 segmented, 100–110
µm long. Frontal tubercle present just anterior to basal antennal segment. Eyes situated
on venter near margin. Anal lobes not developed, but position of each anal lobe with an
apical seta 60 µm long.
Venter: Labium 3 segmented, 50–60 µm long; much smaller than clypeolabral shield;
basal segment with a single pair of minute setae. Legs small but developed. Hind
trochanter + femur 70–75 µm long, hind tibia 35–40 µm long, hind tarsus 30–35 µm
long, claw 16 µm long, curved with a denticle near apex. Hind coxa noticeably larger
than anterior coxae, with a few large translucent pores. Slender setae only slightly longer
than dorsal setae, except on head where they are much longer. Quinquelocular pores not
numerous on head and thorax but more numerous on abdomen. Macrotubular ducts of
two sizes; a larger type about 20 µm long, tapering to narrow orifice, the cup wider than
a setal base of largest setae, around margins only except on thorax near spiracles where
they also occupy the submarginal areas; narrower type with sclerotised cup and filiform
tube, tapering to a minute orifice, sparse, on abdominal segments only. Microtubular
ducts 4 µm long, a few on margin. Cruciform pores absent.
Dorsum: Dorsal enlarged setae narrow, conical about 20 µm long, on seventh and anal
lobe segments and usually shorter than anal ring setae; anteriorly setae shorter and more
slender, but stiff, 8 µm long. Quinquelocular pores present in more or less single rows at
posterior edges of segments. Macrotubular ducts fairly numerous, in even distribution,
about 20 µm long, tapering to narrow orifice, the cup wider than a setal base of largest
setae. Microtubular ducts 4 µm long, not numerous, tending to be in rows across the
segments, filiform with small sclerotised ampulla. Anal ring in form of crescentic
Cryptococcidae 611
Figure 212. Pseudochermes fraxini (Kaltenbach, 1860), female, first instar on top left.
After Williams (1985) with modifications.
sclerotised plate, 30 µm wide, with a few pores and 6 setae, each about 30 µm long, blunt
and stiff (afer Williams (1985a).
Other stages
Detailed description of different stages, including adults given by Newstead (1903),
Williams (1985a), Afifi (1968).
Ecology
Host plant: Fraxinus sp., F. angustifolia, F. caroliniana, F. excelsior, F. mandschurica, F. ornus,
Ligustrum sp., Populus tremula, Syringa sp., Sorbus aucuparia.
Distribution: Austria, Belgium, Bulgaria, China, Croatia, former Czechoslovakia,
Denmark, Finland, France, Germany, Greece, Hungary, Iran, Italy, Lithuania,
Luxembourg, Netherlands, Norway, Poland, Portugal, Romania, Russia, Slovenia, Spain,
Sweden, Switzerland, Turkey, Ukraine, United Kingdom, former Yugoslavia .
Biology: In British Isles females had completed egg-laying by the 21st of May, and on
the 10th of June the first instars were swarming. Pupation takes place in September and
the males appear early in October, and continue to emerge throughout the month and
during the early part of November. Males are very active (Newstead, 1903). According to
Danzig (1986), in Russia females are the overwintering stage and eggs are laid from May
to June. P. fraxini feeds in bark crevices of trees, often a pest, especially in towns. The
access of air to spiracles is limited since it remains pressed with the ventral side into a
tight space when feeding. Thereofre, spiracles are situated in deep cavities of sclerotized
walls. In addition, the clypeo-labral shield is very large which is the adaptation to feeding
on woody parts of plants, and to living in unfavorable oxic conditions (Podsiadlo, 2006).
Cryptococcidae 613
Pseudochermes williamsi Kozár & Konczné Benedicty, 2008 (Fig. 213)

Pseudochermes williamsi Kozár & Konczné Benedicty, 2008b: 250.
Holotype: Female. Spain (Canary Island, Tenerife, Orotaro Botanical Garden),

on Coffea arabica, 13.iv.1976, by D.J. Williams. By original designation. Deposited in
BMNH and one paratype deposited in HNHM.
Lit.: Kozár & Konczné Benedicty, 2008b: 250; Ouvrard & Kozár, 2009: 105 (Miller
et al., 2013).
Description
Unmounted female
Unknown.
Mounted female
Body of slide-mounted specimens, elongate-oval, 0.72–0.74 mm long, 0.44–0.56 mm
wide. Antennae 6 segmented; length segments as follows: I: 17–20, II: 14–15, III: 10–12,
IV: 7–10, V: 7–9, VI: 14–18 µm long, all segments with few setae, second segment with
1 sensory pore, segment VI with apical setae, each 34–41 µm long, and with 3 sensory
falcate setae, each 17–20 µm long, segments IV and V each with single falcate seta, each
10–14 µm long. Apical segment with two coeloconic sensilla, each 3–4 µm long. Frontal
tubercles present. Eyes present on ventral margin.
Venter: Labium 3 segmented, 37–41 µm long, basal segment not well developed,
with a pair of setae. Stylet loop not seen. Legs well developed; lengths of segments
of prothoracic legs; coxa: 15–18, trochanter: 12–15, femur: 37–40, tibia: 23–26, tarsus:
23–26, claw: 11–15 µm long; lengths of segments of mesothoracic legs; coxa: 16–19,
trochanter: 11–16, femur: 36–40, tibia: 23–27, tarsus: 24–28, claw: 11–15 µm long;
lengths of segments of metathoracic legs; coxa: 18–21, trochanter: 13–20, femur: 39–43,
tibia: 27–32, tarsus: 28–30 µm long, tarsal digitules knobbed, 19–25 µm long, claw 11–15,
claw digitules: 10–16 µm long, slightly knobbed. Meso-and metacoxae with spinulae on
anterior (ventral) surfaces, hindcoxae with large translucent pores on posterior (dorsal)
surface; claw with denticle. All legs with few flagellate setae. Mesothoracic spiracles 8–9
µm in diameter, with some quinquelocular pores at spiracular opening. Quinquelocular
pore 3 µm in diameter, scattered in small numbers on much of surface. Slender short
setae present, scattered on abdomen. Some longer flagellate setae, present on head and
thorax. Macrotubular ducts of 2 sizes; about 3–5 µm wide, 10–16 µm long, present on
all segments, each duct with sclerotized rim surrounding orifice, particularly discernible
on larger ducts; inner ductule of larger type of duct longer than duct, ending in a flower-
shaped gland, on venter ducts are shorter and narrower. Smaller macrotubular ducts
with 6-locular inner orifice, 2 µm wide, 11 µm long, inner ductule shorter than duct. Few
microducts present around margin. Oviduct or internal genital organ barely discernible.
Dorsum: Setae on last abdominal segments robust, spine-like, of 2 sizes, 15–17 µm long,
present mostly in single rows across segments, 1 or 2 present on margin of each segment.
Other setae 7–8 µm long scattered all over the body. Macrotubular ducts 10–16 µm long,
about 4–8 on each segments. Microducts present, each about 7 µm long and less than 1
µm wide, with normal orifice, scattered among dorsal setae. Quinquelocular pores same
as those on venter, scattered on margin. Anal ring situated on dorsum, well developed,
19–22 µm wide, 16–17 µm long, with single row of pores and 6 strong setae each 18–20
µm long. Anal lobes not well developed, not sclerotized, with 2 spine-like setae, as long as
marginal ones. Apical setae each 72–90 µm long. Cauda absent (after Kozár & Konczné
Benedicty (2008b)).
Other stages
Mounted first instar nymph (Fig. 213, top left)

with strong sensory setae as in adult female. Dorsum with dome-shaped spines of one
sizes. Usually with six spinose setae on each segment. Tubular ducts absent. Venter with
transverse rows of six small hair-like setae on each abdominal segment; median setae
longer than others. Cruciform pores absent. With 1 pair of quinquelocular pores on each
thoracic segment, 1 near each spiracle, plus 1 pair on frons. Anal ring with 6 spine-like
setae. First instar nymph with a pair of rows of equal long, strong spines in mid-dorsum.
Ecology
Host plant: Coffea arabica.
Distribution: Spain (Canary Islands).
Biology: Unknown.
Comments
Danzig (1980) mentioned populations of P. fraxini from the Far East of Russia, which
differs somewhat from the European populations by having smaller numbers of tubular
ducts and quinquelocular pores, and that they are somewhat similar to P. williamsi. It
deserves further study. Spines of the first instar nymph of P. williamsi are conical, sharply
pointed, not truncate as in P. fraxini, may serve as additional help in the identification of
the two populations.
Cryptococcidae 615
Figure 213. Pseudochermes williamsi Kozár & Konczné Benedicty, 2008, female, first
instar on top left. After Kozár & Konczné Benedicty (2008).
Family:
ERIOCOCCIDAE
Signoret, 1875
Type genus: Eriococcus Targioni Tozzetti, 1868.
Common name: Felt scales, box scales.
Lit.: Borchsenius, 1948: 502; 1949: 322; Cook & Gullan, 2001: 59; 2004: 541; Gullan
& Cook, 2007: 413; Danzig, 1962: 839; 1975: 62, 1980: 203; Fernald, 1903: 70; Ferris,
1955: 94, 1957: 81; Foldi & Kozár, 2007: 51; Goux, 1948: 5; Gwiazdowski et al., 2006:
9; Hardy et al., 2008: 502; Hardy & Gullan, 2007b: 106; Henderson, 2006: 37, 2007b:
25, 2007a: 10; Hodgson & Henderson, 1996: 143; Hodgson & Miller, 2010: 191; Hoy,
1962:10, 1963: 62; Kaydan & Kozár, 2008: 16; Kondo et al., 2006: 19; Kosztarab &
Kozár, 1978: 64, 1988: 274; Koteja, 1974a: 297, 1974b: 76; Kozár, 2009: 96; Kozár
& Konczné Benedicty, 2008b: 250, Kozár et al., 2007: 88, 2009: 1; Kozár & Walter,
1985: 74; Köhler, 1998: 371; Leonardi, 1920: 425; Matesova, 1967: 1193; Miller, 1970:
157; Miller & Gonzales, 1975: 136; Miller & Gimpel, 1996: 597; 1999: 213; Miller &
Miller, 1992: 2, 1993: 9; Miller & Williams, 1976: 118; Morrison & Morrison, 1966:
69; Newstead, 1903: 105; Ouvrard & Kozár, 2009: 101; Schmutterer, 1952: 68, 2008:
84; Tang & Hao, 1995: 415; Tereznikova, 1981: 45; Williams, 1969b: 325; 1985a: 356;
2007: 48; Wu, 2000: 251 (Miller et al., 2013).
Eriococcini Cockerell, 1899a: 389.

Eriococcinae Cockerell; Maxwell-Lefroy, 1909: 758.
Eriococcidae Cockerell; Brues & Melander, 1932: 134.
Eriococci Cockerell; Silvestri, 1939: 638.
Comment
According to Miller & Williams (1976), Eriococcus crispus (Boyer de Fonscolombe, 1834) is
not an eriococcid, because of this here we did not treat it. It is possible that this species
is a member of the family Dactylopiidae genus Dactylopius.
The status of this diverse family, its definition, and borderlines, constitute much
controversy among coccidologists all time, see also discussion under the family of
Acanthococcidae. Although presently only three genera with six species are included
in this family in the Palaearctic region, in the future after a detailed study of about 170
species left by Kozár (2009) in his so called “Eriococcus” genus, will probably result in
some new genera and several species to be be transferred to the Eriococcidae family sensu
stricto. According to molecular data of Cook & Gullan (2004) and Gullan & Cook (2007),
the type genus Eriococcus and type species E. buxi is a representative of the BSE clade and
Eriococcidae 617
for this reason the present distribution records do not indicate the real distribution and
origin of this family, only the collection places of some members.
Description
Ovisac felt-like, encloses female entirely.
Mounted female
Adult female elongate oval, tapering posteriorly. Labium 1 or 3 segmented, basal segment,
if present, with a pair of setae. Antennae 3-7 segmented. Legs well developed; tibia with
four setae, claw with a denticle. Discodial pores present on venter or dorsum, or on
both surfaces. Microtubular ducts, if present, could be found on dorsum. Macrotubular
ducts, if present, on both, dorsum and venter; enlarged macrotubular ducts with a unique
shape, present on dorsal margin and on midline of dorsum and head near eyes. Enlarged
conical setae with unusually sharp-pointed apex on dorsum. Anal ring with pores and
6-8 setae. Anal lobes long, if developed, almost cylindrical each with a long apical seta.
Host plant
The members of Eriococcidae family collected on the leaf of Buxus spp. and other
woody plants.
Distribution
World distribution of this family is not clear, found in the Palaearctic, Oriental and
Neotropic regions. In the Palearctic Region known by 6 species in three genera.
Biology
Live mostly on the leaves and stems of different deciduous hostplants. One yearly
generation. Some of them (Aculeococcus) form galls on the leaves.
Key to genera
1. – Mounted adult female pear shaped, abdomen sclerotized, legs short, deformed
..............................................................................................................................Aculeococcus
– Mounted adult female oval, abdomen not sclerotized, legs short, not deformed....2
2. – Antennae five or less segmented................................................................Asiacornococcus
– Antennae six segmented...................................................................................... Eriococcus
Genus:
Aculeococcus Lepage, 1941
Type sp.: Aculeococcus morrisoni Lepage, 1941, by monotypy.
Lit.: Ferris, 1957a: 63; Gullan et al., 2005: 166; Hao, et al., 1997: 71; Hodgson & Miller,
2010: 100; Hodgson et al., 2004: 52; 2011: 54; Hoy, 1963: 31; Koteja, 1974a: 301;
Kozár, 2009: 111; Kozár & Konczné Benedicty, 2008a: 141; Miller & Gimpel, 2000:
20; Morrison & Morrison, 1966: 3; Tang & Hao, 1995: 436; Wang, 1974: 329 (Miller
et al., 2013).
Description
Body approximately pear shaped, venter becoming highly swollen anteriorly. Abdomen
heavily sclerotized, mouthparts with large apodemes arising from the tentorial box, heavily
sclerotized. Slide-mounted adult female pear shaped; posterior abdominal segments
sclerotized; anal lobes not developed; legs small partly deformed; antennae reduced to
a 3 segmented stub, hind coxae greatly enlarged, dorsal abdomen with stout, acute setae
as long as segment that bears them; apparently without macro- and microtubular ducts;
quinquelocular pores present. First instar with: nipple-shaped enlarged setae; antennae 3
segmented; distinct anal lobes (Ferris, 1957a; Hodgson & Miller, 2010; Miller et al., 2013)).
Host plant
Unknown, probably Lauraceae.
Distribution
Only two species known, one from Brazil and one from China.
Biology
Members of this genus form galls, induces elongate, pointed, conical galls on upper
surface of leaves of hostplant.
Aculeococcus yongpingensis Tang & Hao, 1995 (Fig. 214)

Aculeococcus yongpingensis Tang & Hao, 1995: 437.
Holotype: Female. China (Yunnan, Shidian and Yongping), Deposited in EISC.

Notes: Tang & Hao (1995) first described the species with no author specified,
therefore they must be considered the authors of the species. Hao et al (1997) cite
this species as "sp. nov." and authored by Tang, but this is incorrect.
Eriococcidae 619
Figure 214. Aculeococcus yongpingensis Tang & Hao, 1995, female. After Tang & Hao
Lit.: Hao, et al., 1997: 71; Kozár, 2009: 94; Miller & Gimpel, 2000: 20; Tang & Hao,
1995: 437; Tao, 1999: 31 (Miller et al., 2013).
Description
Unmounted female
Body approximately pear shaped, in leaf gall which open upward, about 0.8–1.1 mm
in diameter and 1.2 mm in depth. Body fills the gall and with sclerotized dorsum of
abdomen covering the gall mouth, body is bulb shaped.
Mounted female
Antenna 3 segmented.
Venter: Labium reduced, directed upward, stylet loop short. Legs small, but well
developed: tibia and tarsus equal long, reduced, tarsal and claw digitules slightly knobbed,
longer than claw. Anterior legs situated far from other legs. Setae in median areas few,
short. Multilocular pores with 5 loculi, numerous, on last abdominal segments and in
goup near spiracles. Macrotubular ducts, cruciform pores absent.
Dorsum: Strong pointed setae in rows and bands on abdominal segments, marginal setae
absent. Macrotubular and microtubular ducts absent. Anal lobes not developed. Anal
ring sclerotized, with 6 short spine-like setae. Cauda not shown. Dorsum of abdomen
sclerotized (after Tang & Hao (1995), with modifications).
Ecology
Host plant: Unknown, probably Lauraceae.
Distribution: China (Yunnan).
Biology: This species induces elongate, pointed, conical galls on upper surface of leaves
of host plant. Adult female completely fills the gall.
Eriococcidae 621
Genus:
Asiacornococcus Tang & Hao, 1995
Asiacornococcus Tang & Hao, 1995: 587.
Type sp.: Eriococcus exiguus Maskell, by monotypy and original designation.

Lit.: Kozár, 2009: 96; Miller & Gimpel, 2000: 68; Tang & Hao, 1995: 587; Xie, 1998:
Description
Adult female body oval in form, red in color, enclosed in white waxy sac like rice.
Antennae 3–5 segmented. Eyes visible on the body margin. Mouthparts developed. Basal
segment of labium with one pair of setae. Legs well developed; tibia with four setae, tarsus
and claw with digitules, slightly knobbed. Thoracic spiracles slightly sclerotised around.
Anal lobes produced, with inner margins smooth and apical setae longer than anal ring
setae. Anal ring with pores in rows and 6-8 setae. Dorsal setae on body acorn-shaped,
distributing 4 or 5 longitudinal rows, marginal, submarginal and medial ones, but absent
on the last abdominal segment. Dorsum with macrotubular ducts and microtubular ducts
in quantity, quinquelocular pores on both surfaces.
Host plant
Woody plants from different plant orders, as Rosaceae, Moreaceae, Theaceae.
Distribution
Eastern-palearctics, and Oriental with 3 species.
Biology
On the leaves and plant surface.
Key to species
1. – Antennae 5 segmented......................................................................................A. japonicus

– Antennae 3 or 4 segmented...............................................................................................2
2. – Anal lobe with three setose enlarged setae....................................................... A. exiguss
– Anal lobe with one strong spine-like, and one setose enlarged setae..............A. kaki
Asiacornococcus exiguus (Maskell, 1897) (Fig. 215)

Eriococcus exiguus Maskell, 1897: 243.
Syntype: Female. China (Hong Kong), on unidentified host. Deposited in Auckland:

USNM Notes: According to Miller et al. (1973) there is also type material in UCDC.
Lit.: Ali, 1970: 76; Cheo, 1935: 98; Deitz & Tocker, 1980: 46; Fernald, 1903: 74; Ferris,
1936: 11; Hoy, 1963: 89; Hua, 2000: 137; Kozár, 2009: 96; Kozár & Walter, 1985: 74;
Kuwana, 1927: 70; Lindinger, 1933a: 108; Martin & Lau, 2011: 45; Maskell, 1898:
243; Miller et al, 1973: 7; Stoetzel & Miller, 1979: 14; Tang, 2001: 3; Tang & Hao,
1995: 439; Tao, 1999: 31; Wang, 1974: 329; 2001: 207; Wu, 1935: 176; Yang, 1982: 104
Nidularia exiguus; Lindinger, 1933a: 108. Change of combination.

Acanthococcus exiguus; Kozár & Walter, 1985: 74. Change of combination.
Asiacornococcus exiguus; Tang & Hao, 1995: 588. Change of combination.
Acanthococcus exiguus; Köhler, 1998: 376. Revived combination.
Asiacornococcus exiguus; Miller & Gimpel, 2000: 68. Revived combination.
Description
Unmounted female
Sac of adult female yellow, elliptical, rather loosely felted. Adult female is brownish-
yellow and elliptical in form.
Mounted female
Body hemispherical oval, extremely small, 0.75 mm long. Antennae 3 segmented. Eyes
situated on venter near margin. Anal lobes developed, long, with one enlarged setae and
two hair-like setae.
Venter: Labium 3 segmented, short. Legs well developed, tibia with four setae. Setae
short, flagellate. Multilocular pores very small on mid abdominal segments.
Dorsum: Dorsal setae on body acorn-shaped, distributed in 4 longitudinal middorsal
rows, in marginal row two setae present on segments. Macrotubular ducts very small,
present throughout. Discoidal pores not mentioned. Anal ring almost circular, with six
setae, anal lobe setae nearly twice as long as the anal ring setae (after Ferris (1936), with
modifications).
Ecology
Host plant: Rosa? sp.
Eriococcidae 623
Figure 215. Asiacornococcus exiguus (Maskell, 1897), female. After Ferris (1937) with
modifications.
Comment
According to Maskell (1898) antennae “with six subequal joints”. Here we follow Ferris
(1936) drawing.
Asiacornococcus japonicus (Kuwana, 1902) (Fig. 216)

Eriococcus japonicus Kuwana, 1902: 50.
Syntype: Female. Japan (Kyushu, Chikujo-gun), on Symplocos myrtacea, by I. Kuwana.

Deposited in USDC and ITLJ.
Lit.: Fernald, 1903: 75; Hoy, 1963: 97; Kawai, 1972: 4; 1977: 152; 1980: 131; Kozár,
2009: 96; Kuwana, 1917: 138; Lindinger, 1933a: 116; Lindinger, 1943: 264; Stoetzel &
Miller, 1979: 19; Takahashi, 1957: 7 (Miller et al., 2013).
Acanthococcus japonicus; Kozár & Walter, 1985: 74. Change of combination.

Asiacornococcus japonicus; Tang & Hao, 1995: 439. Change of combination.
Acanthococcus japonicus; Köhler, 1998: 378. Revived combination.
Asiacornococcus japonicus; Miller & Gimpel, 2000: 69. Revived combination.
Description
Unmounted female
Female sac is convex, elliptical, and pale straw colored, 1.7 mm long, 1 mm wide. Female
is oval, 0.65 mm long and 0.4 mm wide, and eggs are brown (Kuwana, 1902).
Mounted female
Slide-mounted adult female with 5 segmented antennae (Tang & Hao, 1995). Anterior
legs shorter than others, tarsus longer than tibia, claw large, curved, digitules on tarsus
long and fine, digitules on claw stout and short, posterior coxae very large, almost as long
as femur. Margin and dorsum with rows of strong spines. Anal lobe with a few spines,
anal ring with six setae.
Ecology
Host plant: Symplocos myrtacea.
Biology: Unknown.
Eriococcidae 625
Comment
Kuwana (1902) mentioned the species as having 6 segmented antennae, but on his
drawing he drew only 5 segments, so here we accept Tang & Hao’s (1995) drawing. This
species has been confused with Nidularia japonica Kuwana, 1918 by Hoy (1963) and was
also incorrectly considered a senior secondary homonym by Lindinger (1943). Nidularia
japonica is currently placed in the Kermesidae.
Figure 216. Asiacornococcus japonicus (Kuwana, 1902), female. After Kawai (1980) with
modifications.
Asiacornococcus kaki (Kuwana, 1931) (Fig. 217)

Eriococcus kaki Kuwana in Kuwana & Muramatsu, 1931: 659.
Syntype: Female. China (Taken in quarantine in Japan, Kyushu, Nagasaki), on

Diospyros kaki, by K. Tanaka. Deposited in ITLJ.
Lit.: Chen, et al., 1987: 203; Hoy, 1963: 97; Hsu, 1935: 580; Hua, 2000: 137; Hu, et al.,
1992: 180; Wang, 1992: 180; Kozár, 2009: 96; Kozár & Walter, 1985: 74; Kuwana &
Muramatsu, 1931: 659; Tang, 1977: 41; 1984: 126; 2001: 3; Tang & Hao, 1995: 416;
Tao, 1999: 31; Wang, 1974: 329; 1980: 115; 2001: 207; 1982: 41; 1982a: 143; 2001: 207;
Wu, 1935: 176; Xie, 1998: 99; Yang, 1982: 104; Zhang and Huang, 1980: 149 (Miller
et al., 2013).
Acanthococcus kaki; Kozár & Walter, 1985: 74. Change of combination.

Asiacornococcus kaki; Tang & Hao, 1995: 439. Change of combination.
Acanthococcus kaki; Köhler, 1998: 379. Revived combination.
Asiacornococcus kaki; Miller & Gimpel, 2000: 69. Revived combination.
Description
Unmounted female
Body of female oval in outline.
Mounted female
Body of female oval in outline. Antennae 3 segmented. Eyes situated on venter near
margin. Anal lobes slightly developed, with two enlarged setae one larger, other spinelike,
smaller than dorsal setae.
Venter: Labium 3 segmented, short, basal segment with a pair of setae, median setae on
apical segment strong, short. Legs well developed; tibia with four setae, tarsal and claw
digitules slightly knobbed, longer than claw. Setae hair-like on the middle of abdominal
segment, spinelike setae in a band on margin. Quinquelocular pores strongly sclerotised,
in rows on abdominal segments, on thorax and head. Cruciform pores numerous on
submarginal area. Macrotubular ducts of two sizes, smaller ones scattered on abdominal
segment; larger ones with microtubular ducts in rows on margin.
Dorsum: Dorsal enlarged setae numerous, stout and pointed; marginal enlarged setae
of two kinds, short, strong, blunted, as long as wide, this kind is present mainly on
dorsum, other kinds narrower and longer, two times longer than wide. Distributed in
longitudinal rows and bands. Quinquelocular pores present, scattered. Macrotubular
ducts present throughout, in rows on abdominal segments. Microtubular ducts long,
scattered throughout. Anal ring almost circular, with small number of pores and with 8
setae (after Tang & Hao (1995) with modifications).
Eriococcidae 627
Figure 217. Asiacornococcus kaki (Kuwana 1931), female. After Tang & Hao (1995) with
modifications.
Ecology
Host plant: Camellia oleosa, Diospyros kaki, Ficus carica, Lagerstroemia speciosa, Prunus
armeniaca, Ternstroemia sp.
Distribution: China, Japan (?).
Biology: Specimens were collected in Nagasaki customs on a plant from China (Kuwana
& Muramatsu, 1931). Its establishment in Japan has not been verified.
Comments
According to Tang & Hao (1995) adult female in the key have 4 segmented antennae.
However on their drawing there are only 3, but in the drawing by Tang (1977) there are
four segments drawn. On the drawing of Wang (1982a), the antennae is also shown as
3 segmented. The antennal segment number was 3 in slides which we examined. It is
thought that there might be some cryptic species covered under this name or there might
be big individual variation among the populations.
Eriococcidae 629
Genus:
Eriococcus Targioni Tozzetti, 1868
Type sp.: Coccus buxi Boyer de Fonscolombe, 1834: 218. Type data: France: Aix-en-
Provence, on Buxus sempervirens, ?.v.1834. Syntypes, female. Subsequently designated
by Opinion, 1203, 1982: 95. Notes: The International Commission on Zoological
Nomenclature ruled that the type species of Eriococcus be C. buxi in Opinion
1203. A detailed history of the genus and its various possible type species was
presented by Miller & Williams (1976). Notes: Williams (1985a) noted that the type
material has probably been lost (see remarks). Matile-Ferrero & Danzig visited the
Naturhistorisches Museum, Vienna in June of 1997 and were unable to locate any
type material of this species.
Common name: Box scale.
Lit.: Fernald, 1903: 70; Borchsenius, 1949: 322; Danzig, 1975: 42; Ferris, 1955: 94;
Hoy, 1963: 62; Kosztarab & Kozár, 1988: 287; Koteja, 1974a: 248; 1974b: 77; Kozár &
Walter, 1985: 74; Miller, 1970: 4190; Miller & Gonzales, 1975: 136; Miller & Williams,
1976: 118; Morrison & Morrison, 1966: 69; Schmutterer, 2008: 84; Williams, 1985a,
356; Williams, 1969b: 325 (Miller et al., 2013).
Thekes Maskell, 1892: 28.

Type sp.: Thekes multispinus Maskell, 1879: 217.
Syntype: Female. Australia (South Australia, Adelaide), Deposited in NZAC, BMNH,
USNM. Notes: Probably this genus should be re-established, including the T. eucalypti
Maskell, 1892, as it was mentioned by Maskell (1892).
Comment
All other generic synonyms and specific names known from the Palaearctic Region,
mentioned by Miller & Gimpel (2000) are treated here in Acanthococcidae family.
Description
Ovisac felt like, encloses female entirely. Adult female elongate oval, tapering posteriorly,
with well developed and heavily sclerotised anal lobes. Labium 3 segmented, basal
segment a pair of setae. Antennae 6 or 7 segmented. Hind coxa, femur and tibia often
with translucent pores, claw with denticle. Discodial pores on venter only, no discodial
pores on dorsum. Microtubular ducts scattered on dorsum. Among macrotubular ducts
on both dorsum and venter; some usually shaped macrotubular ducts on dorsal margin
and on midline of dorsum and head near eyes. Enlarged conical setae with unusually
sharply pointed apex on dorsum. Anal ring with pores and 8 setae. Anal lobes long,
almost cylindrical each with a long apical seta.
Host plant
Woody plants from Buxaceae.
Distribution
Western part of Palearctic, with 2 species. The real number of species and distribution of
the genus, not clear because of the unsolved situation in “Eriococcus” sensu lato (see more
detailes in the comments of the Eriococcidae family).
Biology
Live under bark flakes or in crevices of bar and on the leaves. One yearly generation.
Key to species
1. – With large numbers of submarginal enlarged ducts, many surrounded by groups

of setae, and other ducts on the midline....................................................... E. williamsi
– With small numbers of submarginal enlarged ducts, none surrounded by groups
of setae, and other ducts on the midline..............................................................E. buxi
Eriococcus buxi (Boyer de Fonscolombe, 1834) (Figs. 218, 219)

Coccus buxi Boyer de Fonscolombe, 1834: 218.
Syntype: Female. France (Aix-en-Provence), on Buxus sempervirens, ?.v.1834. Notes:

Williams (1985a) notes that the type material has probably been lost (see remarks).
Matile-Ferrero & Danzig visited the Naturhistorisches Museum, Vienna in June of
1997 and were unable to locate any type material of this species.
Common name: Box scale.
Lit.: Afifi, 1968: 175; Archangelskaya, 1937: 27; Balachowsky, 1937a: 340; Ben-Dov,
1977: 8; Bielenin & Weglarska, 1990: 377; Borchsenius, 1934: 13; 1936: 111; 1937: 60;
1949: 18; Cockerell, 1896: 323; Cook & Gullan, 2001: 60; 2004: 444; Danzig, 1964:
632; 1971a: 821; 1975: 42; Fernald, 1903: 72; Fetykó, et al., 2010: 296; Foldi, 2001:
305; Froggatt, 1921: 75; Gómez-Menor Ortega, 1937: 346; 1946: 103; Goux, 1931:
72; 1931a: 331; 1943: 128; Hadzibejli, 1983: 269; Hodgson, 2002: 135; Hodgson &
Henderson, 1996: 192; Hoy, 1963: 76; Kaydan et al., 2007: 90; Kiritchenko, 1928: 112;
Kondo et al., 2006: 23; Kosztarab & Kozár, 1988: 287; Koteja, 1974a: 248; Kozár,
2009: 97; Kozár & Walter, 1985: 74; Kozár et al., 2009: 1; Köhler, 1998: 387; Lindinger,
1958: 368; Martin-Mateo, 1985: 91; Maskell, 1895: 21; Miller, 1991: 333; Miller &
Gimpel, 2000: 150; Milonas et al., 2008: 143; Ouvrard & Kozár, 2009: 101; Pellizzari
& Kozár, 2011: 66; Tang & Hao, 1995: 417; Signoret, 1875: 30; Tereznikova, 1981: 48;
1982: 36; Ter-Grigorian, 1962: 130; 1966: 372; 1969: 6; Tranfaglia & Esposito 1985:
Eriococcidae 631
115 Tsalev, 1968: 207; Williams, 1969a: 93; 1985a: 358; Williams and Ben-Dov, 2009:
12; Zahradník, 1977: 121 (Miller et al., 2013).
Eriococcus buxi; Targioni Tozzetti, 1869: 726. Change of combination.

Nidularia buxi; Lindinger, 1935: 135. Change of combination.
Eriococcus buxi; Borchsenius, 1948: 501. Revived combination.
Description
Unmounted female
Ovisac grayish, encloses female entirely. Adult female ovoid, dark red.
Mounted female
Adult female broadly oval, 1.9 mm long, 1.35 mm, broad, not nodulose. Antennae 6
segmented, 160–175 µm long. Frontal lobes well developed, about as long as width of
basal antennal segment. Anal lobes about twice as long as wide, conical, each lobe with
apical setae 140 µm long. Dorsally with an outer enlarged seta situated submarginally
rather than on edge, and 2 inner enlarged setae. Ventrally with a single slender seta and a
suranal seta shorter than anal ring setae.
Venter: Labium 80 µm long, shorter than clypeolabral shield, basal segment with
a single pair of setae. Stylet loop reaching behind median coxae. Legs robust, hind
trochanter+femur 170–190 µm long, hind tibia 80–90 µm long, hind tarsus 80–90 µm
long, the tibia+tarsus either same length as trochanter+femur or slightly shorter, claw 25
µm long, straight with a minute denticle near apex; hind coxa with a few small translucent
pores. Setae normal, slender, on median areas; in submarginal areas the setae are shorter,
stiff and blunt. Enlarged setae, around margins only. Discodial pores quinquelocular,
numerous across abdominal segments almost to margins, occasional pores present in
median areas of thorax and head and around spiracular openings. Macrotubular ducts of
two sizes, a larger type, about 20 µm long, the cup about two-thirds width of diameter
of setal base of enlareged seta, the inner end of filament small and barely perceptible;
numerous in a wide zone around margins and submargins, on thorax reaching to spiracles,
and on head occupying area between clypeolabral shield and antennal bases; a narrower
type on abdomen only, in transverse rows or bands across median areas of segments.
Microtubular ducts, 12 µm long, around margins only. Cruciform pores absent.
Dorsum: Dorsal enlarged setae scattered densely, lanceolate, 32–52 µm long, pointed,
the sides concave but curved in profile and thorn-like, covering surface of dorsum, all
approximately same size. Macrotubular ducts of one size, evenly distributed, about 20
µm long, the cup about two-thirds width of diameter of setal base of enlareged seta,
the inner end of filament small and barely perceptible. Enlarged ducts present, about 35
µm long, the sides almost parallel, with inner end flat and rim of orifice sclerotised, but
in profile thicker towards one side; present either singly or sometimes in pairs on head
margin where 2-4 present at most. Microtubular ducts in a regular arrangement, about 12
µm long, very slender, the sides parallel, with single sclerotized area, orifice widely bifid.
Anal ring with 8 setae, each about 110 µm long. Cauda only lightly sclerotised, rounded
(Williams, 1985a).
Other stages
First instar nymph (Fig. 218, top right)

Venter: Labium 3 segmented, basal segment with 1 pair of short setae, apical segment
with 5 pairs of equal length hair-like setae, and 1 pair short apical setae. Legs normal:
tarsal and claw digitules slightly knobbed, longer than claw. Claw with a denticle. Legs
with a few hair-like setae; tibia with 4 setae. Multilocular pores quinquelocular, in one
longitudinal row on venter. Microtubular ducts absent on venter. Cruciform pores absent.
A few hair-like setae present on submedian venter. Suranal setae hair-like.
Dorsum: Enlarged setae pointed, conical; dome-shaped, in a row on margin, one on
margin of each segment, and a group on head margin, shorter setae in four longitudinal
rows on dorsum. Microtubular ducts scattered, in four longitudinal rows. Anal ring
situated on margin of venter; oval, sclerotized, with one row of pores and with 6 setae.
Anal lobes well developed, membranouos, with 3 blunted conical spines, ventrally with
two flagellate setae; and a long apical seta. Cauda wide, plate-like.
Second instar female nymph (Fig. 219 a)

Frontal lobe and frontal tubercle present. Eyes situated on venter near margin.
Venter: Labium 3 segmented, basal segment with 1 pair of short setae, apical segment
with 5 pairs of equal length hair-like setae, and 1 pair short apical setae. Stylet loop not
seen. Legs normal: tarsal and claw digitules slightly knobbed, longer than claw. Claw with
a denticle. Legs with a few hair-like setae; tibia with 4 setae. Median and posterior coxae
with spinulae, posterior coxae and femur with large oval pores, too. Multilocular pores
with 3-5 loculi, mostly quinqueloculars; arranged in six longitudinal rows. Microtubular
ducts absent on venter. Cruciform pores absent. A few hair-like setae present on
submedian venter in six longitudinal rows. Suranal setae hair-like.
Dorsum: Enlarged setae sharply pointed, conical; in a row on margin, two on margin of
each segments, shorter setae in 6-8 longitudinal rows on dorsum. Microtubular ducts very
long with oval opening, scattered among spines. Anal ring situated on margin of venter;
oval, sclerotized, with one row of pores and with 6 setae. Anal lobes well developed,
membranouos, with 3 pointed, conical spines, ventrally with two strong flagellate setae;
and a long, strong apical seta. Cauda wide, plate-like.
Eriococcidae 633
Figure 218. Eriococcus buxi (Boyer de Fonscolombe, 1834), female, after Williams (1985).
First instar on top right, original.
Second instar male nymph (Fig. 219 b)

Frontal lobe and frontal tubercle present. Eyes situated on venter near margin.
Venter: Labium 3 segmented, basal segment with one pairs of short setae, apical segment
with 5 pairs of equal length hair-like setae, and 1 pair short apical setae. Stylet loop not
seen. Legs normal: tarsal and claw digitules slightly knobbed, longer than claw. Claw with
a denticle. Legs with a few hair-like setae; tibia with 4 setae. Median and posterior coxae
with spinulae. Multilocular pores quinqueloculars, and narrow, long macrotubular ducts
scattered all over venter. Microtubular ducts forming a marginal row. Cruciform pores
absent. A few hair-like setae present on submedian venter, short spine-like setae forming
a row on submargin. Suranal setae hair-like.
Dorsum: Enlarged setae pointed, conical; in a row on margin, two on margin of each
segments, and in 6-8 longitudinal rows on dorsum. Macrotubular and microtubular ducts
scattered all over dorsum. Anal ring situated on margin of venter; oval, sclerotized, with
one row of pores and with 6 setae. Anal lobes well developed, membranouos, with 3
pointed, conical spines, ventrally with two flagellate setae; and a long apical seta. Cauda
wide, plate-like.
Adult male described by Afifi (1968).
Ecology
Host plant: Buxus sempervirens.
Distribution: Bulgaria, former Czechoslovakia, France, Germany, Greece, Italy,
Romania, Russia, Spain, Switzerland, Turkey, Ukraine, Uzbekistan, former Yugoslavia.
Biology: On the stems, braches, twigs, and leaves. Sometimes becoming a pest. This
species is monophagus, and has 2 generations per year in Crimea, overwinters as second
instar. First generation females lay eggs in the first part of June, and second generation
females lay in mid-August. Reproduction biparental (Kosztarab & Kozár, 1988).
Comments
"There seems to be no doubt about the identity of this species, even though the original
material cannot be traced. The species was described from Aix-en-Provence and the
specimens at hand collected in Orange, and from Lyons, not far from the type-locality,
are considered by French workers to be this species. A further specimen is available
from material collected in the south of France and this was identified by V. Signoret as
E. buxi " (Williams, 1985a). Marginal setae of first instar nymph and female of E. buxi
are triangular dome-, or thorn shaped and have enlarged tubular ducts, differing from all
other Acanthococcidae genera and species known in the Palearctic Region, with longer,
narrow, spine-like dorsal setae. They are somewhat similar to spines of “Eriococcus”
(Thekes) eucalypti Maskell, 1892 (Kozár, 2009; Ouvrard & Kozár, 2009) from Australia.
Eriococcidae 635
Figure 219. Eriococcus buxi (Boyer de Fonscolombe, 1834), second instar, a female, b
male. Original.
Eriococcus williamsi Danzig, 1987 (Fig. 221)

Eriococcus williamsi Danzig, 1987: 118.
Holotype: Female. Ukraine (Crimea, Massandra), on Buxus sp., 24.vii.1951, by T.

Bushchik (type no. 43-86). By original designation. Deposited in ZMAS. Notes: 1
paratype female on holotype slide; approximately 45 adult female paratypes in ZMAS.
Paratypes also in the BMNH and MNHN.
Lit.: Cook & Gullan, 2001: 61; Cook & Gullan, 2004: 444; Germain, 2008: 78;
Germain et al., 2007: 472; Kozár, 2009: 97; Miller & Gimpel, 2000: 377; Williams,
1985a: 361 (Miller et al., 2013).
Description
Unmounted female
Unknown.
Mounted female
Adult female broadly oval, 2.3 mm long, 1.3 mm. wide, not nodulose. Anal lobes about
twice as long as wide, conical, each lobe with apical setae 180 µm long. Dorsally with
an outer enlarged seta situated submarginally rather than on edge, and 2 inner enlarged
setae; ventrally with a single slender seta and a suranal seta shorter than anal ring setae.
a single pair of setae. Antennae 150–210 µm long, with 6 segments. Legs robust, hind
trochanter+femur 160–190 µm long, hind tibia 80–95 µm long, hind tarsus 80–95 µm
long, the tibia+tarsus either same length as trochanter+femur or slightly shorter, tibia
with four setae, claw 25 µm long, straight with a minute denticle near apex; hind coxa
with a few small translucent pores. Frontal lobes well developed, about as long as width
of basal antennal segment. Setae normal, slender, on median areas and laterally towards
margins. Enlarged setae, around margins only. Quinquelocular pores, numerous on
abdomen where they almost reach margins; others present in median areas of head and
thorax and around spiracular openings, those next to first spiracle extending to margin.
Macrotubular ducts of two sizes, a larger type, about 25 µm long, present around margins
and submargins, reaching to spiracles, a narrower type on abdomen only, in transverse
rows or bands across median areas of segments. Microtubular ducts, 12 µm long, around
margins only. Cruciform pores absent. Anal lobes strongly sclerotized; membraneous
plate present on dorsum anterior of anal ring; microtubular ducts elongate with 1
sclerotized area; orifice bifurcate (Williams, 1985a).
Dorsum: Dorsal enlarged setae scattered numerous, lanceolate, 25–45 µm long, broadly
lanceolate, pointed, curved in profile; group of these setae usually present submarginally
on mid-head region, behind eyes, on prothorax, first and seventh abdominal segments,
each group associated with a single enlarged duct or sometimes 2 each duct 40 µm long
with inner end flat and with sclerotised rim raised from surrounding integument. Other
Eriococcidae 637
Figure 220. Eriococcus williamsi Danzig, 1987, female. After Williams (1985) with
modifications.
enlarged large ducts, other than macroducts present in marginal areas of head, thorax
and abdomen, surrounded by small cluster of enlarged setae, usually with similar ducts in
medial areas of body; sometimes present on submargins without the groups of setae up
to 17 marginal ducts present in some specimens and others present on midline, varying in
number but there is usually one on mesothorax and another on first abdominal segment.
Macrotubular ducts of one size, evenly distributed, about 25 µm long, with a clear circular
rim surrounding orifice, the cup about half width of setal base of dorsal setae, the inner
end filament resembling a buch of grapes. Microtubular ducts in a regular arrangement,
about 12 µm long, very slender, the sides parallel, with orifice widely bifid. Anal ring with
8 setae, each about 185 µm long. Cauda only lightly sclerotised, rounded (after Williams
(1985a); Danzig (1987) with modifications).
Ecology
Host plant: Buxus sp., Buxus sempervirens.
Distribution: Greece, Turkey, Ukraine.
Biology: On the stems, braches, twigs, and leaves. Eriococcus williamsi was originally
confused with E. buxi since they both attack Buxus sempervirens. E. williamsi is a more
eastern species (Danzig, 1987).
Comments
The difference from E. buxi was mentioned by Williams (1985a) as Eriococcus sp. near
buxi, unavailable name.
Kuwaninidae 639
Family:
KUWANINIDAE
Kozár Family nova
Type genus: Kuwanina Cockerell, 1903.

Common name: Bark felt scale.
Lit.: Fernald, 1903: 70; Hardy et al., 2011: 502; Hardy & Gullan, 2007b: 106; Henderson,
2007b: 10; Hoy, 1963: 164; Kawai, 1980: 134; Koteja, 1974a: 297; 1974b: 78; Kozár,
2009: 114; Kozár & Walter, 1985: 76; Lindinger, 1937: 187; Miller, 1970: 157; Miller
& Gimpel, 2000: 390; Morrison & Morrison, 1922: 58; Morrison & Morrison, 1966:
100; Takahashi, 1958: 1; Williams, 1985a: 384; Wu & Li, 2009: 221 (Miller et al., 2013).
Description
In life, adult female hemispherical or spherical, pink (but dead body pale yellow), covered
with white cottony secretion of fine waxythreads, living in crevices of bark on main
trunk of the host.
Mounted female
Adult female oval, dorsal abdominal segments sclerotised. Antennae reduced, generally 3
or 4 segmented. Labium apparently two segmented, with strong basal ring. Legs absent,
all legs represented by vestigial patches. Anal lobe absent. Setae spine like, small, scattered
more numerous posteriorly. Discodial pores generally quinquelocular but varying from
3–7 loculi, also varying in size; the apparently unique invaginated on dorsum and/or
venter. Anal ring either simple or with a non-cellular anal ring, with 0–6 small setae.
Host plant
The members of Kuwaninidae family collected from deciduous plant.
Distribution
Members of this family are known from the Eastern-palearctic-Oriental, and Australasian
regions, with one genus and four species.
Biology
Generally live under bark in fine white sac.
Genus:
Kuwanina Cockerell, 1903
Kuwanina Cockerell in Fernald, 1903: 121.
Type sp.: Sphaerococcus parvus Maskell, by monotypy and original designation.

Lit.: Afifi and Kosztarab, 1967: 3; Boratynski & Williams, 1964: 91; Borchsenius,
1949: 43; Fernald, 1903: 121; Ferris, 1918: 324; 1941: 26; Gullan et al., 2005: 166;
Hardy et al., 2011: 502; Hardy & Gullan, 2007b: 106; Henderson, 2007b: 10; Hoy,
1963: 164; Kawai, 1980: 134; Koteja, 1974a: 269; 1974b: 78; Kozár, 2009: 114; Kozár
& Walter, 1985: 76; Lindinger, 1937: 187; Miller & Gimpel, 2000: 390; Morrison &
Morrison, 1922: 58; Morrison & Morrison, 1966: 100; Takahashi, 1958: 1; Williams,
1985a: 384; Wu & Liu, 2009: 221 (Miller et al., 2013).
Cryptococcus; Lindinger, 1937: 187. Incorrect synonymy; discovered by Kozár & Drozdják,
1998: 413.
Comments
Although many authors have considered this genus to belong in the Pseudococcidae
(Morrison & Morrison, 1966) it is clearly part of the Eriococcidae s.l. (Williams, 1985a,
Hoy, 1963). Kosztarab (1968) considered it to be in a separate family, Cryptococcidae.
Koteja (1974a, b) elevated it to a family status.
Description
Adult female oval, dorsal abdominal segments heavily sclerotised with intersegmental
furrows. Antennae much reduced, 3 or 4 segmented with or without setae. Labium
3 segmented, basal segment with a pair of setae. Legs absent, all legs represented by
vestigial patches. Anal lobes absent. Setae spine-like, small, scattered more numerous
posteriorly. Discodial pores generally quinquelocular but varying from 3-7 loculi, also
varying in size; the apparently unique invaginated on dorsum and/or venter. Anal ring
either simple or with a non-cellular anal ring, with 0-6 small setae.
Kuwaninidae 641
Host plant
Woody plants from Fabaceae, Myrtaceae, Rosaceae, Betulaceae.
Distribution
Eastern-Palearctic, Oriental, and Australasian regions; with 2 species known from the
Far-Eastern part of the Palaearctic Region.
Biology
Generally living under bark within a fine white sac.
Key to species
1. – Metathoracic legs replaced by a ball-shaped, pore bearing flap,

without no pores......................................................................................................K. parva
– Metathoracic legs replaced by a flat pore-plate or vestigial leg
flap with few pores................................................................................................K. betulae
Kuwanina betulae Wu & Liu, 2009 (Fig. 221)

Kuwanina betulae Wu & Liu, 2009: 221.
Holotype: Female. China (Henan Province, Songxian County, Mt. Baiyun, N: 34º08’,
E:112º05’, on Betula ablosinensis Burkill, 15.8.2008, by S. Wu & J. Liu. Deposited
Beijing: Forestry University.
Lit.: Wu & Liu, 2009: 221 Miller et al., (2013).
Description
Unmounted female
In life, adult female hemispherical or spherical, pink (but dead body pale yellow), covered
with white cottony secretion of fine waxy threads.
Mounted female
Body of adult female circular oval, 0.76–0.87 mm long. Antenna reduced, stub-like, 3
segmented, apical segment with 4 small setae. Frontal tubercle and frontal lobes not seen.
Eyes situated on venter near margin. Anal lobes absent.
Venter: Labium 3 segmented, basal segment not well developed, with a pair setae; broad
50–60 µm long, clypeolabral shield nearly rectangular, 100–110 µm long, 80–90 µm wide.
Posterior and mid-legs absent, each hind leg reduced to transverse oval pore plate, 40–50
µm long, 30 µm wide. Spiracles surrounded by a sclerotised plate, each anterior spiracle
with 7-9, posterior ones each with 9-12 quinquelocular pores on surface near atrium.
Setae few, short, small, spinelike, Quinquelocular pores 4 µm in diameter, on margin, and
around mouthparts. Invaginated quinquelocular pores with short tubes, 4 µm long, on
the body margin. Cruciform pores absent. Vulva circular, sclerotised.
Dorsum: Dorsal setae few, short, conical. Quinquelocular pores scattered on body
surface. Invaginated disc pores with short tubes, 4 µm long, scattered through. Anal ring
almost circular, with 9 pores and 6 thick setae, each 13–15 µm long, surrounded setae all
in sclerotised plate. anal ring surrounded by 10 thick setae, of which 8 setae, each 8–9
µm long, 2 setae, 10–15 µm long, anal ring and surrounding setae all in a sclerotised plate
(after Wu & Liu (2009)).
Ecology
Host plant: Betula albosinensis.
Biology: It lives in the crevices of bark on the main trunk.
Kuwaninidae 643
Figure 221. Kuwanina betulae Wu, 2008, female. After Wu & Wei (2003) with modifications.
Kuwanina parva (Maskell, 1897) (Fig. 222)

Sphaerococcus parvus Maskell, 1897: 244.
Syntype: Female. Japan, on Prunus sp., 1896, by A. Koebele. Deposited in CASC,

NZAC, UCDC, and USNM.
Lit.: Afifi & Kosztarab, 1967: 3; Boratynski & Williams, 1964: 91; Deitz & Tocker,
1980: 20; Fernald, 1903: 121; Ferris, 1919: 252; Green, 1915: 181; Hendricks &
Kosztarab, 1999: 165; Hoy, 1963: 165; Kanda, 1941: 12; Kawai, 1972: 5; 1977: 153;
1980: 134; Kosztarab, 1968: 12; Koteja, 1974a: 249; 1974b: 78; 1980: 77; Kozár, 2009:
103; Kozár & Walter, 1985: 76; Kuwana, 1902: 56; 1907: 185; Lindinger, 1932: 188;
Maskell, 1897: 244; Miller & Gimpel, 2000: 391; Miller et al., 1998: 297; Morrison &
Morrison, 1922: 58; Paik, 1978: 169; Williams, 1985a: 384; Wu & Liu, 2009: 221; Yang
& Kosztarab, 1967: 10 (Miller et al., 2013).
Kuwanina parvus; Cockerell in Fernald, 1903: 121. Change of combination.

Sphaeracoccus parvus; Kuwana, 1907: 185. Misspelling of genus name.
Kuwanina parva; Green, 1915: 181. Change of combination.
Cryptococcus nudatus; Brittin, 1915: 160. Green (1916) incorrectly stated that it was a
synonym.
Cryptococcus parvus; Lindinger, 1932: 188. Change of combination.
Kuwanina parva; Hoy, 1962: 26. Revived combination.
Description
Unmounted female
Insects are covered by very loose white cotton which forms distinct sacs, midpart
yellowish-brown, hard, adult female reddish-brown, globular, larva red (Maskell, 1897).
Mounted female
Body of adult female circular oval. Antennae reduced, stub-like, apical segment with
several small setae. Frontal tubercle, lobes and eyes not shown. Anal lobe absent.
Venter: Labium 1 segmented, details not shown, clypeolabral shield nearly rectangular.
Stylet loop reaches behind posterior spiracles. Posterior and mid-legs absent, each hind
leg a ball-shaped, pore bearing flap. Spiracle surrounded by a sclerotised plate, each
anterior spiracle with 2–4, on surface near its atrium. Setae few, short, small, spinelike,
Discodial pores generally quinquelocular but varying from 3–7 loculi, also varying in size,
scattered on submargin and margin, and around mouthparts. Invaginated quinquelocular
pores with short tubes, on the body margin and a few around mouthpart. Cruciform
pores absent. Vulva not shown.
Kuwaninidae 645
Figure 222. Kuwanina parva (Maskell, 1887), female, after Wu & Wei (2003) with
modifications. First instar on top right, after Afifi & Kosztarab (1967) with modifications.
Dorsum: Dorsal setae few, short, conical. Quinquelocular pores few, scattered on surface.
Invaginated quinquelocular pores with short tubes, in rows on abdominal segment, and
on margin. Anal ring almost circular, with 6 thick setae (after Maskell (1897); Morrison &
Morrison (1922); Wu & Liu (2003)).
Other stages
Detailed descriptions and illustrations of first and second-instar nymphs given by Yang
& Kosztarab (1967).
Ecology
Host plant: Prunus sp., P. pseudocerasus.
Distribution: Japan, South Korea, United Kingdom.
Biology: Unknown.
References 647
REFERENCES
Afifi, S. A. (1968): Morphology and taxonomy of the adult males of the families Pseudococcidae and
Eriococcidae (Homoptera: Coccoidea). Bulletin of the British Museum (Natural History) (Entomology)
supplement 13: 1-120.
Afifi, S. A. & Kosztarab, M. (1967): Studies on the morphology and taxonomy of the males of Antonina and one
related genus (Homoptera: Coccoidea, Pseudococcidae). Research Division Bulletin Virginia Polytechnic
Institute and State University, Blacksburg 15: 1-43.
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TAXONOMIC INDEX
A brachypodii 282
brevenniae 418
abaii 184 brucius 464
abeliceae 78 buxi 630
Acalyptococcus 64
Acanthococcidae 59 C
Acanthococcidae FG 57
Acanthococcites 59 cactearum 421
Acanthococcus 72 cantium 422
acericola 80 castanopus 102
aceris 82, 597 caudatus 426
aceris kurdica 82 centaurea 338
Aculeococcus 618 chaoticus 218
acutispinatus 384 cingulatus 193
acutus 397 cingulatus orientalis 193
adzharicus 186 cistacearum 428
agavacearum 375 coccineus 432
agavium 376 confusus 197
agenjoi 302 corniculatus 105
agropyri 189 coronillae 436
alpina 280 costatus 108
altaicus 87 crassispinus 438
ammophilus 400 Cryptococcidae 594
Anophococcus 180 Cryptococcus 595
araucariae 580 cynodontis 202
arboisi 484
D
arbuti 176
armeniacus 90 Dactylopiidae 59
artemisiarum 403 Dactylopites 59
arthrophyti 406 danzigae 197
artiguesi 408 deformis 66
Asiacornococcus 621 desertus 440
astragali 411 devoniensis 443
atlihani 336 distincta 299
azaleae 92 duzgunesae 258
azumae 96 dziedzickae 522
B E
baldonensis 414 echinatus 446
baltica 312 ericae 110
Balticococcus 587 erinaceus 448
betulae 642 Eriococcidae 59, 616
betulaefoliae 98 Eriococcini 59
bezzii 100 Eriococcus 629
borchsenii 416 Eriokermes 260
Borchseniococcus 257 erwini 285
borealis 92 eugeniae 64
Taxonomic index 667
evelinae 200 I
evinae 451
exiguus 622 iljinae 212
exoticus 380 inermis 214
insignis 217
F integricornis 604
isacanthus 116
fagisuga 600 istresianus 484
festucae 454
festucarum 454 J
filifer 358
formicicola 202 japonicus 624
franceschinii 254 juniperi 270
fraxini 609 Jutlandicoccus 590
G K
gassinus 456 kaiseri 344

gavrilovi 458 kaki 626
Gedanicoccus 589 kaschgariae 118
glanduliferus 113 Kaweckia 310
glyceriae 311 Kermidae 59
gnidii 461 Kerminae 59
Gossyparia 261 kilinceri 120
Gossypariella 260, 269 koelreuterius 124
gouxi 286 kondarensis 221
gracilis 589 korotyaevi 326
gracilispinus 341 Kotejacoccus 324
graminicola 189 kotejai 222
graminis 68 Kuenowicoccus 592
gramuntii 265 kurdicus 488
granulatus 205 Kuwanina 640
greeni 464 KUWANINIDAE 639
Greenisca 278
L
Greenoripersia 295
Gregoporia 295, 296 lactucae 490
guesinus 469 laeticoris 352
Gymnococcus 373 lagerstroemiae 126
lanigera 265
H latialis 129
hassanicus 470 lauri 369
helichrysi 473 lerzanae 226
hellenica 316
M
henmii 476
henryi 478 macedoniensis 130
herbaceus 209 marginalis 493
heteroacanthus 481 mariannae 584
Hispaniococcus 300 matesovae 318, 496
hoheriae 365 melnikensis 82, 134
Hujinlinococcus 307 micracanthus 499
minimus 502
miscanthi 504 R
multispinatus 506
multispinosus 509 reynei 528
mumtazi 511 Rhizococcus 390
munroi 513 ribesiae 146
roboris 148
N rosaceus 299
rosannae 152
nedimi 517 rubra 354
nematosphaerus 308 rugosus 534
Neoacanthococcus 334
Neokaweckia 351 S
Neotrichococcus 357
ningxianensis 520 sachalinensis 464
notabilis 361 sachalinensis 536
nudatus 644 salicicola 155
salicicola 262
O salicis 155
salsolae 246
oblicus 587 sanguinairensis 248
oblongus 228 saratogensis 217
obscurus 189 sasae 158
oligacanthus 304 saxatilis 538
onukii 140 saxidesertus 540
orbiculus 328 Scutare 64
oreophila 290 selmae 251
orientalis 320 shiraiwai 160
Orontesicoccus 368 siakwanensis 543
osbeckiae 387 siamensis 274
Ovaticoccus 373 Sisyrococcus 64
oxyacanthus 231 socialis 252
sojae 546
P
spiniferus 548
palustris 522 spinosus 588
pamiricus 345 spiraeae 163
pannonicus 234 spuria 265
Parrotia 373 spurius 265
parva 644 subterraneus 551
parvispinus 239
pauper 591 T
perfectus 591 tamaricicola 348
phyllanthi 273 targassonensis 554
pietrzeniukae 593 terrestris 193
placida 292 teucriicolus 556
podhalensis 522 thaleri 559
polyphagus 478 Thekes 72, 629
populi 142 thymelaeae 562
Proteriococcus 383 thymi 166
pseudinsignis 242 timidus 566
Pseudoacanthococcus 386 tokaedae 170
Pseudochermes 608 tounetae 513
puymorensis 525
Taxonomic index 669
transversus 172 W
trispinatus 70
turcicus 331 wangi 534
turkmenicus 568 williamsi 613, 636
U Y
Uhleria 579 yongpingensis 618

ulmarius 174
Z
ulmi 265, 606
uvae-ursi 177 zernae 573
uvaeursi 176 zygophylli 576
V
vanensis 323
variabilis 570
HOST PLANT INDEX
A Albizia kalkora 118

Alhagi maurorum 578
Abelicea hirta 80 Alnus crispa 268
Acantholimon sp. 358, 360 Alnus glutinosa 268
Aceraceae 594, 596 Alnus sp. 268
Acer campestre 82, 86 Aloe sp. 378
Acer cincinatum 86 Alopecurus pratensis 243
Acer platanoides 86 Alopecurus sp. 243
Acer pseudoplatanus 86, 598 Ammophila arenaria 402, 468
Acer sp. 82, 95, 132, 598 Ammophila arenaria arundinacea 402
Acer trifidum 172 Ammophila sp. 243
Acer truncatum 82 Anaphalis sp. 544
Achillea distans 516 Andropogon sp. 220, 243
Achillea millefolium 196, 243, 450, 516 Aneurolepidium sp. 192
Achillea ponticum 243 Anogeiussus latifolia 128
Achillea sp. 243, 444 Anogeiussus sp. 128
Acroptilon repens 444 Anthoxanthum odoratum 220
Aegilops geniculata 468 Anthoxanthum sp. 220, 243, 316
Aeluropus littoralis 550 Anthusa sp. 168
Aesculus hippocastanum 86, 150 Araucaria angustifolia 582
Aesculus sp. 150 Araucaria araucata 582
Agave americana 378, 382 Araucaria bidwillii 582
Agave beaueriana 378 Araucariaceae 580
Agave decipiens 378 Araucaria columnaris 582
Agave lecheguilla 378 Araucaria cunninghamii 582
Agave lophantha var. poselgeri 378 Araucaria heterophylla 582
Agave parryi 378 Araucaria hunsteinii 582
Agave sisalana 378 Araucaria orientalis 582
Agave sp. 378 Araucaria sp. 582
Agave triangularis 378 Arbutus sp. 178
Agave utahensis var. nevadensis 378 Arctostaphylos uva-ursi 178
Agropyron capillaris 220, 316, 354 Ariocarpus trigoni 422
Agropyron cristatum 192, 220 Arrhenaterum elatius 243
Agropyron curtisii 220 Arrhenatherum elatius 220, 316
Agropyron fragile 192, 320 Artemisia absinthium 196
Agropyron intermedium 468 Artemisia argyi 504
Agropyron repens 192, 220, 222, 243, 294, 316, Artemisia austriaca 196
444, 468, 574 Artemisia dracunculus 196
Agropyron rupestris 220 Artemisia frigida 498, 540
Agropyron sibiricum 192 Artemisia gallica 242
Agropyron sp. 192, 222, 230, 239, 243 Artemisia gmelini 428
Agropyron tsukushiense var. transiens 316 Artemisia gurganica 404
Agrostis canina 316 Artemisia herba-alba 168
Agrostis capillaris 243 Artemisia maritima 498
Agrostis curtisii 216 Artemisia marschalliana 540
Agrostis gigantea 316 Artemisia nana 168
Agrostis sp. 192, 220, 316, 354, 356 Artemisia ordosica 196, 508
Ajuga sp. 540
Host plant index 671
Artemisia schrenciana 196 Buxus microphylla koreana 128

Artemisia sp. 178, 239, 242, 302, 304, 418, 428, Buxus sempervirens 634, 638
440, 444, 458, 460, 508, 516, 536, 552, Buxus sinica 80
569, 578 Buxus sp. 617, 638
Artemisia sphaerocephala 440, 442
Artemisia sublissingiana 196 C
Artemisia terrae-albae 404, 498
Calamagrostis arundinacea 220
Artemisia vulgaris 540, 574
Calamagrostis brachytricha 468
Arthrocnemium macrostachyum 431
Calamagrostis canadensis var. langsdorffii 468
Arthrocnemum macrostachyum 114
Calamagrostis epigeios 468
Arundinaria debilis 174
Calamagrostis sp. 210, 243, 284, 288, 316
Arundinaria hindsii var. graminae 142
Calluna sp. 168, 516
Arundinaria sp. 174
Calluna vulgaris 112, 416, 420, 444
Asplenium ceterach 542
Calystegia hederacea 548
Asteraceae 374, 486, 556
Camellia oleosa 106, 628
Astragalus sp. 196, 412
Camphorosma lessingii 442
Astrophytum ornatum 422
Camphorosma monspeliacum 442
Astrophytum sp. 434
Camphorosma sp. 196
Austrocylindropuntia vestita 510
Carex sp. 210, 220, 243, 288, 316, 462, 468,
Avena fatua 230
524
Avena flavescens 294
Carpesium abrotanoides 544
Avenastrum pubescens 220
Carpesium sp. 544
Avenastrum sp. 220
Carpinus betulus 86, 132
Azalea hybrid ‘hino de giri’ 95
Castanea sativa 150, 154
Azalea indica 95, 178
Castanea sp. 150
Azalea nudiflora 95
Castanopsis sp. 102, 104
Azalea sp. 95, 178
Celtis sinensis 128
B Celtis sp. 95
Centaurea cyanus 196
Bambusa dissemulator var. hispida 174 Centaurea orientalis 196
Bambusa nana var. normalis 142 Centaurea sibirica 540
Bambusa sp. 64, 66, 68, 96 Centaurea solstitialis 168, 340
Bambusa textilis 174 Centaurea sp. 335
Bambusa vulgaris 174 Cephalanoplos segetum 548
Betula albosinensis 642 Cereus sp. 422, 434
Betulaceae 641 Chamaecyparis obtusa var. breviramea 105
Betula fruticosa 164 Chrysanthemum sp. 477, 516
Betula platyphylla 164 Chrysopogon gryllus 468
Brachypodium pinnatum 243, 284, 288, 294, 300, Chrysopogon sp. 68
398, 448, 516 Cichorium intybus 220, 444
Brachypodium sp. 210, 220, 284, 286, 288, 292, Cichorium sp. 220
316, 468 Cissus sp. 192, 230
Brachypodium sylvaticum 243, 294, 422, 424 Cistus albidus 431
Bromus erectus 284, 300 Cistus monpeliensis 431
Bromus inermis 220, 243 Cistus salviifolius 486
Bromus mollis 220 Cleistocactus sp. 434
Bromus sp. 202, 220, 222, 228, 239, 243, 284, Clematis songorica 196
288 Clematis sp. 196
Buxaceae 630 Coffea arabica 614
Buxus microphylla 80 Conyza canadensis 516
Coronilla varia 436 Erica multiflora 112, 444

Corylus sp. 268 Erica scoparia 168
Corynephorus canescens 316, 468 Erica sp. 178, 444
Crataegus coccinea 95 Erica tetralix 112, 168, 444, 566, 655
Crataegus sp. 95 Eriophorum vaginatum 524
Crepis sp. 516 Eriophyllum sp. 178
Crithmum maritimum 178 Erodium sp. 444
Crithmum sp. 178 Eubotryoides grayana 178
Cupressus sp. 272, 582 Eubotryoides sp. 178
Cycas sp. 582 Euphorbia characias 480, 532
Cynodon dactylon 192, 202, 205, 210, 220, 224, Euphorbia pithyusa 480
230, 243, 256, 572 Euphorbia sp. 452, 516, 518, 540, 578
Cynodon sp. 239, 300 Euphorbia spinosa 480
Cyperaceae 279
F
D
Fabaceae 641
Dactylis glomerata 220, 239 Fagaceae 383, 594, 596
Dactylis glomerata hispanica 398 Fagus grandifolia 601
Dactylis hispanica 398 Fagus orientalis 601
Dactylis sp. 468 Fagus sylvatica 86, 601
Dalbergia sp. 128 Festuca montana 188
Daphne gnidium 168, 462 Festuca ovina 208, 216, 220, 288, 316, 468
Dendranthema sp. 544 Festuca parvigluma 455
Deschampsia flexuosa 216, 220, 243 Festuca pratensis 220, 468, 516
Deschampsia sp. 220, 468, 516 Festuca rubra 208, 220, 316
Dianthus capitatus 540 Festuca sp. 205, 220, 230, 243, 284, 294, 316,
Dianthus carthusianorum 480 354, 356, 468, 516
Dianthus crinitus 243 Festuca sulcata 316, 354, 516
Dianthus sp. 196, 444, 500 Festuca supina 316
Diospyros kaki 128, 628 Ficus carica 128, 628
Dodartia orientalis 578 Ficus sp. 276
Dracaena sp. 378 Foeniculum vulgare 500
Duschekia fruticosa 164 Fragaria sp. 516, 540
Fraxinus angustifolia 612
E Fraxinus caroliniana 612
Fraxinus excelsior 612
Echinocactus sellowii 422
Fraxinus mandschurica 612
Echinocactus sp. 434
Fraxinus ornus 612
Echinocereus sp. 434
Fraxinus sp. 268, 612
Echinopsis sp. 422, 434
Fremontodendron sp. 95
Elaeagnus angustifolia 86
Fumana glutinosa 482
Eleusine indica 582
Furcraea selloa 378
Elymus angustus 192
Elymus arenarius 316 G
Elymus giganteus 192
Elymus sp. 192, 205, 220, 354, 356 Galium sp. 196, 462, 500
Elytrigia sp. 316 Gaylussacia sp. 95
Empetrum nigrum 416 Genista sagittalis 480
Erica arborea 112, 168, 444 Glochidion puberum 128
Erica carnea 112, 168, 560 Glyceria maritima 316
Erica cinerea 444 Glyceria sp. 316
Glycine max 128, 548 L

Glycine soja 548
Lactuca sp. 492
H Lagerstroemia indica 128
Lagerstroemia japonica 128
Haloxylon ammodendron 408 Lagerstroemia sp. 388
Haloxylon sp. 186, 408 Lagerstroemia speciosa 128, 628
Helianthemum polifolium 431 Lauraceae 618, 620
Helianthemum sp. 516 Laurus nobilis 369, 372
Helichrysum sp. 486 Lavandula sp. 444
Helichrysum stoechas 474 Leontodon crispus 516
Hieracium pilosella 220 Leptospermum scoparium 586, 661
Hieracium sp. 480, 516 Ligustrum obtusifolium 128
Hierochloa odorata 316 Ligustrum sp. 612
Hoheria populnea 366 Linnaea borealis 220
Hoheria sp. 366 Liquidambar sp. 95
Holcus lanatus 220, 243 Lithocarpus sp. 385
Holcus mollis 243 L. nemorosa 210
Holcus sp. 243 Lolium perenne 284, 316, 468
Hordeum sp. 222 Lolium sp. 192, 284, 468
Hylocereus sp. 434 Lotus cystisoides 458
Hylotelephium telephium 540 Luzula pilosa 524
Hyparrhenia hirta 205 Luzula sp. 210, 220, 243
Hypericum sp. 220
Hypochoeris sp. 516 M
Hystrix patula 217, 220
Mallotus japonicus 128
I Malus pumila 128
Malus sp. 86
Ilex sp. 444 Malus sylvestris 220
Iljinia regelii 212, 214 Mammillaria sp. 422, 434
Medicago sativa 196
J
Melica sp. 468
Juniperus phoenicea 272 Melica turczaninowiana 468
Juniperus sp. 582 Melilotus sp. 232
Mentha sp. 540
K Minuartia anatolica 516
Minuartia sp. 442, 516
Kalidium gracile 192, 214, 306 Miscanthus sp. 506
Kalidium sp. 304 Molinia caerulea 288
Kaschgaria komarovi 120 Moreaceae 621
Kochia prostrata 442, 494 Myrtaceae 580, 641, 654, 661
Koeleria askoldensis 216, 220 Myrtus communis 86, 137
Koeleria glauca 243 Myrtus sp. 128
Koeleria macrantha 196, 468
Koeleria marchantha 462 N
Koeleria sp 196, 220, 316
Koelreuteria paniculata 126 Nardus sp. 220
Kunzea capitata 582 Nardus stricta 468
Neobuxbaumia polylopha 434
O Potentilla chinensis 500

Potentilla megalantha 516
Oleaceae 594, 608 Potentilla sp. 196, 468, 516
Opuntia ficus barbarica 434 Poterium spinosum 564
Opuntia sp. 434 Prunus armeniaca 90, 628
Osbeckia sp. 388 Prunus pseudocerasus 646
Prunus sp. 646
P
Pteridium sp. 220
Panderia pilosa 260 Pteris sp. 220
Pasania sp. 385 Pterocarya pterocarpa 150
Pelecyphora sp. 434 Puccinellia sp. 316
Perotis indica 64, 66 Punica granatum 128, 442
Perotis sp. 64, 66 Pyrethrum sp. 196
Phalaris sp. 220, 252 Pyrus betulaefolia 98
Phleum phleoides 220 Pyrus sp. 86, 598
Phleum pratense 220, 316
Phleum sp. 286, 468 Q
Phragmites communis 70 Q. robur imeretina 150
Phragmites sp. 243 Quercus coccifera 564
Phyllanthus emblica 274 Quercus ilex 132
Phyllostachys glauca 310 Quercus petraea 150
Phyllostachys nigra 310 Quercus pubescens 86, 137, 150
Phyllostachys nuda 310 Quercus robur 86, 150
Phyllostachys praecox 310 Quercus sp. 123, 137, 150, 332, 385
Phyllostachys propinqua 310
Phyllostachys pubescens 310, 536 R
Phyllostachys rubella 310
Phyllostachys sp. 310 Reaumuria soongarica 328
Phyllostachys viridis 310 Rebutia sp. 434
Pinus pinea 582 Rhipsalis sp. 434
Pinus sp. 198 Rhododendron catawbiense 95
Pinus sylvestris 601 Rhododendron ferrugineum 102
Piptatherum miliaceum 210 Rhododendron sp. 95, 178
Piptatherum multiflorum 398 Rhododendron tolmachevii 178
Piptatherum songaricum 316 Rhododendron tomentosum 416
Pistacia weinmannifolia 544 Ribes meyeri 148
Plantago sp. 540 Ribes sp. 95
Platanus orientalis 86, 489 Rosaceae 594, 596, 608, 621, 641
Poa angustifolia 316 Rosa sp. 622
Poaceae 198, 250, 279, 282, 296, 298, 322, 324, Rubiaceae 608
374, 410, 462, 469, 472, 512, 537 Rubus sp. 128
Poa compressa 316 Rumex sp. 220
Poa pratensis 316 Russelia sp. 196
Poa sp. 220, 288, 468 Ruta montana 564
Polygonum sp. 350
S
Populus euphratica 144
Populus pruinosa 144 Salicaceae 594, 608
Populus sp. 95, 144 Salicornia fruticosa 114, 444
Populus tremula 612 Salicornia sp. 168, 444, 552
Potentilla anserina 540 Salix alba 156
Salix capitata 264 Ternstroemia japonica 106

Salix caprea 86 Ternstroemia japonica var. wightii 106
Salix sp. 88, 95, 144, 156, 164, 196, 264, 268 Ternstroemia sp. 628
Salix wilhelmsiana 144 Tetragonolobus sp. 516
Salsola sp. 248, 362 Teucrium chamaedrys 540
Salvia sp. 444, 500 Teucrium polium 558
Santolina rosmarinifolia 168 Teucrium scorodonia 480
Santolina sp. 444 Teucrium sp. 168, 516
Sarcopoterium spinosum 516 Theaceae 621
Sasa peniculata 142 Thelocactus bicolor 434
Sasa sp. 142, 160 Thuja sp. 95
Saussurea japonica 516 Thymelaea hirsuta 564
Saussurea sp. 516 Thymelaea villosa 168
Saxifraga sp. 220 Thymelaea virgata var. broussonetii 564
Scabiosa columbaria 480 Thymus caespititius 562, 564
Scabiosa fischeri 500 Thymus glabrescens 410, 462
Scabiosa lachnophylla 500 Thymus mastichina 168
Scabiosa sp. 192, 442, 518 Thymus quinquecostatus var. przewalskii 516
Scabiosa ucranica 196 Thymus serpyllum 532
Secale cereale 316 Thymus sp. 220, 444, 516
Sedum sp. 205, 316, 480, 516 Thymus vulgaris 168
Selenicereus sp. 434 Tilia amurensis 606
Seseli sp. 462 Tiliaceae 594, 596
Setaria sp. 205, 243 Tilia sp. 598
Silene brahuica 498 Trifolium medium 220
Silene gallica 562, 564 Trigonella ruthenica 196
Silene sp. 220, 412 Trigonella sp. 196
Sorbus aucuparia 612 Trinia sp. 196
Spiraea crenata 164 Triticum aestivum 316
Spiraea douglasii 220 Triticum turanicum 574
Spiraea hypericifolia 164 Triticum vulgare 316
Spiraea media 164
Spiraea salicifolia 118, 164 U
Spiraea sp. 164
Ulex europaeus 444
Stipa baicalensis 356
Ulex sp. 220
Stipa pennata 468
Ulmus americana 268
Stipa sp. 196, 220, 284, 354, 356
Ulmus angustifolia 268
Suaeda vera 168
Ulmus campestris 86
Symplocos myrtacea 624
Ulmus campestris var. pyramidalis 268
Syringa sp. 612
Ulmus davidiana 108, 110, 174, 176
T Ulmus davidiana var. japonica 110, 176
Ulmus glabra 268
Tamarix gracilis 330 Ulmus glabra var. camperdownii 268
Tamarix leptostachys 330 Ulmus laevis 268
Tamarix mannifera 344 Ulmus minor 268
Tamarix ramosissima 330 Ulmus procera 268
Tamarix sp. 335, 338, 342, 346, 350 Ulmus pumila 110, 176, 268, 606
Taraxacum mongolicum 508 Ulmus rubra 268
Taraxacum sp. 196, 243, 444 Ulmus sp. 80, 220, 268, 606
Ternstroemia gymnanthera 106 Ulmus thomasii 268
Ulmus x hollandica Dampieri 268 Vicia sp. 516

Urtica dioica 220 Viscum sp. 268
Urtica sp. 220 Vitex sampsoni 230
Vitis sp. 196
V
W
Vaccinium hirtum 178
Vaccinium macrocarpon 95 Wilcoxia sp. 434
Vaccinium myrtillus 178
Vaccinium praestans 178 Y
Vaccinium sp. 95, 444
Youngia tenuicaulis 440, 442
Vaccinium uliginosum 178, 416
Yucca sp. 376, 378
Vaccinium vitis-idaea 178, 416
Valeriana officinalis 516 Z
Valeriana sp. 516
Veronica chamaedrys 243 Zerna cappadocica 574
Veronica multifida 444 Ziziphora clinopodioides 540
Veronica sp. 243, 462, 516 Zygophyllum fabago 578
Distribution index 677
DISTRIBUTION INDEX
A D
Algeria 112, 205, 444, 482, 564, 582 Denmark 220, 268, 444, 587, 588, 590, 591,
Argentina 582 592, 602, 612
Armenia 150, 220, 440, 468, 494, 540, 552,
567, 602 E
Australia 434, 582
Egypt 344, 434, 564, 582
Austria 86, 112, 178, 220, 244, 268, 316, 444,
Eritrea 380
560, 602, 612
Ethiopia 380
Azerbaijan 150, 540, 598
F
B
Fiji 582
Belgium 95, 602, 612
Finland 268, 416, 566, 612
Brazil 434, 582
France 86, 112, 114, 150, 168, 178, 208, 216,
Bulgaria 33, 86, 137, 192, 205, 220, 244, 268,
220, 242, 244, 250, 256, 268, 272, 288,
288, 442, 516, 532, 602, 612, 634
300, 316, 380, 398, 402, 410, 420, 431,
C 434, 444, 448, 458, 461, 462, 468, 469,
474, 480, 486, 516, 526, 532, 556, 572,
Canada 95, 268, 468, 602 582, 586, 602, 612, 634
Chile 582 France (New Caledonia) 582
China 68, 70, 80, 82, 90, 110, 118, 126, 128,
142, 144, 156, 164, 174, 176, 178, 192, G
196, 222, 230, 268, 310, 354, 385, 388,
Georgia 150, 164, 188, 216, 220, 268, 288,
418, 428, 440, 442, 494, 504, 508, 522,
602
532, 536, 542, 544, 548, 578, 598, 606,
Germany 86, 95, 102, 112, 168, 178, 216, 220,
612, 622, 628, 642
244, 268, 294, 316, 434, 444, 510, 516,
China (Beijing) 70
532, 582, 598, 602, 612, 634
China (Guangdong) 66, 667
Greece 192, 202, 205, 210, 224, 268, 318, 416,
China (Guangxi) 104
468, 486, 488, 516, 532, 564, 582, 612,
China (Hainan) 66
634, 638
China (Inner Mongolia) 264, 316, 354
Guatemala 582
China (Ningxia) 98
China (Taiwan) 174 H
China (Tibet) 66
China (Yunnan) 106, 276, 620 Hungary 33, 86, 137, 150, 168, 192, 196, 205,
Cook Islands 582 208, 210, 220, 239, 244, 268, 284, 288,
Costa Rica 582 294, 298, 316, 354, 380, 410, 416, 424,
Croatia 86, 205, 268, 298, 434, 582, 602, 612 431, 442, 444, 448, 462, 468, 486, 488,
Cuba 582 500, 516, 532, 536, 556, 574, 598, 602, 612
Cyprus 86, 137, 205, 564
Czechoslovakia (former) 86, 150, 220, 244, I
268, 284, 294, 316, 354, 434, 602, 612, 634 Iceland 208
Czech Republic 582 India 128, 174, 274, 582
Iran 186, 196, 222, 250, 268, 346, 532, 542,
582, 602, 612
Iraq 86, 220, 268, 489, 512, 558
Ireland 444 Poland 86, 132, 192, 198, 210, 216, 220, 244,
Israel 53, 137, 168, 434, 444, 564, 582 268, 284, 288, 316, 354, 424, 444, 468,
Italy 86, 102, 112, 130, 150, 154, 168, 178, 516, 524, 587, 588, 589, 590, 592, 593,
192, 196, 205, 220, 244, 268, 282, 284, 602, 612
288, 292, 294, 300, 316, 376, 380, 382, Portugal 150, 168, 268, 564, 582, 612
398, 422, 434, 444, 448, 468, 482, 500, Puerto Rico 582
516, 532, 582, 586, 602, 612, 634
R
J
Romania 86, 150, 192, 196, 216, 220, 268,
Japan 68, 80, 96, 105, 128, 142, 162, 172, 193, 284, 316, 340, 450, 468, 516, 578, 602,
268, 434, 455, 477, 506, 548, 624, 628, 646 612, 634
Russia 86, 95, 110, 118, 142, 148, 150, 156,
K 160, 164, 176, 178, 192, 196, 198, 205,
Kazakhstan 88, 144, 148, 150, 164, 192, 196, 210, 216, 220, 222, 232, 268, 284, 288,
220, 264, 284, 316, 330, 342, 356, 404, 294, 300, 316, 320, 356, 416, 418, 422,
442, 468, 498, 516, 540 436, 468, 472, 500, 516, 537, 582, 598,
Kenya 582 602, 606, 612, 634
Kyrgyzstan 264, 356 S
L Slovakia 288
Latvia 268, 284, 316, 416 Slovenia 86, 202, 205, 244, 268, 602, 612
Lithuania 216, 268, 284, 416, 612 South Africa 434, 582
Luxembourg 612 South Korea 128, 196, 316, 548, 646
Spain 102, 112, 168, 205, 268, 304, 434, 444,
M 468, 552, 564, 582, 612, 614, 634
Sri Lanka 174, 388, 582
Macedonia 132 Sweden 178, 192, 220, 244, 268, 288, 444,
Malaysia 582 468, 602, 612
Malta 112, 205, 422, 444, 582 Switzerland 86, 132, 150, 168, 178, 205, 268,
Mauritius 582 286, 288, 292, 444, 468, 524, 602, 612, 634
Mexico 162, 434, 582
Moldova 86, 192, 268, 468, 516, 532 T
Monaco 582
Mongolia 120, 128, 156, 176, 196, 214, 268, Tajikistan 192, 222, 230, 248, 264, 346, 360,
306, 328, 330, 356, 408, 418, 442, 578 362, 492, 542, 550
Morocco 268, 422, 564, 582 Thailand 276
Tunisia 244, 564
N Turkey 123, 150, 156, 168, 192, 196, 205, 210,
222, 228, 244, 250, 252, 260, 268, 324,
Netherlands 86, 112, 150, 198, 216, 220, 268, 332, 338, 372, 412, 444, 452, 468, 486,
444, 468, 582, 602, 612 500, 516, 518, 532, 540, 542, 572, 574,
New Zealand 366, 434, 582, 594 582, 602, 612, 634, 638
Nicaragua 582 Turkmenistan 86, 264, 350, 408, 442, 460,
North Korea 118, 176, 322, 418, 500 569, 578
Norway 220, 268, 612
U
P
Ukraine 86, 150, 178, 192, 196, 205, 210, 216,
Panama 582 220, 244, 268, 316, 354, 380, 416, 436,
Papua New Guinea 582 450, 468, 500, 516, 540, 574, 598, 602,
Philippines 582 612, 634, 638
Distribution index 679
United Kingdom 128, 216, 220, 244, 268, 294, Y

316, 380, 424, 434, 444, 468, 516, 582,
602, 612, 646 Yugoslavia (former) 86, 244, 268, 298, 316,
United Kingdom (Scilly Isles) 366 434, 582, 602, 612, 634
United Kingrom (Bermuda) 582
Z
Uruguay 582
USA 95, 220, 268, 380, 582, 602 Zimbabwe 582
USA (Hawaiian Islands) 434
Uzbekistan 192, 222, 264, 268, 552, 578, 634
V
Venezuela 582

Kozar Et Al - Acanthococcidae and Related Families of The Palaearctic Region

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ACANTHOCOCCIDAE

AND RELATED FAMILIES

*Plant Protection Institute, Centre for Agricultural Research,

Plant Protection Institute, Centre for Agricultural Research,

All rights reserved.

Technical editor: Éva Szita

Printed by Akaprint Nyomdaipari Kft., Budapest, Hungary

CRYPTOCOCCIDAE Kosztarab, 1967.............................................................................594

This work on the FAMILY GROUP “ACANTHOCOCCIDAE” deals with 4 families

Unmounted adult female

Mounted adult female

Figure 1 a-d. a – Gossyparia spurious, b – Rhizococcus coronillae male test,

Figure 1 e-h. e – Eriococcus sp., f – Ovaticoccus agavium, g – Ovaticoccus agavium male

Figure 2. Antennae of Acanthococcidae family group: a – Acanthococcus aceris,

Figure 3. Labium of Acanthococcidae family group. a – Acanthococcus aceris, b –

Figure 4. Legs of Acanthococcidae family group. a – Acalyptococcus deformans, b –

Figure 5. Varieties of pores of Acanthococcidae family group, a – Anophococcus formicicola,

Figure 7. Macrotubular ducts of Acanthococcidae family group: a – Acanthococcus

Figure 8. Different kinds of enlarged setae of Acanthococcidae family group: a –

Figure 9. Different kinds of anal lobes of Acanthococcidae family group: a – Acanthococcus

Morphology of the slide-mounted male

Although, In “A Systematic Catalogue of the Eriococcidae (Felt Scales) (Hemiptera:

The Acanthococcidae FG is considered sister to all other neococcids and the

Anophococcus herbaceus Van - Gevaş - Tatvan ,

Greenisca placida Bükkszentlélek,

According to the survey by Kozár (2009), the Acanthococcidae FG contains 628

Host plant relationships

present knowledge that the Acanthococcidae FG originated in the Upper Cretaceous.

COLLECTING AND MOUNTING METHODS

For mounting, Wilkey’s method is suggested (Kosztarab & Kozár, 1988), as

DEPOSITORIES OF TYPES AND STUDIED INSECTS

Abbreviations of depositories according to ScaleNet:

AAKA: Institute of Zoology, Kazakhstan Academy of Science, Almaty, Kazakhstan

UCD: The Bohart Museum of Entomology, University of California, Davis, California,

Key to the superfamilies

1 – Abdominal spiracles present............................ Orthezioidea (= Archaeococcoids)

Key to the family groups in Coccoidea

Acanthococcites Signoret. 1875: 314.

Family group diagnosis

Key to the families in the Acathococcidae FG present in the Palaearctic Region

1. – Basal segment of labium with two pairs of setae ..........................Acanthococcidae

Type genus: Acanthococcus Signoret, 1875: 16.

Acanthococcites Signoret. 1875: 314.

Mounted adult female

Mounted adult male

1. – Base of enlarged setae with fused microtubular ducts..................................................2

13. – Dorsum with cruciform pores or discoidal pores or with both................................14

1. – Middorsal setae much smaller than the marginal ones.............................. A. trispinatus

Acalyptococcus deformis (Wang, 1974) (Fig. 26)

Acanthococcus deformis; Kozár & Walter, 1985: 74. Change of combination.

Acalyptococcus graminis (Maskell, 1897) (Fig. 27) Combination nova

Syntype: Female. China (Hong Kong). Deposited in UCDC, USNM, NZAC.

Nidularia graminis; Lindinger, 1933a: 116. Change of combination.

Acalyptococcus trispinatus (Wang, 1974) (Fig. 28) Combination nova

Holotype: Female. China (Beijing). By original designation. Deposited in IEBC.

Rhizococcus trispinatus; Kozár & Walter, 1985: 75. Change of combination.

Acanthococcus Kiritchenko 1936: 156 is a junior homonym of Acanthococcus Signoret 1875

1. – Enlarged dorsal setae on abdominal segment VIII present.........................................2