System of generic groups in mealybugs (Homoptera: Coccinea: Pseudococcidae)

2015·0.78 MB·

by  I. A. Gavrilov-Zimin

ZOOSYSTEMATICA ROSSICA, 24(2): 236-259 25 DECEMBER 2015
System of generic groups in mealybugs (Homoptera: Coccinea:
Pseudococcidae)
Система групп родов мучнистых червецов (Homoptera: Coccinea:
Pseudococcidae)
I.A. GAVRILOV-ZIMIN
И.А. ГАВРИЛОВ-ЗИМИН
Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia.
E-mail: [email protected]
The paper provides a brief conspectus of the system of morphological generic groups, elaborated ear-
lier by the author basing on the total taxonomic revision of Palaearctic mealybugs. Here the system is
complemented by the analysis of all 249 genera of the world fauna. Borders of two generic groups are
reconsidered and two else groups (with mainly Oriental and Australasian genera) are included in the
system. Main taxonomic characters of generic rank are discussed and illustrated.
Статья представляет собой краткий конспект системы морфологических групп родов
мучнистых червецов, предложенной автором ранее на основе тотальной ревизии
палеарктической фауны. В настоящей работе эта система дополнена обзором всех 249 родов
мировой фауны псевдококцид. Объем двух родовых групп пересмотрен; еще две группы,
включающие преимущественно рода ориентальной и австралазийской фаун добавлены в
систему. Рассмотрены и проиллюстрированы основные таксономические признаки родового
уровня.
Key words: scale insects, mealybugs, systematics, morphology, ovoviviparity
Ключевые слова: кокциды, мучнистые червецы, систематика, морфология, яйцеживо-
рождение
INTRODUCTION parative analysis of the basic morphologi-
cal characters of mealybugs, discussed in
The Pseudococcidae or mealybugs are detail all main modern contradictions in
one of two largest scale insect families and the higher classification of the family and
comprises more than 2000 species in world introduced a system of 19 morphological
fauna. Unfortunately, general classifica- generic groups, most of which seem to
tion and phylogeny reconstruction for the be monophyletic (holophyletic or para-
family have been poorly elaborated till now phyletic) and will be probably considered
in view of the numerous problems with in future as taxa of the formal nomen-
the correct, unequivocal evolutional inter- clatural rank (for example, tribes). Some
pretation of the mealybug morphological other morphological groups are perhaps
(and some other) characters as plesiomor- polyphyletic and combine genera which
phic, apomorphic, convergent or parallel demonstrate convergent similarity only;
(homoplasy). In our recent total review of however, the decision of this problem is
Palaearctic mealybugs (Danzig & Gavrilov- impossible basing on the present basis of
Zimin, 2014; 2015) we have provided com- our knowledge.
© 2015 Zoological institute, Russian Academy of Sciences
I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 237
Here I am providing a synopsis of the DISCUSSION
elaborated system, a discussion of some ad-
The main taxonomic characters used
ditions to the system, basing on analysis of
in the analysis of the genera (Fig. 1)
non-Palaearctic genera and a conspectus
of all mealybug genera of the world fauna [authorship of the generic names see below in
with preliminary grouping of them accord- the system of generic groups]
ing to the system. Two generic groups used
Wax glands (Table). The structure and
in the revision of Palaearctic fauna (Danzig
distribution of wax glands are the main di-
& Gavrilov-Zimin, 2014, 2015), g/g Trabu-
agnostic characters in the systematics of
tina and g/g Planococcus, are included here
mealybugs and all scale insects. Cuticular
correspondingly to g/g Nipaecoccus and
sclerotized parts of these glands are clearly
g/g Trionymus (see below). On the other
visible because they absorb stain during
hand, two other groups, which combine
preparation of scale insects and can be sub-
mainly Oriental and Australasian genera,
divided into 3 main groups: discoidal glands,
g/g Pedronia and g/g Allomyrmococcus are
cylindrical glands and glands connected
added to the system.
with conical setae (“spines” in non-English
The analysis and discussion below are
literature). {In the English coccidological
based on the study of the type species of
tradition the term “wax glands” is usually
249 genera, most of which are represented
not used, probably because the soft parts of
(as type and non-type specimens) in the
the glands are lost during preparation and
collections of Zoological Institute, Rus-
only “ducts” and “pores” are visible on spec-
sian Academy of Sciences (St Petersburg)
imens mounted in Canada-balsam. In Rus-
and Muséum National d’Histoire Naturelle
sian and other old European coccidological
(Paris). Some material was also loaned by
schools the term “glands” is widely used, be-
me from the collection of the British Mu-
cause there are no special terms to indicate
seum of Natural History (London) and
the soft parts of the wax glands.}
Plant Protection Institute of Hungarian
Discoidal glands produce powdery wax
Academy of Sciences (Budapest). In addi-
and can be subdivided in the following sub-
tion to the study of the materal I used for
types.
the analysis the morphological descriptions
1) Simple discoidal pores; have a single
and illustrations published in the compre- opening (loculus). Such pores are often
hensive faunistic reviews by following au- scattered in more or less quantity on both
thors for the main non-Palaearctic zoogeo- body surfaces of mealybugs and/or present
graphical regions of the world: for Nearctic in groups with other wax glands or associat-
(Ferris, 1950, 1953; McKenzie, 1967), for ed with different types of setae. Monotypic
Neotropical (Williams & Granara de Wil- genus Hopefoldia is characterized by very
link, 1992), for Oriental (Williams, 2004) peculiar compact groups of discoidal pores,
and for Australasian (Ferris, 1948; Beards- so-called “poraria” located on sclerotized
ley, 1964, 1966, 1971; Williams, 1985; Cox, plates on both body sides.
1987; Williams & Watson, 1988). The Afro- 2) Trilocular (3-locular) pores have 3
tropical region has not been covered by the excretory openings and approximately tri-
general taxonomic revision till now, but the angular in form. These pores are one of the
following basic publications help to unders- synapomorphic characters of mealybugs and
tand the main morphological trends in the the monotypic family Phenacoleachiidae
evolution of the mealybugs of the region: and can be found in majority of mealybugs
Mamet, 1940, 1951, 1954, 1962, 1967; De on all body surface. Only some mealybug
Lotto, 1957, 1958, 1964, 1967, 1969, 1977; genera, such as from g/g Heterococcus, g/g
Millar, 2002). Boreococcus, g/g Neotrionymus, g/g Metad-
© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259
238 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS
Quinquelocular pore
Eye
Claw digitules
Anterior ostiolus
Claw with denticle
Anterior spiracle
Trilocular pore
Simple pore
Conical seta
Circuli
Flagellate setae
Simple tubular duct
Pores
Spinulae
Anus
Multilocular pore
Cerarius C Vulva
18
Setae of anal
apparatus
Fig. 1. Main taxonomic characters of mealybug females (Fonscolombia tomlinii (Newstead, 1892),
Russia: Voronezh Prov.).
© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259
I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 239
enopus, and also some genera from other Cylindrical glands (Table 1) produce
groups demonstrate reduction in the num- wax filaments. There is a great diversity of
ber of trilocular pores or their secondary these glands, probably, exceeding the diver-
total absence. Sometimes trilocular pores sity of such glands in other scale insects all
are scattered on body very irregularly; thus, together.
they can be absent or very few in medial 1) Simple tubular ducts. They have one
zone of the thoracic and anterior abdominal simple excretory duct. The opening of this
sternites (as, for example, in some species of duct usually lies on the surface of the cu-
the species-rich genus Phenacoccus) or form ticle or more rarely it is slightly raised or
dense marginal or medial bands (as in Trim- slightly invaginated from the surrounded
erococcus and Kiritshenkella, respectively). cuticle area. Most genera of subfamily Phe-
The size of the trilocular pores is usually nacoccinae have only this type of cylindri-
equal throughout the body. However, in the cal glands. The deviation of this type is so-
species of g/g Puto, dorsal trilocular pores called “bottle-shaped” ducts in the species
(and especially cerarian ones) are larger of g/g Coccidohystrix: distal (distant from
than ventral pores. Trilocular pores of pe- the excretory opening) part of such duct is
culiar structure are known in some species enlarged in the form of a bottle.
and genera, for example, in Cucullococcus 2) Collar tubular ducts. In these glands
arrabidensis (Neves, 1954) (pores without the duct is surrounded by collars of differ-
fringe) and in the species of Iberococcus ent sizes and forms. The greatest diversity
(pores are strongly convex and located in a of collar ducts takes place in the subfam-
deep cup). ily Pseudococcinae, whereas among more
3) Quinquelocular (5-locular) pores primitive Phenacoccinae these ducts are
have one central loculus and 5 peripheral known in several genera only, for example
loculi; sometimes central loculus absent. in Heliococcus, Seyneria, Ehrhornia. There
These pores are present mainly in the species are at least the following main variants of
of the subfamily Phenacoccinae (in Pseu- collar ducts.
dococcinae they are known mainly in g/g a) Fungiform ducts. The collar of these
Metadenopus) and usually scattered on the ducts is very wide and similar to a cap of
ventral surface of body, especially around fungus. Most of species with fungiform
mouthparts and near spiracles. In some cas- ducts are members of g/g Pseudococcus
es (for example, in g/g Heterococcus and g/g and only some species with such ducts are
Metadenopus) 5-locular pores are numerous known in other genera, for example, in the
on both body sides and replace partly or to- genera Mirococcopsis and Humococcus.
tally reduced trilocular pores. b) Oral collar ducts, with narrow collar
4) Multilocular pores usually have one which surrounds the excretory opening and
central loculus and 8–10 peripheral loculi part or all tube of the duct. Such ducts are
and form transverse rows on abdominal most common and known in most genera of
sternites. More rarely multilocular pores the subfamily Pseudococcinae. The length
present on other surface of body and/or of the collar can be rather different, from
show a peculiar distribution, structure and very small to longer than the length of the
size. For example, in Mirococcus sphaeroides duct itself.
Danzig, 1975 some of multilocular pores c) Middle collar ducts, with collar lo-
have several central loculi; species of the cated in the middle part of the duct (for
genera Pelionella and Rosebankia show 2 example, in Volvicoccus volvifer Goux,
different types of multilocular pores in the 1945, Metadenopus caudatus (Borchsenius,
same female. In g/g Neotrionymus, multiloc- 1958), etc.).
ular pores replace partly or totally reduced d) Double collar ducts. Proximal part of
trilocular pores. such collar is always more sclerotized than
© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259
240 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS
Table 1. Wax glands in Pseudococcidae.
Discoidal wax gland s
Simple discoidal pore s
Nudicauda nigra
Trilocular pore s

Cucullococcus Iberococcus
Quinquelocular por es

Multilocular pore s

Pelionella
Cylindrical wax glands
Simple tubular ducts

Coccidohystrix

Fungiform ducts

Oral collar ducts

Middle collar ducts

Collar tubular ducts

Double collar ducts

Crateriform duct s

Heliococcus
Mitral duct s

Ehrhornia, Discococcus

Bitubular /tritubular
ducts
(in g/g Rhizoecus)
Peculiar types

Nudicauda Ferrisia
© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259
I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 241
the distal part (for example, in some Triony- along body margin, cerarii, which is one of
mus, Paraputo, etc.). unique apomorphic characters of Pseudo-
f) Crateriform ducts. The collar of such coccidae. Structure and number of cerarii
ducts bears one or several conical setae. are important taxonomic characters which
These are known in the genus Heliococcus are used in the classification and identifica-
only. tion at the species level and at higher taxa
g) Mitral ducts; resemble a headwear level. The number of cerarii usually ranges
(“mitre”) of orthodox religious superiors. from 1 to 18 pairs (first pair on head and
These ducts are known, for example, in the the last pair on last abdominal tergite) and
genera Ehrhornia, Discococcus, Paradisco- their exact number is usually stable inside
coccus. of the species. More rarely there are small
3) Bitubular and tritubular ducts are individual variations of this number. An
structures, consisted of 2 or 3 crossed ducts, extraordinary variability of the number is
with or without general collar; these are currently known in some species only. For
present in many (but not in all!) species of example, Dysmicoccus angustifrons (Hall,
g/g Rhizoecus. 1926) has from 2 to 10 pairs of cerarii, Atro-
4) Peculiar types of tubular ducts are coccus alhagii (Hall, 1926) – from 2 to 15
known in different genera. For example, in pairs. Evolutional and individual reduction
the species of the genus Nudicauda, and in in the number of cerarii always occurs in the
the monotypic genus Malekoccus, the duct following succession: thoracic and anterior
opening is encircled by a flat sclerotized abdominal cerarii become absent initially;
ring; in the species of pantropical genus Fer- followed by the cerarii on head, and at the
risia, the opening is encircled by a sclero- end the anal pair (С ) is reduced.
18
tized zone of the cuticle with several associ- In some mealybugs the normal cerarii
ated flagellate setae. are totally absent or in the place of the last
In addition to these rather large ducts cerarian pair there are one or several flag-
which are well visible under usual trans- ellate setae, accompanied by group of tri-
lucent microscope, many mealybugs have locular pores. We consider such structures
numerous poorly visible minute microtubu- as pseudocerarii. Numerous pseudocerarii,
lar ducts, which can be located on different distributed along all body margin are a very
parts of the body and sometimes on the hind rare character which is known in monotypic
coxae. Fine structure and function of these Madagasia and Telocorys and in Dysmicoc-
microducts are unknown till now. cus trispinosus (Hall, 1923), which is prob-
Conical setae have different forms and ably must be transferred to Telocorys.
sizes (Fig. 2) and produce consistent wax. In g/g Puto and g/g Paraputo, the mar-
Especially large and numerous conical setae ginal cerarii are more numerous, up to 26
are known in the species of g/g Puto, g/g pairs. Moreover, additional (non-marginal)
Coccidohystrix and g/g Pedronia. In con- cerarii can present in the in medial zone of
trast, in some desert mealybugs, conical se- dorsum. In the monotypic genera Trimero-
tae are totally lost and the body is covered coccus and Eastia and in many species of g/g
by flagellate and/or hair-like setae only. Paraputo the marginal cerarii are fused in a
In most of mealybugs the conical setae solid marginal band.
have pointed or slightly roundish apex and In some species of Coccidohystrix, normal
only in the genera Rastrococcus and Stemma- cerarii are replaced by groups of large coni-
tomerinx the conical setae are sharply trun- cal setae on cuticular tubercles, without tri-
cated (see Fig. 2). The conical setae with locular or 5-locular pores – mamelons.
associated discoidal glands (usually with Flagellate and hair-like setae of differ-
trilocular pores, more rarely with 5-locu- ent size are more or less numerous on ven-
lar pores) form multiple binate structures tral body side of all species. Dorsal flagellate
© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259
242 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS
and/or hair-like setae are present mainly Fig. 3. The compound structure of the anal
in the species of subfam. Pseudococcinae. apparatus is connected with the mode of
Amongst Phenacoccinae these setae (usual- mealybug nourishment. Constant sucking of
ly together with conical setae) are present, sweet sap from the plant floem leads to prob-
for example, in Palaearctic genera Fonscol- lems with eliminating the surplus of water
ombia and Perystrix. and sugar (honeydew) from the organism.
Ostioles represented by slit-like open- The honeydew would easily cover the anal
ings, located symmetrically as one or 2 pairs segment and occlude the anus if lacking spe-
on dorsal (rarely on margin of ventral) sur- cial morphological structures that envelop
face of body. These structures are one of the honeydew drop with wax in order to
the unique synapomorphic characters of discard it from the body. Secondary simpli-
mealybugs (including “Putoidae” of some fication of the anal apparatus can be con-
authors) and the monotypic family Phen- nected with the feeding of sap directly from
acoleachiidae. The position of the ostioles the plant cells or by obligate symbiosis with
is constant; anterior pair always located on ants which feed on the excreted honeydew.
anterior part of cephalothorax and the pos- Legs usually small in comparison with
terior pair – on VI or VII abdominal tergites body; in the species of g/g Antonina legs
or between them. Margins (lips) of ostioles significantly reduced or totally absent.
usually slightly raised above surrounded Tarsus 1-segmented, with 2 thin setae at
cuticle and bearing the same cuticular the apex, so-called “tarsal digitules”. Claw
structures, i.e., different setae and pores. In with or without a denticle; this is impor-
the Oriental mealybugs of g/g Allomyrmo- tant taxonomic character that is tradition-
coccus, the ostioles are highly convex and ally used for the separation of two subfami-
strongly sclerotized. In very rare cases, the lies of Pseudococcidae. Claw digitules are
ostioles are secondary absent though they usually slightly longer than claw and have
are known in related species and genera or a clavate apex; very rarely they are short
even in the larvae of the same species. and/or pointed. In the species of g/g Allo-
Anal apparatus of a compound struc- myrmococcus claw digitules are widely ex-
ture: flat sclerotized ring around anus, one panded. All legs more or less equal in size,
inner row of round or oval pores, one outer but in Mongolian endemic, Mirococcus fos-
row of spinulae and 6 setae, which are usu- sor Danzig, 1983, anterior legs of burrowing
ally slightly longer than the diameter of anal type: femur, tibia and tarsus strongly en-
ring. Such most common type of anal appa- larged and claw elongated, with 2 denticles;
ratus we name as complete apparatus (Fig. claws of other legs of this species are also
3). In some species the number of spinulae elongated, but without denticles. Claws
is more numerous – 2–4 rows; such type of with 2 denticles are also known in some
anal apparatus we consider as complicated. other mealybugs, for example, in Perystrix
On the contrary, in some species, especially ulmaria Gavrilov, 2004; the second denticle
in deserts, anal apparatus is strongly sim- is always smaller than the first one. Unusu-
plified, with reduced number of pores and ally enlarged legs are known in the species
spinulae or even with their total absence. of Australian Pseudoripesia and South Af-
Additionally, many species have more or less rican Diversicrus. Hind coxae (more rarely
developed anal tube, invaginated inside of hind femurs and tibiae) of many species,
anal segment of body. In most species of g/g mainly from the subfamily Pseudococcinae,
Rhizoecus, pores and spinulae of anal appa- bearing so-called “translucent pores” or,
ratus are enlarged and irregular-shaped. In more rarely, very small microtubular ducts.
the genera Rhodania and Pararhodania, anal In the species of g/g Antonina numerous
ring strongly enlarged and sclerotized. Dif- and closely spaced translucent pores or mi-
ferent types of anal apparatus are shown on crotubular ducts form two compact groups
© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259
I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 243
Conical setae Flagellate setae Hair-like setae
Cerarius with multiple
Usual cerarius Pseudocerarius Mamelon
setae on sclerotized plate
Fig. 2. Setae, cerarii and cerarius-like structures.
on metathorax and anterior abdominal ster- cies in nature is higher. Some species-rich,
nites instead of lacked hind legs. but morphologically heterogenous genera
Ovoviviparity. We (Gavrilov-Zimin & such as Phenacoccus, Pseudococcus, Triony-
Danzig, 2012; Danzig & Gavrilov-Zimin, mus include oviparous and obligate ovovi-
2014) consider ovoviviparity as a plesio- viparous species and also species with facul-
morphic character of all Pseudococcidae. tative ovoviviparity.
This character was probably inherited by
mealybugs from the obligate ovoviviparous Family Pseudococcidae Cockerell, 1905
ancestral family Phenacoleachiidae and is
known in all primitive groups of mealybugs: Putoidae Tang, 1992: 382 (as a tribe of Pseudo-
coccidae with several genera). Williams et
g/g Puto, species-rich genera Heliococcus,
al., 2011: 11 (as a separate monotypic family
Coccura, Fonscolombia, Paraputo, Formi-
without Ceroputo). Gavrilov-Zimin & Dan-
cococcus and in many other genera, for ex-
zig, 2012: 108 (synonymy).
ample in legless mealybugs (g/g Antonina),
Rhizoecidae Williams, 1969 (as a tribe of Pseu-
species-rich genus Mirococcopsis, in numer- dococcidae). Hodgson, 2012: 8 (as a separate
ous small and monotypic genera: Nudicau- family). Gavrilov-Zimin & Danzig, 2014: 124
da, Adelosoma, Antoninella, Cucullococcus, (synonymy).
Lacombia, Ritsemia, Ehrhornia, Paratriony- Pennygullaniidae Koteja et Azar, 2008: 158.
Gavrilov-Zimin & Danzig, 2012: 108 (syn-
mus, Lankacoccus, Stricklandina, Madacan-
onymy).
thococcus, Mauricoccus, Mascarenococcus,
Cirnecoccus, Pleistocerarius, etc. To date Two subfamilies of the Pseudococcidae
more than 500 obligate ovoviviparous spe- generally differ in the following characters.
cies of mealybugs from more than 60 genera Species of the subfamily Phenacoccinae
have been found (Trapeznikova & Gavrilov, usually possess claw denticle, dorsal coni-
2008; I. Gavrilov-Zimin, unpublished), that cal setae, 9-segmented antennae, 5-locular
is about 25% of the species diversity in the pores and only simple tubular ducts. Species
global fauna. Moreover, there is no doubt of the subfamily Pseudococcinae without
that the real number of ovoviviparous spe- claw denticles, with flagellate dorsal setae,
© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259
244 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS
6–8-segmented antennae and tubular ducts ability row of general taxonomic characters
of different types (not only simple); 5-locu- with vast g/g Phenacoccus.
lar pores absent. However, there are numer- Nominal genera (6): Eastia De Lotto,
ous exclusions from this general definition. 1964 (Afrotropical), Mombasinia De Lotto,
Moreover, some genera and groups of gen- 1964 (Afrotropical), Nairobia De Lotto,
era (for example, g/g Metadenopus) have a 1964 (Afrotropical), Puto Signoret, 1876
questionable phylogenetic and taxonomic (mainly Holarctic), Rastrococcus Ferris,
position – see the scheme of estimated re- 1954 (Oriental, Australasian), Trimerococ-
lationships from Danzig & Gavrilov-Zimin, cus Balachowsky, 1952 (Palaearctic).
2014 with minor changes and additions Deviating genera/species. Several spe-
(Fig. 4). cies of Puto (Palaearctic P. borealis Borch-
senius, 1948 and Neotropical P. paramoensis
System of the morphological Matile-Ferrero, 1985 + P. lamottei Matile-
generic groups (g/g) Ferrero, 1985) and monotypic genera Mom-
basinia and Nairobia lacked claw denticle.
1 g/g Puto Signoret, 1876 Moreover, Mombasinia lacked ostioles and
demonstrates very peculiar large dorsal
Combination of diagnostic characters
groups of simple tubular ducts, enclosed by
(plesiomorphic+apomorphic): claw with a
multilocular pores.
denticle; most or all cerarii with multiple
All species of Rastrococcus have peculiar
(more than 4) conical setae or all cerarii
truncate conical setae.
merged in a solid marginal band; 9-seg-
mented antennae; both pairs of ostioles well
2 g/g Phenacoccus Cockerell, 1893
developed; complete anal apparatus (with
one inner row of pores, one outer row of Combination of diagnostic characters
spinulae and 6 long setae); tubular ducts of (plesiomorphic + apomorphic): cerarii
simple type; trilocular pores numerous and each with 2–3 conical setae, never merged
evenly scattered on all body surface. in marginal band; 9-segmented antennae;
Additional characters: marginal cerarii quinquelocular pores present; both pairs of
are usually supernumerary (more than 18 ostioles present.
pairs) in comparison with other Phenacoc- Additional characters (plesiomorphic):
cinae or even merged in the solid marginal well developed dorsal conical setae (often
band of conical setae and trilocular pores organized in several medial cerarii), claw
(as in Trimerococcus and in South African with a denticle; simple type of tubular ducts;
endemic Eastia). well-developed legs and antennae, complete
All species of the group are highly mov- anal apparatus; trilocular pores numerous
able insects possessing well developed an- and evenly scattered on all body surface.
tennae and legs and living openly on their Nominal genera (10): Coccura Šulc, 1908
host plants. Probably all species are ovovi- (Palaearctic), Fonscolombia Lichtenstein,
viparous and do not construct wax ovisacs. 1877 (Palaearctic), Malekoccus Matile-
There is no doubt that the group or at least Ferrero, 1988 (Palaearctic), Mammicoccus
the genus Puto itself are the most ancient Balachowsky, 1959 (Neotropical), Octococ-
and primitive mealybugs and neococcids cus Hall, 1939 (Afrotropical), Oxyacanthus
(superfamily Coccoidea) as a whole. The De Lotto, 1970 (Afrotropical), Perystrix
species of g/g Puto demonstrate all apomor- Gavrilov, 2004 (Palaearctic), Phenacoccus
phic characters of the family Pseudococ- Cockerell, 1893 (global distribution), Stem-
cidae, plesiomorphic characters inherited matomerinx Ferris, 1950 (Nearctic), Syna-
from estimated ancestral family Phenaco- canthococcus Morrison, 1920 (Palaearctic
leachiidae and almost uninterrupted vari- and Oriental).
© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259
I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 245
Usual complete anal apparatus Complicated anal apparatus
Different examples of simplified anal apparatus
Mirococcopsis ehrhornioidea
Ritsemia pupifera
Mirococcopsis kalaginae
Cucullococcus arrabidensis
Paratrionymus halocharis Ehrhornia fodiens
Some peculiar types of anal apparatus
Ripersiella brevipes Pararhodania armena
Rhodania flava
Fig. 3. Different types of anal apparatus.
© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259