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A new African soft scale genus, Pseudocribrolecanium
gen. nov. (Hemiptera: Coccoidea: Coccidae), erected
for two species, including the citrus pest P. andersoni
(Newstead) comb. nov.
Takumasa Kondo1
1Department of Entomology, University of California, 1 Shields Avenue, Davis, California 95616-8584,
U.S.A.
Abstract
A new African genus of soft scale insects, Pseudocribrolecanium gen. nov. is erected to accommodate
Akermes colae Green & Laing and Cribrolecanium andersoni (Newstead). The adult females and
first-instar nymphs of the two species are redescribed and illustrated. Taxonomic keys to separate the adult
females and first-instar nymphs are provided. The affinity of Pseudocribrolecanium with the tribe
Paralecaniini in the subfamily Coccinae is discussed.
Résumé
Un nouveau genre africain de cochenille, Pseudocribrolecanium gen. nov. est créé pour les espèces
Akermes colae Green & Laing et Cribrolecanium andersoni (Newstead). Les femelles adultes et les larves
du premier stade des deux espèces sont redécrites et illustrées. Une clé dichotomique est proposée pour les
femelles adultes ainsi qu'une clé pour les larves. L'affinité du genre Pseudocribrolecanium avec la tribu des
Paralecaniini, dans la sous-famille des Coccinae, est discutée.
Correspondence: tkondo@ucdavis.edu
Received: 6.9.2005 | Accepted: 20.10.2005 | Published: 3.31.2006
Keywords: coccids, new genus, taxonomic keys
Creative Commons Attribution 2.5 http://creativecommons.org/licenses/by/2.5/
ISSN: 1536-2442 | Volume 6, Number 1
Cite this paper as:
Kondo T. 2006. A new African soft scale genus, Pseudocribrolecanium gen. nov. (Hemiptera: Coccoidea: Coccidae),
erected for two species, including the citrus pest P. andersoni (Newstead) comb. nov. 16pp. Journal of Insect Science
6:01, available online: insectscience.org/6.01
Journal of Insect Science | www.insectscience.org ISSN: 1536-2442
Journal of Insect Science: Vol. 6 Article 1 1
Introduction
There are about 270 species in 56 genera of soft
scale insects (Hemiptera: Coccoidea: Coccidae)
hitherto described from the African continent
(Ben-Dov et al. 2005). These have been described
mainly during two periods: (i) from 1899–1941,
during which more than one hundred species were
named by such early workers as Newstead, Hall,
Green, and Laing, and (ii) from 1954–1979 when
about another hundred species were added mainly
by De Lotto and Hodgson (Ben-Dov et al. 2005).
Since then, various authors, including Hodgson,
have sporadically contributed to the knowledge of
the coccid fauna of Africa. Despite its rich flora,
which includes 6 biodiversity hotspots (Myers et al.
2000; Myers 2001), the number of described coccid
species recorded in the African continent has not
increased in the last 10 years, undoubtedly because
of the lack of taxonomic specialists in the region.
The author had the opportunity to travel to Ghana
in June, 2005, and collected scale insects of various
families including the Coccidae, Diaspididae,
Margarodidae, Ortheziidae, Pseudococcidae and
Stictococcidae. Among the coccids, a scale insect
later identified as Akermes colae was collected on
the leaf of a cacao tree. The genus Akermes
Cockerell (1902) currently includes 14 species
distributed in Central and South America (10
species), India (1 species), Australia (2 species), and
Africa (1 species). The author has studied the genus
Akermes previously and considers that it is
endemic to the New World (Kondo & Williams
2004). Attempts were therefore made to allocate A.
colae to an appropriate genus. When Green & Laing
(1924) described Akermes colae, they pointed out
that A. colae was structurally very similar to A.
andersoni (referred to as Lecanium andersoni). De
Lotto (1968) redescribed A. bruneri Cockerell, the
type species of Akermes, and stated that A.
andersoni had little in common with the genus
Akermes and transferred it to Cribrolecanium
Green (1921). However, De Lotto (1968) did not
study A. colae, and it has remained in the genus
Akermes until the present.
Cribrolecanium currently contains 3 species, 2
from the Oriental region and C. andersoni from the
Afrotropical region (Ben-Dov et al. 2005) although
the latter species has been considered only
doubtfully congeneric (Hodgson 1994) based on
adult female morphology. However, studies based
on crawler morphology suggest that C. andersoni is
definitely not congeneric (Kondo and Williams
2002; Williams and Kondo 2002). As indicated by
Green & Laing (1924), C. andersoni and A. colae
are closely related, but do not fit into any known
genus, and thus a new genus
Pseudocribrolecanium is erected here to
accommodate them.
Materials and Methods
Specimens were slide mounted according to
methods discussed by Kosztarab (1996) and Kondo
& Gullan (2005). Morphological terminology
follows mostly that of Hodgson (1994).
Measurements of specimens were made using an
ocular micrometer on an Olympus phase-contrast
microscope. The illustrations of the coccids follow
the style adopted for the Coccoidea, with the dorsal
surface drawn on the left side of the drawing and
the ventral surface drawn on the right, with
enlargements of important features placed around
the illustration. The total number of specimens
used for each description is given in parentheses,
e.g. (n=48). In the material studied, the number of
slides and specimens on each slide are given as the
number of slides followed by the total number of
specimens and their corresponding stages; for
example, 1 slide with 1 second-instar female and 1
adult male specimen would be: 1(2: 1 second-instar
female + 1 adult male). The stage is not given if all
specimens are adult females.
Depositories
AUCC: Auburn University Coccoidea Collection,
Alabama, U.S.A.
BMNH: the Natural History Museum, London,
U.K.
Results
Taxonomy
Pseudocribrolecanium gen. nov. keys out to the
subfamily Coccinae, tribe Coccini, in the keys to
subfamilies, tribes and type species of genera of
Coccidae provided by Hodgson (1994). However, it
does not fit well into any known genus.
Subfamily Coccinae, Tribe Coccini
Pseudocribrolecanium Kondo, new genus
Type species
Akermes andersoni Newstead, 1917, by present
designation.
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Generic diagnosis, adult female
Insects oval, pyriform to asymmetrical in shape,
flat, becoming sclerotized at maturity.
Dorsum. Derm membranous, becoming highly
sclerotized at maturity. Dorsal setae short, sharply
spinose. Simple pores present or absent. Dorsal
microducts present, rather few, scattered
throughout dorsum. Translucent furrows present,
associated with stigmatic areas and often
elsewhere; furrows either touching or not touching
margin. Dorsum usually with several cribriform
plates, although occasionally absent on P. colae.
Dorsal tubercles and tubular ducts absent.
Preopercular pores usually present in a longitudinal
line anterior to anal plates. Anal plates together
quadrate or inverted pyriform, dorsal surface with
3 or 4 apical or subapical setae, plus 1 seta on inner
margin of each plate. Anal ring with 4 pairs of setae
and an irregular row of translucent wax pores. Eyes
present, represented by clear areas located on
dorsal submargin.
Margin. Derm heavily sclerotized in a narrow
band around margins. Marginal setae numerous,
slender, flagellate, arranged in a single row.
Stigmatic clefts shallow or absent, with 3, rarely
with fewer or up to 5 stigmatic setae; with either
one seta longer than others or all subequal in
length.
Venter. Ventral derm membranous. Ventral
microducts present, small, scattered evenly on
venter. Mouthparts well-developed, often displaced
to one side; labium 1 segmented, with 4 pairs of
setae. Pregenital disc-pores each with 5–11 loculi,
mostly 10-locular; scarce, present around vulvar
area and sparsely distributed on abdominal
segments. Ventral tubular ducts absent. Spiracular
disc-pores with 3–5 loculi, present in a single line
extending laterally from each spiracle to body
margin. Antennae each 2–5 segmented,
segmentation indistinct; fleshy setae present only
on last antennal segment and setae often knobbed.
Legs greatly reduced, segmentation poorly defined;
claws without a denticle. Spiracles rather small.
First-instar nymph.
Body elongate oval.
Dorsum. Dorsal derm membranous. Dorsal setae
short, present in 2 longitudinal parallel rows of 5
setae. A trilocular pore present on each side of head
region near margin. Dorsal microducts present in a
submarginal and 2 submedian longitudinal rows,
with a few additional microducts on thorax between
the submedian and submarginal rows. Simple pores
each closely associated with a microduct. Anal
plates each triangular, dorsal surface with a
shingled texture and 3 apical setae, 1 seta along
inner margin of each plate, with 1 fringe seta and 1
ventral subapical seta. Anal ring with 3 pairs of
setae and an irregular row of translucent wax pores.
Eyes located on margin on area laterad to each
antennal scape.
Margin. Outline smooth. Marginal setae slender,
total 34, with 8 anteriorly between eyes and, on
each side, 3 between each eye and anterior
stigmatic setae, 2 between each anterior and
posterior stigmatic setae and 8 between each
posterior stigmatic setae and posterior end.
Stigmatic setae in groups of 3, well differentiated
from marginal setae; median seta bluntly spinose,
or clavate; lateral setae short, bulbous, length about
1/5–1/11 of median seta.
Venter. Ventral derm membranous. Pairs of
submedian abdominal setae present on last 3
abdominal segments. Submarginal setae arranged
in a parallel row of 7 on each side of abdomen, plus
1 submarginal seta between each anterior and
posterior spiracle, and 1 submarginal pair of setae
near apex of head. Interantennal setae 1 pair. With
1 ventral microduct present between each pair of
submarginal setae on abdomen and 1 between each
anterior and posterior spiracle. Spiracular
disc-pores each with 3 loculi; 3 or 4 pores between
each anterior and posterior spiracle and margin.
Mouthparts normal; labium with 4 pairs of labial
setae. Legs well-developed; microctenidia present
on tibial apex. Prothoracic tarsal digitules
dissimilar, one knobbed and other spiniform;
mesothoracic and metathoracic tarsal digitules
similar, both knobbed. Spiracles normal, with a
well developed muscle plate. Claws each with a
denticle; claw digitules knobbed, one slightly
broader than other. Antennae each 6-segmented,
with 3rd antennal segment longest; fleshy setae
present on last 3 apical segments; 1 single fleshy
seta on segment 4, 1 fleshy seta plus 1 setose seta on
segment 5, and several fleshy and setose setae on
terminal segment.
Etymology
The genus name is derived from the prefix Pseudo-
(Greek), meaning something superficially
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resembling the original subject, and the genus
name Cribrolecanium Green.
Key to species of Pseudocribrolecanium, gen. nov.
based on adult females
1. Dorsum with both large and small cribriform
plates, smaller plates near margin each 10–80 µm
wide, larger plates farthest from margin each
100–180 µm wide; present in about 2 or 3 irregular
rows totaling 12–22 around body
P. andersoni (Newstead), comb. nov.
–Dorsum with or without cribriform plates, when
present, each plate small and subequal in size,
15–50 µm wide; present in 1 irregular row totaling
7–9 plates around body
P. colae (Green & Laing), comb. nov.
Key to species of Pseudocribrolecanium, gen. nov.
based on first-instar nymphs
1. Median stigmatic seta of each cleft bluntly
spinose, apex round, not swollen; head rounded
P. andersoni (Newstead), comb. nov.
–Median stigmatic seta of each cleft clavate, with a
swollen apex; body narrowing at head
P. colae (Green & Laing), comb. nov.
Pseudocribrolecanium andersoni
(Newstead), comb. nov.
Figures 1, 2
The White Powdery Scale
Akermes andersoni Newstead, 1917: 347; Almeida
1969: 135; De Lotto, 1965: 178; Hall 1937: 122
Lecanium andersoni Newstead; Green & Laing
1924: 419
Cribrolecanium andersoni (Newstead); De Lotto
1968: 83; Brink & Bruwer 1989: 9; Hodgson 1969:
10
Parakermes andersoni (Newstead); Fonseca, 1973:
247
Material studied
Lectotype. In order to preserve stability and the
nomenclatural status of this species, a lectotype is
here designated from the syntypes of this species as
follows: “Lecanium andersoni”, Kenya (as British
East Africa), 1(1): collecting date not given
(according to the Type description, the date should
be i.1914), coll. T.J. Anderson, ex orange, No. 18,
14/252, labeled cotype, B.M. 1945, 121 (BMNH).
Paralectotypes. Kenya (as B.E. Africa), 1(5):
Kabete, i.1914, ex orange, coll. T.J. Anderson, B.M.
1945, 121, (BMNH); 1(3): collecting date not given,
slide mounted from Newstead’s collection (the
collector T.J. Anderson does not appear on the
label), ex citrus, labeled part of type material, B.M.
1940, No. 180 (BMNH); 1(4): Limoru, i.1914, coll.
T.J. Anderson, ex orange, No. 14/252, No. 18,
labeled cotype, Reg. 1916.30 (BMNH); 1(5: 2 adult
ᥜᥜ + 1 third-instar nymph + 2 Diaspididae):
Limoru, i.1914, coll. T.J. Anderson, ex orange, No.
14/252, No. 18, labeled cotype, 1916-30 (BMNH);
1(3 first-instar nymphs): Limoru, i.1914, coll. T.J.
Anderson, ex orange, No. 14/252, No. 18, labeled
“Type larvae”, 1916–30 (BMNH).
Non-type material. Ghana, 1(1): Tofo, 21.v.1976,
coll. not given, ex cacao, COPR, C.I.E.A. 9866
(BMNH); Kenya, 1(2): Westlands, 23.viii.1973, coll.
not given, ex Ficus , No. 2108, G.W. 0100, C.I.E.A.
6593 (BMNH); 1(1): Juja, 23.ix.1978, coll. T.C.
Griesbach, 2175, C.I.E.A.10823 (BMNH); 1(2):
Nairobi, 8.i.1982, coll. Alice, ex passionfruit, No.
60/83 TC 2219, CIEA 13866 (BMNH); Tanzania (as
German East Africa), 19(19): 2.iii.1918, coll.
Lamborn, S.W.R., host not given, AL-133-99, slide
mounted from dry material (BMNH); 11(11:
first-instar nymphs): 2.iii.1918, coll. S.W.R.
Lamborn, det. Newstead, host not given,
slide-mounted from BMNH dry material,
AL-133-99c–m (AUCC); Uganda, 1(2: 1 adult ᥜ+1
adult ᥞ): Kampala, 25.viii.1919, coll. C.C. Gowdey,
ex Citrus nobilis, B.M. Reg. No. 1919.253 (BMNH).
Zambia (as Northern Rhodesia), 1(2): Nkana,
6.vi.1948, coll. J. Fourfooz, det. W.J. Hall, host not
given, I.I.E. 1521/1969, BM1958-758 (BMNH).
Zimbabwe (Southern Rhodesia), 2(2): Mazoe,
Umba, 24.iii.1935, coll. M.C. Mossop, ex grape fruit
leaves, B.M. 1958-229, No. 822 (BMNH); 1(1):
Mazoe, Umba, 26.v.1935, coll. Mossop per W.J.
Hall, via Green, det. F. Laing, ex grape fruit leaves,
No. 822 (BMNH); 1(1): Harare (as Salisbury),
27.ix.1962, coll. not given, ex citrus, det. C.J.
Hodgson, (815), Acc. No. 9877, BM1967-758
(BMNH). Country not given (probably Zimbabwe),
1(3): Imbija, 10.xi.1938, Pres. Dr. W.J. Hall, ex
grapefruit, B.M. 1958.229 (BMNH).
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Figure 1. Pseudocribrolecanium andersoni (Newstead), adult female. A, cribriform plate; B, enlargement of dorsal
derm; C, enlargement of sclerotized marginal band with marginal and stigmatic setae; D, marginal setae; E, dorsal
setae; F, dorsal microduct; G, simple pore; H, anal plates; I, pregenital disc-pores; J, anal ring (right half); K, reduced
legs; L, ventral setae; M, ventral microduct; N, spiracular disc-pores; O, antenna.
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Figure 2. Pseudocribrolecanium andersoni (Newstead), first-instar nymph. A, trilocular pore; B, dorsal seta; C,
marginal seta; D, stigmatic setae; E, simple pore; F, dorsal microduct; G, anal ring (right half); H, ventral seta; I, ventral
microduct; J, claw; K, spiracular disc-pore; L, ventral submarginal cephalic seta.
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Description
Adult female (Fig. 1)
Unmounted material. Insects completely
covered with rather dense, dusky-white, mealy
secretion. Color on removal of secretion, rich dark
piceous or very dark castaneous, shining; younger
specimens varying from reddish brown to dusky
buff. Form irregularly oval, asymmetrical, more or
less narrowed in front; sometimes broadly ovate or
subcircular. Flat or very low convex, with faint
median keel in abdominal region; sides well within
margin. Derm densely sclerotized, especially
towards margins; with innumerable minute,
translucent poreless cells (Newstead 1917).
Mounted material. Body outline oval, pyriform
to slightly asymmetrical; body 2.1–3.8 mm long,
1.4–3.2 mm wide (n=48).
Dorsum. Derm often with areolations, particularly
expressed submarginally (Fig. 1B). Dorsal setae
(Fig. 1E) sharply spinose or conical, each 5–9 µm
long. Simple pores (Fig. 1G) about 1.8 µm wide,
scattered on dorsum. Dorsal microducts (Fig. 1F)
each about 1.8 µm wide, scattered on dorsum.
Translucent stigmatic furrows present as in
diagnosis, usually with an additional translucent
lateral furrow on head and two on abdominal
region. Cribriform plates (Fig. 1A) oval to elongate
oval, occurring either submarginally or
submarginally and submedially, totaling 12–22;
varying in size, plates closer to margin smallest,
each about 10–80 µm wide, innermost plates
largest, each 100–180 µm wide. Preopercular pores
present in a linear group of about 10–40 pores
anterior to anal plates, often displaced anteriorly
onto the thoracic and head region with pores
completely absent just anterior to anal plates (as
reported by Almeida 1969), or form a long line of
pores extending from area just anterior to anal
plates up to head region. A sclerotic area often
present around anal plates. Anal plates (Fig. 1H)
together inverted pyriform or quadrate, with round
angles, located at about 1/6 of body length from
posterior margin, each plate 117–121 µm long,
53–58 µm wide; anterolateral margin 62–70 µm
long, posterolateral margin 98–107 µm long, with 3
or 4 setae on dorsal surface, plus 2 pairs of fringe
setae and about 2 ventral subapical setae. Anal ring
(Fig. 1J) as in diagnosis. Eyes present, represented
by clear subcircular areas located on dorsal
submargin (see arrow on Fig. 1).
Margin. A strongly sclerotized narrow band
present around margins (Fig. 1C) in older
specimens. Marginal setae (Fig. 1D) flagellate, each
46–115 µm long, arranged in a single row,
numerous, with 24–28 on each side between
anterior and posterior stigmatic areas. Stigmatic
clefts shallow, with 3, rarely less or up to 5
stigmatic setae (Fig. 1C), each seta bluntly spinose;
5–10 µm long, often with one seta much longer
than others, 10–15 long.
Venter. Ventral setae (Fig. 1L) slender, straight or
slightly bent, each 7–16 µm long. Ventral
microducts (Fig. 1M) scattered evenly on venter,
small, each about 1.5 µm wide. Mouthparts
centered or less often displaced to one side;
clypeolabral shield 145–156 µm wide. Pregenital
disc-pores (Fig. 1I) each 3.6–4.4 µm wide, with
6–11 loculi, mostly 10-locular, present around
vulvar area and anteriorly across all abdominal
segments, becoming gradually scarce on anterior
segments. Spiracular disc-pores (Fig. 1N) each 3–4
µm wide, with 3–5 loculi, in a single line extending
laterally from each spiracle to body margin.
Antennae (Fig. 1O) rather short, each 111–145 µm
long, segmentation indistinct, with 2–4 discernible
segments; with fleshy setae present on last segment
only and often apically knobbed. With 2 pairs of
interantennal setae. Legs (Fig. 1K) each 23–44 µm
long (total length); tarsal digitules not detected;
claw digitules, slender, setose. Anterior spiracular
peritremes each 31–34 µm wide, posterior
peritremes each 37–41 µm wide.
Morphological variation. Two specimens from
Kenya labeled as A. andersoni collected on Ficus
sp. are strongly asymmetrical and resemble P.
colae. Their largest cribriform plates are rather
small, each 40–80 µm wide, smallest cribriform
plates each 10–40 µm wide. The cribriform plates
in these species are arranged mostly in a single
marginal row although there are places with
definitely an inner and outer row as in the rest of
the material of P. andersoni. The total number of
cribriform plates in the Kenyan specimens are 14
and 16 respectively and fall also in the range of P.
andersoni. Whether the specimens collected on
Ficus sp. should belong in a different species needs
further study. Measurements of the above
specimens were excluded from the description.
First-instar nymph (Fig. 2)
Unmounted material. Not seen.
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Mounted material. Body outline elongate oval,
anterior end rounded; body 431–485 µm long,
189–226 µm wide (n=14).
Dorsum. Dorsal setae (Fig. 2B) short, each 4–5
µm long, distributed as in diagnosis. A trilocular
pore (Fig. 2A) present on each side of head region
near margin. Dorsal microducts (Fig. 2F) each
2.7–3.6 µm wide, distributed as in diagnosis.
Simple pores (Fig. 2E) each about 1.8 µm wide and
closely associated with a microduct. Anal plates
each 48–50 µm long, 22–24 µm wide; anal plate
setae and anal ring (Fig. 2G) as in diagnosis.
Margin. Marginal setae (Fig. 2C) slender, each
13–33 µm long, those on anterior and posterior
ends longest; number and distribution as in
diagnosis. Each stigmatic area with a group of 3
stigmatic setae (Fig. 2D), well differentiated from
marginal setae; each median seta bluntly spinose,
15–17 µm long; lateral setae bulbous, each 1.8–2.7
µm long.
Venter. Submedian abdominal setae in pairs on
posterior 3 segments, each seta 13–22 µm long,
those on last abdominal segments longest.
Submarginal setae (Fig. 2H & L) distributed as in
diagnosis. Ventral microducts (Fig. 2I) each about
1.5 µm wide, distributed as in diagnosis. Spiracular
disc-pores (Fig. 2K) each 2–3 µm wide, with 3
loculi; with 3 or 4 pores present in a line from each
spiracle to margin. Clypeolabral shield 67–71 µm
wide. Trochanter + femur of each leg: 68–74 µm
long, tibia + tarsus 65–68 µm long, claw (Fig. 2J)
13–15 µm long. Antennae 6-segmented, each
130–138 µm long.
Biology (from Brink 1998)
The white powdery scale is a polyphagous insect
known as a pest of citrus. The adult female is
completely covered with a dense white powdery
secretion, which also spreads over the surrounding
parts of the host plant giving infested leaves an
almost uniform white powdery appearance. P.
andersoni feeds on leaves and young shoots, but
does not feed on the fruit and branches. The
honeydew serves as a substrate for the saprophytic
sooty mold fungi of the genus Capnodium. In South
Africa, the insect is found throughout the year,
where it appears to have three field generations per
year. Various natural enemies of P. andersoni are
known, including the hymenopterous parasitoids
Coccophagus philippiae (Silvestri), C. pulvinariae
Compere, C. lycimnia (Walker), Coccophagus sp.
(Aphelinidae), Metaphycus spp., Neastymachus
dispar Prinsloo (Encyrtidae), Tetrastichus sp.
(Eulophidae), and chrysopid larvae (Neuroptera).
P. andersoni was considered parthenogenetic by
Brink (1998) and Brink & Brewer (1989). However,
Almeida (1969) described the adult male and an
adult male was observed in the present study (see
Material studied).
Diagnostic features
The adult female of P. andersoni (Newstead) can be
easily separated from P. colae by the following
combination of characters: (i) presence of both
large and small cribriform plates on P. andersoni
(all small on P. colae ), (ii) difference in number of
cribriform plates, with 10–22 in P. andersoni, but
only 0–9 on P. colae, (iii) body shape, which is oval
to only slightly asymmetrical in P. andersoni but
highly asymmetrical in P. colae, and (iv) the
presence of obvious simple pores (not detected on
P. colae ). Furthermore, the first-instar nymphs of
P. andersoni have (v) bluntly spinose stigmatic seta
whilst they are clavate on P. colae, and (vi) the head
has a round contour whereas it is tapering in P.
colae.
Distribution
Afrotropical Region: Angola, Cameroon, Ghana,
Kenya, Mozambique, South Africa, Swaziland,
Uganda, Zambia, Zimbabwe, Mauritius (Brink
1998; De Lotto 1965; Grové 2001; Hall 1937;
Hodgson 1969; Matile-Ferrero & Nonveiller 1984).
Host plants
Anacardiaceae: Mangifera indica; Araliaceae:
Schefflera sp.; Boraginaceae: Ehretia silvatica;
Lauraceae: Persea americana; Loganiaceae:
Anthocleista grandiflora,Strychnos
madagascariensis; Meliaceae: Toona ciliata;
Moraceae: Ficus benjamina,F. elastica,F. sur,F.
verrucocarpa; Myrtaceae: Callistemon sp., Psidium
guajava; Passifloraceae: Passiflora edulis;
Rubiaceae: Psychotria zombamontana,Coffea
canephora; Rutaceae: Citrus paradisi,C.
reticulata,Citrus spp.; Sterculiaceae: Theobroma
cacao; Strelitziaceae: Strelitzia nicolai (Almeida
1969; Ben-Dov 1993; Brink 1992a, b, 1998; De
Lotto 1965; Grové 2001; Hall 1937; Hodgson 1969;
Matile-Ferrero & Nonveiller 1984).
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Pseudocribrolecanium colae (Green &
Laing), comb. nov
Figures 3, 4, 5
Lecanium (Akermes)colae Green & Laing, 1924:
419
Akermes colae Green & Laing, Ben-Dov 1993: 5
Material studied
Lectotype. In order to preserve the stability and
nomenclatural status of this species, a lectotype of
this species is here designated from the syntypes of
this species as follows: “Lecanium (Akermes)
colae”, Ghana, 1(1): Gold Coast, Aburi, 24.i.1922,
coll. W.H. Patterson, det. Laing, ex Cola
acuminata, No. 610 (BMNH).
Paralectotypes. GHANA, same data as lectotype,
mixed slide, 1(6: 1 hymenopteran larva + 4 adult ᥜᥜ
Parasaissetia nigra (Nietner) + 1 adult ᥜP. colae ):
paralectotype conspecific with lectotype clearly
indicated on label: longest of all specimens on slide
ca. 3.5 mm (at greatest length) (BMNH).
Other material studied. Sierra Leone, 4(8: 3
adult ᥜᥜ + 5 second-instar ᥞᥞ): Freetown, Sama,
25.xi.1924, coll. E. Hargreaves, ex. avocado pear,
No 1268 (BMNH). Ghana, 2(2: 1 adult ᥜ+1
first-instar nymph): Eastern Region, Bunso
Arboretum, 06°16’02” N, 00°27’44” W, 19.vi.2005,
coll. G.E. Morse & T. Kondo, ex Theobroma cacao,
underside leaf along vein (BMNH).
Note. The collection data of the lectotype and
paralectotype herein designated matches well the
data given in the original description by Green &
Laing (1924). However, the paralectotype slide is
composed of a mixture of specimens of which only
one specimen is conspecific with the lectotype. The
remaining specimens include four adult females of
the coccid Parasaissetia nigra (Nietner) and one
hymenopteran larva (probably a parasitoid). It is
common to find one or two, and even more species
together on one host plant. The mixing of species
on this one slide probably occurred because the
adult females of the two coccid species resemble
each other superficially in life, e.g., P. nigra and P.
colae both become heavily sclerotized at maturity
and their body shape is somewhat similar being
pointed at both extremities.
Description
Adult female (Figs 3 & 4)
Unmounted material. Morphology not
recorded.
Mounted material. Body outline asymmetrical,
head acuminate; body 2.0-2.2 mm long, 3.3-3.7
mm wide (n=4).
Dorsum. Derm with numerous areolations in a
broad submarginal band; area of areolation often
narrower or less pronounced on one side of body
(enlargement of dorsal derm shown in Figs 3A, 4E).
Areolated submarginal band often with lighter
subcircular patches of less sclerotized areas present
at regular intervals. Dorsal setae (Figs 3F, 4D)
spinose, short, each 2–3 µm long. Simple pores not
detected. Dorsal microducts (Figs 3D, 4F) small,
each about 2 µm long, inner ductule not detected,
scattered throughout dorsum. Cribriform plates
(Fig. 3F) present or absent (Fig. 4), if present,
subcircular, each 15–45 µm wide, located
submarginally; total 7–9: with 1–3 on head region,
1 between stigmatic areas, and 2 between posterior
stigmatic area and body apex. Translucent
stigmatic furrows present as in diagnosis, usually
with an additional submarginal lateral furrow on
head and 2 on abdominal region. Preopercular
pores (Figs 3G, 4G) each 4.5–8.0 µm wide, in 1 or
rarely 2 groups totaling 12–25 pores in a straight
longitudinal line 1 or 2 pores wide anterior to anal
plates; pore line extending up to area about half
way between anal plates and posterior spiracular
furrows; pores absent from area immediately
anterior to anal plates. Anal plates (Figs 3H, 4H)
together quadrate, each with round angles; located
about 1/7 of body length from posterior margin;
each plate 43–45 µm long, 25–26 µm wide;
anterolateral margin 32–33 µm long, posterolateral
margin 37–39 µm long; each with 4 setae on dorsal
surface, plus 1 pair of fringe setae and about 2
ventral subapical setae. Anal ring (Figs 3J, 4J) as in
diagnosis. Eyes present, represented by clear
subcircular areas on dorsal submargin.
Margin. Margins becoming heavily sclerotized in
a narrow band around entire body on older
specimens. Marginal setae (Figs 3C, 4C) flagellate,
each 62–72 µm long, arranged in a single row,
numerous, with 23–31 on each side between
anterior and posterior stigmatic areas. Stigmatic
clefts very shallow or absent; each with 3, rarely 4
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Journal of Insect Science: Vol. 6 Article 1 9
Figure 3. Pseudocribrolecanium colae (Green & Laing), adult female (drawn from type material). A, enlargement of
dorso-submarginal derm; B, stigmatic setae (top) and enlargement of sclerotized marginal band with marginal and
stigmatic setae (below); C, marginal seta; D, dorsal microduct; E, preopercular pores; F, dorsal setae; G, cribriform
plate; H, anal plate; I, pregenital disc-pores; J, anal ring (right half); K, ventral seta; L, reduced leg; M, ventral
microduct; N, spiracular disc-pores; O, antenna.
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Figure 4. Pseudocribrolecanium colae (Green & Laing), adult female (drawn from specimen collected in Ghana). A,
stigmatic setae; B, enlargement of sclerotized marginal band with marginal and stigmatic setae; C, marginal seta; D,
dorsal setae; E, enlargement of dorso-submarginal derm; F, dorsal microduct; G, preopercular pore; H, anal plate; I,
pregenital disc-pores; J, anal ring (right half), wax pores not illustrated; K, ventral seta; L, reduced leg; M, ventral
microduct; N, spiracular disc-pores; O, antenna.
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stigmatic setae (Figs 3B, 4A&B), often absent or
broken off in some stigmatic areas; each setae
sharply spinose, rarely bifurcate, 7.5–20 µm long;
median seta longest, or all subequal in length.
Venter. Ventral setae (Figs 3K, 4K) slender,
straight or slightly bent, each 8–18 µm long,
sparsely distributed submarginally, near
appendages, and on mid areas of venter. Ventral
microducts (Fig. 3M, 4M) sparse, scattered evenly
on venter, small, each about 1.5 µm wide.
Mouthparts displaced to one side; clypeolabral
shield 54–55 µm wide. Pregenital disc-pores (Fig.
3I, 4I) each 3.6–4.5 µm wide, with 5–10 (mostly 10)
loculi, present around vulvar region and sparsely
distributed on mid areas of abdomen. Spiracular
pores (Figs 3N, 4N) each 2.7–3.6 µm wide, with
3–5 loculi, present in an irregular line 1 or 2 pores
wide extending laterally from each spiracle to body
margin. Antennae (Figs 3O, 4O) small, each 38–44
µm long, segmentation indistinct, with about 3–5
discernible segments; setal distribution as in
diagnosis. With 2 pairs of interantennal setae. Legs
(Figs 3L, 4L) greatly reduced, each 8–16 µm long
(total length), all segments fused; claws rarely
visible; claw and tarsal digitules not discernible.
Spiracles rather small; anterior spiracular
peritremes each 25–27 µm wide, posterior
peritremes each 30–32 µm wide.
Morphological variation. A specimen recently
collected in Ghana (Fig. 4) has fewer translucent
furrows, no lighter patches around submargins and
no cribriform plates. The degree of sclerotization
appears to be highly variable in P. colae. Thus the
main difference between the specimen from Ghana
and those from the type series is the presence or
absence of cribriform plates. More material is
needed to determine if the specimen without
cribriform plates is part of the morphological
variation of A. colae or a separate closely-related
species.
First-instar nymph (Fig. 5)
Unmounted material. Morphology not
recorded.
Mounted material. Body outline elongate oval,
tapering towards head; body 575 µm long, 275 µm
wide (n=1).
Dorsum. Dorsal setae (Fig. 5B) short, each 4–5
µm long, distributed as in diagnosis. A trilocular
pore (Fig. 5A) present on each side of head region
near margin. Dorsal microducts (Fig. 2E) each
about 2.0 µm wide, distributed as in diagnosis.
Simple pores (Fig. 5F) each about 1.5 µm wide, each
present near a microduct. Anal plates each 48 µm
long, 18 µm wide; setae as in diagnosis. Anal ring
(Fig. 5G) as in diagnosis.
Margin. Marginal setae (Fig. 5C) slender, each
13–41 µm long, longest towards posterior apex;
number and distribution as in diagnosis. Stigmatic
setae (Fig. 5D): median seta longest, clavate, each
22–25 µm long; lateral setae bulbous, short, each
2–4 µm long.
Venter. Abdomen with pairs of setae on posterior
3 segments, each 22–25 µm long, those on
posterior abdominal segment longest. Submarginal
setae (Fig. 5H & L) distributed as in diagnosis.
Ventral microducts (Fig. 5J) each 1.5–2.0 µm wide,
distributed as in diagnosis. Spiracular disc-pores
(Fig. 5K) each about 2 µm wide, with 3 loculi;
present in a line of 3 pores from each spiracle to
margin. Clypeolabral shield 55 µm wide. Legs
well-developed, each trochanter + femur 60–73 µm
long, tibia + tarsus 65–70 µm long, claw (Fig. 5I)
15–16 µm long; prothoracic legs shortest. Antennae
6-segmented, each 133–138 µm long.
Biology
No information on the biology of P. colae has been
published. The single specimen of the form without
cribriform plates collected recently in Ghana was
on the under surface of a leaf of Theobroma cacao
alongside the leaf vein. The presence of
second-instar males (see Material studied) suggests
that the species reproduces sexually.
Diagnostic features
See diagnostic features under P. andersoni.
Distribution
Afrotropical Region: Ghana, Sierra Leone.
Host plants
Sterculiaceae: Cola acuminata,Theobroma cacao.
Lauraceae: Persea americana.
Discussion
Pseudocribrolecanium appears similar to
Cribrolecanium Green (Myzolecaniinae), with
which it shares dorsal cribriform plates and highly
reduced appendages. However, this resemblance is
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Figure 5. Pseudocribrolecanium colae (Green & Laing), first-instar nymph (drawn from specimen collected in Ghana).
A, trilocular pore; B, dorsal seta; C, marginal seta; D, stigmatic setae; E, dorsal microduct; F, simple pore; G, anal ring
(right half); H, ventral seta; I, claw; J, ventral microduct; K, spiracular disc-pore; L, ventral submarginal cephalic seta.
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superficial. Cribriform plates are also present on
some other coccids. Hodgson (1994) describes the
following genera as having cribriform plates:
Eutaxia Green, Hemilecanium Newstead and
Stictolecanium Cockerell (Coccinae: Saissetiini);
and Halococcus Takahashi and Myzolecanium
kibarae Beccari (Myzolecaniinae: Myzolecaniini).
Furthermore, reduction of appendages is observed
frequently on many members of the Coccidae, e.g.
the Myzolecaniinae, Coccinae (Coccini &
Paralecaniini in part), Antandroya spp.,
Lecanochiton spp., Neolecanochiton grevillea
Hempel, Paracardiococcus actinodaphnis
Takahashi, Platinglisia noacki Cockerell,
Pseudokermes spp., Schizochlamidia mexicana
Cockerell & Parrott (Cardiococcinae), Cissococcus
fulleri Cockerell (Cissococcinae), Cyphococcus
caesalpiniae Laing (Cyphococcinae), Scythia
craniumequinum Kiritshenko (Eriopeltinae) and
Physokermes spp. (Eulecaniinae).
Cribrolecanium possesses the following features
that separate it from Pseudocribrolecanium
(character states on Pseudocribrolecanium in
brackets): (i) the presence of a membranous
bag-like structure which contains the spiracular
pores (see Hodgson 1995) (spiracular disc-pores
not within a bag-like structure), (ii) the restriction
of pregenital disc-pores to immediately around the
vulvar region (spreading anteriorly), (iii) the lack of
spiracular setae (3–5), (iv) complete absence of
preopercular pores (present), (v) presence of small
groups of dorsal microducts on area anterior to
anal plates (absent), (vi) antennal length less than
the width or only slightly longer, never twice its
width (length of antennae 3–4 times its width), and
(vii) an anal ring with 16–18 setae (8 in
Pseudocribrolecanium ). The two genera also differ
greatly at the ecological level, with Cribrolecanium
being found on the roots and in hollow branches of
its host, whereas Pseudocribrolecanium feeds on
the leaves.
Pseudocribrolecanium shares important features
with the Paralecaniini (Coccinae), particularly with
species of the genus Paralecanium Cockerell. These
shared features are: (i) the position of the eyes on
the dorsum away from the margin, (ii) a marginal
sclerotized band, (iii) translucent furrows, and (iv)
numerous marginal setae. However, in
Paralecanium, as well as other genera in the
Paralecaniini, the multilocular disc-pores are
restricted to immediately anterior to the genital
opening and the stigmatic clefts are usually quite
distinct (Hodgson 1994). In Pseudocribrolecanium,
the multilocular disc-pores are found also on some
anterior abdominal segments, and the stigmatic
cleft is poorly developed or lacking. In addition, the
morphology of the first-instar nymphs does not
support a close relation with the Paralecaniini. Very
few studies have described or illustrated the
first-instar nymphs of the Paralecaniini, but those
species which have been described (e.g.,
Austrolecanium cappari (Froggatt), A. sassafras
Gullan & Hodgson, Maacoccus cinnamomicolus
(Takahashi), Paralecanium paradeniyense Green,
P. planum (Green), and Xenolecanium takahashii
Kondo) share many important features, i.e., a seta
next to each mesothoracic and metathoracic coxa,
six pairs of ventral submedian setae, and stigmatic
setae positioned submarginally on either side of a
deep stigmatic cleft (Gullan & Hodgson 1998,
Hodgson and Martin 2001, Kondo et al. 2005).
However, on the first-instar nymphs of
Pseudocribrolecanium, the stigmatic clefts are very
shallow, the stigmatic setae are positioned along
the margins and not on the sides of a deep stigmatic
cleft, and no seta is found next to either the meso-
and metathoracic coxae, suggesting that
Pseudocribrolecanium is not closely related to the
Paralecaniini despite the similarities shared by the
adult females. Further studies using additional
material, molecular techniques and adult males,
may help elucidate the phylogenetic position of
Pseudocribrolecanium.
Acknowledgments
I am grateful to P.J. Gullan and C.J. Hodgson for
reviewing the manuscript, to D. Matile-Ferrero for
the French translation of the Abstract, and to one
anonymous reviewer who helped improve the
manuscript. This research was supported by grant
DEB-0118718 from the U.S. National Science
Foundation (Partnerships for Enhancing Expertise
in Taxonomy program) to P.J. Gullan, and in part
by the Department of Entomology and Plant
Pathology, Auburn University.
Editor's Note
Paper copies of this article will be deposited in the
following libraries. Senckenberg Library, Frankfurt
Germany; National Museum of Natural History,
Paris, France; Field Museum of Natural History,
Chicago, Illinois USA; the University of Wisconsin,
Madison, USA; the University of Arizona, Tucson,
Arizona USA; Smithsonian Institution Libraries,
Journal of Insect Science | www.insectscience.org ISSN: 1536-2442
Journal of Insect Science: Vol. 6 Article 1 14
Washington D.C. U.S.A.; The Linnean Society,
London, England.
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