Accepted by C. Hodgson: 31 Jan. 2007; published: 5 Mar. 2007 23
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2007 · Magnolia Press
Zootaxa 1415: 23–33 (2007)
Taxonomy of Toumeyella lomagundiae Hall and T. obunca De Lotto, and their
transfer to Hallicoccus gen. nov. (Hemiptera: Coccoidea: Coccidae)
Department of Entomology, University of California, 1 Shields Avenue, Davis, California 95616-8584, U.S.A.
Two African soft scale insects, Toumeyella lomagundiae Hall and T. obunca De Lotto, are transferred to Hallicoccus gen.
nov. The adult females of both species are redescribed, and the first-instar nymph of T. lomagundiae is described. A
revised taxonomic key to separate the adult females is provided. The affinity of Hallicoccus gen. nov. with the genus
Toumeyella Cockerell is briefly discussed.
Key words: Africa, Coccidae, keys, new genus scale insect, taxonomy
Deux espèces des cochenilles africains, Toumeyella lomagundiae Hall et T. obunca De Lotto sont transférés au nouveau
genre Hallicoccus gen. nov. Les femelles adultes des les deux espèces et la larve du premier stade des T. lomagundiae
sont redécrites (ou décrites) et illustrées. Une clé dichotomique est proposée pour les femelles. L'affinité du genre Halli-
coccus dans du genre Toumeyella Cockerell, est discutée.
The genus Toumeyella Cockerell, 1895, currently contains 18 species (Ben-Dov et al 2006), of which 16 occur
in the New World and 2 in Africa. Kondo & Williams (2002) indicated that the African species are not conge-
neric with the type species, T. mirabilis (Cockerell), based on a morphological study of the adult females and
Toumeyella is well represented in the Nearctic Region (North America and northern Mexico) with 12
described species, but it is also known from the Neotropical region, with 2 species described from Brazil and
1 species from Cuba (Ben-Dov et al, 2006; Heidel and Köhler, 1979; Hempel 1929, 1932; Kondo & Williams,
2002, 2003, 2004). Hall (1935), when describing T. lomagundiae, commented as follows: “It [T. lomagundiae]
has very distinctive characteristics and appears to be referable to the genus Toumeyella, Ckll., where it has
been placed provisionally”, suggesting that the species might not be congeneric with the New World Toum-
eyella. Later, De Lotto (1966) added a second African species, T. obunca, recognizing its close affinities with
T. lomagundiae, and stated that it was uncertain whether the two African species were actually congeneric
with the type of Toumeyella, Lecanium mirabile Cockerell. As noted by De Lotto (1966), the two African spe-
cies are closely related to each other, but have little in common with the New World species; thus a new genus
Hallicoccus is erected here to accommodate them.
24 · Zootaxa 1415 © 2007 Magnolia Press
Material and methods
Specimens were slide-mounted using the method discussed by Kosztarab (1996). The terms for morphologi-
cal features of the adult female follows mostly that of Hodgson (1994), and for the first-instar nymph that of
Kondo (2006). Measurements of specimens were made using an ocular micrometer on an Olympus phase-
contrast microscope. The illustrations of the coccids follow the conventional style used for the Coccoidea,
with the dorsal surface of the body depicted on the left side of the drawing and the ventral surface on the right,
and with enlargements of important features arranged around the illustration. The total number of specimens
used for each description is given in parentheses, e.g. (n=35). In the material studied, the number of slides and
specimens on each slide are recorded 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 recorded when all specimens are adult
females. The two species are here redescribed and illustrated adding previously unreported morphological
features, i.e., preopercular pores (in H. obunca) and ventral tubular ducts (in H. lomagundiae). Although
Hodgson (1969) provided morphological features to separate the two species, a key is given here which incor-
porates the new features given above.
AUCC Auburn University Coccoidea Collection, Alabama, U.S.A.
BMNH The Natural History Museum, London, U.K.
SANC: South African National Collection of Insects, Pretoria, South Africa
The taxonomic position of Hallicoccus remains uncertain. It shares many features with the Eulecaniinae, Coc-
cinae and Myzolecaniinae as defined by Hodgson (1994). Hallicoccus lomagundiae and H. obunca are clearly
closely related. In the keys of Hodgson (1994) to subfamilies, tribes and type species of genera of Coccidae,
H. lomagundiae keys out to the subfamily Coccinae, tribe Coccini, because of the presence of dorsal tuber-
cles. However, H. obunca lacks dorsal tubercles and keys out to the subfamily Myzolecaniinae. Besides the
reduction of limbs, Hallicoccus appears to have little in common with Myzolecanium kibarae Beccari, the
type species of the Myzolecaniinae. Kondo & Williams (2002) suggested that the Myzolecaniinae is com-
posed of several unrelated lineages, and that the morphology of first-instar nymphs indicates that many genera
currently included in the Myzolecaniinae, including H. lomagundiae (as Toumeyella lomagundiae) are not
closely related to the Myzolecanium-group, which group is composed of Cribrolecanium Green, Cryptostima
Ferris, Halococcus Takahashi, Houardia Marchal and Myzolecanium Beccari. Hallicoccus shares some fea-
tures with the Eulecaniinae, particularly to Ericeroides Danzig to which it shares the presence of numerous
stigmatic spines (although these do not extend onto the dorsal submargin as in Hallicoccus) and a discal seta
on each anal plate. It also shares many features with Saissetia coffeae (Walker) [Coccinae: Saissetiini], the
type species of the genus Saissetia, particularly a characteristic H-shape ridge on the dorsum, dorsal submar-
ginal tubercles (absent in H. obunca), and a discal seta on each anal plate.
Further studies using other growth stages, adult males, and molecular data should elucidate the phyloge-
netic relationships of Hallicoccus in the future.
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TAXONOMY OF TOUMEYELLA
Hallicoccus Kondo, new genus
Type species: Toumeyella lomagundiae Hall, 1935, by present designation.
Unmounted material. Body of adult female convex. Living insects with an H-shaped ridge on dorsum
somewhat similar to that found on many members of the genus Saissetia Déplanche.
Dorsum: derm with well-developed areolations. Anal plates each with 4 apical setae. A narrow sclerotized
crescent around anal plates present. Anal ring with 6 or 8 setae. Submarginal dorsal tubercles and preopercular
pores present or absent. Dorsal setae sharply spinose, stout, with straight and pointed tips. Dorsal microducts
heavily sclerotized, with a single central opening. Dorsal simple pores present, small. Eyespots not detected.
Margin: Marginal setae similar to dorsal setae. Stigmatic spines present submarginally on dorsum, total-
ling 3–16 in each group; each stigmatic area with usually more than 6 spines of various sizes.
Venter: Antennae small, 5 or 6 segmented. Mouthparts typical of the family; labium 1 segmented, with 8
labial setae. Legs greatly reduced, with all segments fused into one segment plus claw, or reduced to a minute
sclerotized plate with a few associated setae. Spiracles relatively small. Ventral tubular ducts present on area
between anterior and posterior spiracles. Ventral setae all slender or slender medially and sharply spinose sub-
marginally and submedially; prevulvar setae similar to other ventral setae or only slightly longer. Spiracular
disc-pores with 5–12 loculi, mostly with 5 loculi. Pregenital disc-pores each with 6–12 (mostly 10) loculi,
present around vulva and extending medially on all median areas of abdomen and thorax. Ventral microducts
scattered evenly on venter or most abundant submarginally and around mouthparts.
Etymology. The new genus Hallicoccus is named after the late Dr. Wilfrid John Hall (who collected and
described the type species Toumeyella lomagundiae), and comprises the surname plus “coccus”, the latinized
version of the Greek word “kokkos”, meaning a small grain or seed, an ending commonly used to describe
Key to species of Hallicoccus gen. nov. based on adult females
1. Submarginal dorsal tubercles present; preopercular pores absent; most stigmatic spines bulbous...............
............................................................................................... Hallicoccus lomagundiae (Hall), comb. nov.
– Submarginal dorsal tubercles absent; small preopercular pores present; all stigmatic spines sharply spi-
nose..........................................................................................Hallicoccus obunca (De Lotto), comb. nov.
Hallicoccus lomagundiae (Hall), comb. nov.
(Fig. 1 & 2)
Toumeyella lomagundiae Hall, 1935: 81; De Lotto 1966: 149; Hodgson 1969: 36; Ben-Dov 1993: 330.
Holotype. ZIMBABWE, 1 (1): Sinoia, coll., 15.xii.1927, ex Bauhinia macrantha, (det. date: 11.vii.1934),
det. W.J. Hall, No. 1770 (BMNH).
Paratypes. ZIMBABWE, 1 (2): Same data as Holotype (BMNH); ZIMBABWE, 1 (4): Same data as
Holotype, except No. 185 (Note: not labelled as Paratypes but with same data as Holotype) (BMNH); ZIM-
BABWE, 28 (28): Same data as Holotype, except AL-141-98 (mounted from BMNH dry Type material)
26 · Zootaxa 1415 © 2007 Magnolia Press
FIGURE 1. Hallicoccus lomagundiae (Hall), adult female. A, derm areolation; B, submarginal dorsal tubercle; C, stig-
matic setae; D, dorsal microduct; E, simple pore; F, dorsal and marginal seta; G, anal plate; H, pregenital disc-pores; I,
spinose ventral submarginal seta; J, slender ventral submedial seta; K, anal ring (right half); L, reduced metathoracic
legs; M, ventral microduct; N, top and side view of ventral tubular duct; O, spiracular disc-pores; P, antenna.
Non-type material. MALAWI, 1 (1): Chitedzi Exp. Station, 1.vii.1966, ex Piliostigma thouningii, coll.
C.J. Hodgson, BM 1967-558 (528), det. C.J. Hodgson (BMNH).
Description. Adult female (Fig. 1)
Unmounted material. “Old adult female very highly convex, the height being approximately the same as
Zootaxa 1415 © 2007 Magnolia Press · 27
TAXONOMY OF TOUMEYELLA
the diameter. The sides are roughly vertical with the top more or less evenly rounded but surmounted at the
apex by a minute protuberance. The margin is outwardly turned so as to give a small flattened base. On either
side of the dorsum is a large and usually conspicuous dent, as though the insect has been pinched between the
finger and thumb. In cross-section the adult female is either irregularly broadly oval or circular. In some spec-
imens the dorsum exhibits vertical ridges making it six or seven sided. The colour is a dull brown, darker
brown generally in the vicinity of the ridges. The surface of the dorsum is not smooth but has little irregularly
shaped flattish protuberances at wide intervals. Very young specimens have a distinct dorsal keel and the mar-
gin is set with crystalline outgrowths which are more or less regular and tend to be conical in shape. Similar
conical outgrowths occur submarginally and are directed outwards towards the margin. In somewhat older
specimens three spurs run down to the margin from the dorsal keel on either side, the conical marginal pro-
cesses are present and similar crystalline masses occur at intervals over the dorsum giving a somewhat glassy
appearance. Diameter of adult female, 5.5–7.5 mm; height, 5–7.5 mm.” (Hall, 1935).
Mounted material. Body outline subcircular or elongate oval; body 2.7–6.4 mm long, 2.3–5.0 mm wide
Dorsum. Derm of young adult females only slightly sclerotized; derm areolations well developed. Anal
plates located about 1/5 of body length from body apex. Each plate 156–199 µm long, 81–97 µm wide, ante-
rolateral margin 102–113 µm long, posterolateral margin 156–194 µm long; with 4 dorsal apical setae: 2 short
setae on inner margin, 1 large subapical seta close to outer margin, and 1 short apical seta displaced towards
outer margin, often broken off, but setal sockets clearly present; plus 2 ventral anterior margin and 1 shorter
lateral margin seta. Anal ring with 8 setae and 2 rows of translucent pores. Submarginal dorsal tubercles
present, each about 24 µm in diameter, totalling 6–18 around body. Preopercular pores absent. Setae scattered
over dorsum, each 13–17 µm long. Dorsal microducts elongate oval, each about 5 µm wide, scattered
throughout dorsum. Simple pores each about 2 µm in diameter, scattered evenly over dorsum.
Margin. Marginal setae more or less in two rows, each 11–28 µm long, longest on anal lobes, with about
12–16 setae between groups of anterior and posterior stigmatic setae. Stigmatic spines of two shapes, totalling
11–16 in each group, located on dorsal submargin, longest sharply spinose, slender, 77–98 µm long; others
spinose with greatly swollen bases, each shorter, 13–43 µm long.
Venter. Derm membranous. Antennae reduced, 6 or 7 segmented, each 76–108 µm long. Clypeolabral
shield 162–205 µm wide. Legs greatly reduced, with all segments fused but with a claw; prothoracic legs
shortest; metathoracic legs longest; total length of each leg, 32–76 µm long. Tarsal digitules on each protho-
racic leg dissimilar, one knobbed and the other spiniform; tarsal digitules on each meso- and metathoracic leg
similar, knobbed. Claws simple, with no denticle; each claw with a pair of slender, knobbed digitules. Spira-
cles relatively small, anterior spiracles usually smaller than posterior spiracles, each anterior peritreme 92–
113 µm wide, posterior peritreme 92–124 µm wide. Ventral tubular ducts scarce, present in a narrow band
between anterior and posterior spiracles. Ventral setae of two types, those on mid areas of abdomen and thorax
slender and long, each 13–17 µm long; rest of setae with broader bases, shorter, each 10–12 µm long. With 2–
4 pairs of interantennal setae present. Spiracular disc-pores with 4–12 loculi, mostly with 5 loculi, each 2–5
µm in diameter; spiracular pore bands each 2–5 pores wide. Pregenital disc-pores numerous, each 4–6 µm in
diameter, each with 6–12 (mostly 10) loculi, present medially on all abdominal segments, extending anteriorly
onto both sides of mouthparts, and reaching area around prothoracic legs; often with 1 or 2 multilocular disc-
pores on area between antennae. Microducts each about 4 µm wide, abundant around body margin and
labium, less frequent elsewhere on venter.
Diagnostic features. The adult female of Hallicoccus lomagundiae (Hall) can be easily separated from
Hallicoccus obunca (De Lotto) by the following combination of character states (character states of H.
obunca in brackets): (i) absence of preopercular pores (present), (ii) presence of submarginal dorsal tubercles
(absent), (iii) legs with all segments fused, but with a distinct claw (legs vestigial, composed of a small sclero-
tized plate plus 2 or 3 associated setae), and (iv) presence of bulbous stigmatic spines (stigmatic spines
28 · Zootaxa 1415 © 2007 Magnolia Press
Remarks. The present description of H. lomagundiae differs from previous descriptions by Hall (1935)
and Hodgson (1969) in the following points (character states of present description in parenthesis): (i) dorsal
setae most abundant near the anal plates (dorsal setae not particularly abundant around anal plates), (ii) anten-
nae 7 segmented (6 or 7 segmented), and (iii) ventral tubular ducts absent (present in a narrow band between
anterior and posterior spiracles).
FIGURE 2. Hallicoccus lomagundiae (Hall), first-instar nymph. A, trilocular pore; B, marginal setae; C, dorsal seta; D,
stigmatic spines; E, dorsal microduct; F, simple pore; G, ventral submarginal seta; H, anal ring (right half); I, spiracular
disc-pores; J, ventral microduct; K, ventral submarginal cephalic seta.
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TAXONOMY OF TOUMEYELLA
According to Hall (1935), Hallicoccus lomagundiae (as Toumeyella lomagundiae) was a common species
on Bauhinia in the Sinoia district (Lomagundi) of Zimbabwe. The second host, Piliostigma thonningii, is
closely related to Bauhinia, a genus in which it was formerly included (Missouri Botanical Garden:
Description. First-instar nymph (Fig. 2)
Paratypes. ZIMBABWE, 6(33): Same data as Holotype (slide-mounted from type dry material)
(BMNH); ZIMBABWE, 1(6): Same data as Holotype, except No.185, (BMNH).
Unmounted material. External morphology not recorded.
Mounted material. Body outline elongate oval; body 464–507 µm long, 226–280 µm wide (n=39).
Dorsum. Derm membranous. Anal plates each 47–55 µm long, 21–24 µm wide, with 4 dorsal setae: 1
inner margin seta and 3 apical (including long median apical seta), plus 1 anterior margin seta. Anal ring with
6 setae and an irregular row of pores. Dorsal setae short, each about 3 µm long; in 2 submedian parallel rows
of 4 setae on head and thorax. A trilocular pore present on each side of head region near margin. Dorsal
microducts each about 3 µm wide, present submarginally and in 2 submedian rows. Simple pores each about 2
µm in diameter, usually closely associated with a microduct. Eyespots present on margin of head.
Margin. Marginal setae slender, each 11–15 µm long, totalling about 32 around body, with 8 anteriorly
between eyes, plus (on each side) 2 between eye and anterior stigmatic setae, 2 between anterior and posterior
stigmatic setae, and 8 between posterior stigmatic setae and body apex on each side. Each stigmatic area with
a group of 3 stigmatic spines, each well differentiated from marginal setae; each median spine sharply spi-
nose, 24–28 µm long; lateral spines short, sharply spinose or bulbous, each 5–7 µm long.
Venter. Antennae 6 segmented, each 162–178 µm long. Clypeolabral shield 76–86 µm wide. Legs well
developed, each trochanter and femur with a very long seta; also with a long seta on each meso- and metatho-
racic tibia; trochanter + femur of each leg 81–86 µm long, tibia + tarsus 108–113 µm long; anterior tarsal dig-
itules dissimilar, one long and knobbed, the other short and spiniform; meso- and metathoracic tarsal digitules
both knobbed and subequal in size; claw digitules all similar and knobbed; claw with a small denticle. Ante-
rior and posterior spiracular peritremes similar in size, each 6–9 µm wide. Submedian abdominal setae in
pairs on posterior 3 segments. Submarginal setae arranged in an inner and outer row, each with 7 setae on each
side, between posterior stigmatic areas and posterior body apex, plus 1 seta on each side between anterior and
posterior stigmatic areas, and 1 pair present anteriorly on head region; other ventral setae present in a subme-
dian line, with 1 seta on each abdominal segment. Spiracular disc-pores each about 3 µm in diameter, with 3–
5 loculi (mostly 3 or 5); present in a line between each spiracle and margin, with 3 pores in each anterior line
and 4 in each posterior line. Ventral microducts each about 2 µm wide, present submarginally on each seg-
ment between inner and outer submarginal setae, except absent from between posterior-most pair; also with 2
microducts on each side between anterior and posterior stigmatic areas, and 2 on each side posterior to anten-
nae.Diagnostic features. The first-instar nymphs of H. lomagundiae can be easily diagnosed by the presence
of a very long seta on each coxa and femur, with a long seta found also in the meso- and metathoracic tibiae
(see remarks for additional comments).
Distribution of H. lomagundiae: Afrotropical Region: Zimbabwe, Malawi.
Host plants. Fabaceae: Bauhinia macrantha Oliv.; B. variegata (Ben-Dov, 1993; Hodgson, 1969); Pil-
iostigma thonningii (Schumach. & Thonn.) Milne-Redh.
Remarks. The extremely long setae observed in the legs of H. lomagundiae are unique among the Coc-
cidae in their distribution, being present in each trochanter and femora, and on each tibia of the meso- and
metathoracic legs. Williams & Hodges (1997) discussed the taxonomic features of first-instar nymphs of 52
species in 45 genera representative of 8 out of the 10 subfamilies of the Coccidae as recognized by Hodgson
(1994). In their study, Williams and Hodges reported similar long setae in the leg segments of first-instar
30 · Zootaxa 1415 © 2007 Magnolia Press
nymphs of several coccids, i.e., one long seta on the trochanters of Coccus hesperidum L., two long setae on
the trochanters of Eulecanium tiliae (L.), one long and one short seta on each trochanter of Ceroplastodes
dugesii (Signoret), and the presence of an extremely long seta on the femur which extends to near the apex of
the leg of Etiennea petasus Hodgson, Kilifia americana Ben-Dov, Milviscutulus mangiferae (Green) and Pro-
topulvinaria pyriformis Cockerell. However, long setae on the legs of the first-instar nymphs discussed by
Williams and Hodges were restricted to one segment on each leg, rather than two or more leg segments as
observed on H. lomagundiae.
Hallicoccus obunca (De Lotto), comb. nov.
Toumeyella obunca De Lotto, 1966: 149; Hodgson 1969: 39; Ben-Dov 1993: 381.
Holotype. South Africa, 1 (1): Natal, Richmond, 21.iii.1964, coll. J. Munting, ex Cnestis natalensis, H.C.
Paratypes. South Africa, 3(3): same data as Holotype, H.C. #856/3, 4 & 7 (BMNH).
Description. Adult female (Fig. 3)
Unmounted material. “Living adult females at the beginning of the stage moderately convex, with a well
developed H-mark on the dorsum; colour evenly brown.” (De Lotto, 1966).
Mounted material. Body outline elongate oval; body 1.9–2.3 mm long, 1.6–2.0 mm wide (n=3).
Dorsum. Derm of young adult females membranous, with well-developed derm areolations. Anal plates
located about 1/5–1/6 of body length from body apex. Each plate 117–121 µm long, 55–57 µm wide; antero-
lateral margin 64–66 µm long, posterolateral margin 100–102 µm long; each with 4 dorsal setae: 2 short setae
on inner margin, 1 long subapical seta and 1 short apical seta; plus 2 ventral anterior margin setae and 1 lateral
margin seta. Anal ring with 6 setae and 2 rows of translucent pores. Submarginal dorsal tubercles absent. Pre-
opercular pores each 2–3 µm in diameter, present in a group of 15–40 pores on area anterior to anal plates.
Setae robust, scattered over dorsum, each 15–22 µm long. Dorsal microducts heavily sclerotized, each about 4
µm wide, scattered throughout dorsum. Simple pores each about 2 µm in diameter, scattered evenly over dor-
Margin. Marginal setae each 12–16 µm long, longest at anal lobes, with 10–15 setae between groups of
anterior and posterior stigmatic spines. Stigmatic spines of one type only present on dorsal submargin, each
sharply to bluntly spinose, totalling 3–7 in each group, longest 120–150 µm long, others shorter, each 15–120
Venter. Derm membranous. Antennae reduced; antennal segments indistinct, probably 3–5 segmented,
98–107 µm long. Clypeolabral shield 164–175 µm wide. Legs vestigial, each composed of a small sclerotized
plate plus 2 or 3 associated setae, not easy to detect. Spiracles: anterior pair usually smaller than posterior pair,
each anterior and posterior peritreme 36–43 µm wide. Ventral tubular ducts present in a narrow band between
anterior and posterior spiracles. Ventral body setae slender, each 7–16 µm long. Interantennal setae numbering
about 2 pairs. Spiracular disc-pores each with 5–7 (mostly 5) loculi, rarely with an occasional disc-pore with
10 loculi, 3–5 µm in diameter; spiracular pore bands each 2–4 pores wide. Pregenital disc-pores numerous,
each 5–6 µm in diameter, with 10 loculi, present medially on all abdominal segments, extending anteriorly
onto area near anterior spiracles. Ventral microducts each 3 µm wide, scattered throughout venter.
Diagnostic features. See under diagnostic features of H. lomagundiae comb. nov.
Remarks. The present description of H. obunca differs from that given by De Lotto (1966) in the follow-
ing points (character states of present description in parenthesis): (i) preopercular pores absent (present), (ii)
Zootaxa 1415 © 2007 Magnolia Press · 31
TAXONOMY OF TOUMEYELLA
absence of legs (legs vestigial), and (iii) slightly larger ranges for some measurements. Perhaps De Lotto did
not notice the vestigial legs and the preopercular pores in H. obunca because of their small size, each leg
being represented by a small sclerotized plate plus 2 or 3 associated setae, and the preopercular pores being
only about 0.5–1.0 µm wider than the more abundant dorsal microducts. First-instar nymphs were not avail-
able during the present study.
Distribution of H. obunca: Afrotropical Region: South Africa.
Host plants. Connaraceae: Cnestis natalensis (Hochst.) Planch. ex Sond.
FIGURE 3. Hallicoccus obunca (De Lotto), adult female. A, enlargement of dorsal derm; B, stigmatic spines; C, dorsal
microduct; D, simple pore; E, dorsal and marginal seta; F, preopercular pores; G, anal plate; H, pregenital disc-pore; I,
anal ring (right half); J, metathoracic leg; K, ventral setae; L, ventral microduct; M, ventral tubular duct; N, spiracular
disc-pores; O, antenna.
32 · Zootaxa 1415 © 2007 Magnolia Press
The two African species hitherto included in Toumeyella (i.e., H. lomagundiae and H. obunca) have very little
in common with the New World genus Toumeyella. The adult females of Hallicoccus differ from Toumeyella
mainly in possessing the following features (character states of Toumeyella in brackets): (i) female body with
an H-shaped ridge on dorsum (without an H-shaped ridge on dorsum), (ii) dorsal and marginal setae subequal
in shape (not subequal in shape), (iii) submarginal dorsal tubercles present or absent (always absent), (iv) stig-
matic spines numbering 3–16 in each group, usually more than 6 (0–3, usually 3), (v) ventral tubular ducts
present in a narrow band between anterior and posterior spiracles (usually present around perivulvar area and
restricted to abdomen), and (vi) pregenital disc-pores mostly with 10 (mostly with 5) loculi (Williams &
Kosztarab, 1972; Kondo & Williams 2003, 2004). Furthermore, the first-instar nymphs of the two genera dif-
fer mainly in possessing the following features (character states of To um ey el la in brackets): (i) dorsal setae
present, about 4 pairs present in 2 parallel rows on head and thorax (dorsal setae present or absent, when
present numbering 1 pair on head region), (ii) antennae 6 segmented (5 segmented), and (iii) legs with
extremely long setae on trochanters, femora and tibiae (legs without extremely long setae).
I am grateful to Jon Martin (BMNH) and to Ian Millar (SANC) for the loan of specimens used in this study. I
thank C.J. Hodgson (National Museum of Wales, Cardiff), Y. Ben-Dov (The Volcani Center, Israel), and an
anonymous reviewer for reviewing 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 (University of California, Davis), and in part by the Department of Entomology and Plant Pathology,
Ben-Dov, Y. (1993) A Systematic Catalogue of the Soft Scale Insects of the World (Homoptera: Coccoidea: Coccidae)
with data on geographical distribution, host plants, biology and economic importance. Flora & Fauna Handbook,
No. 9. Sandhill Crane Press, Gainesville, FL., 536 pp.
Ben-Dov, Y., Miller, D.R. & Gibson, G.A.P. (2006) ScaleNet: a database of the scale insects of the world. [Online] http:/
De Lotto, G. (1966) A new genus and four new species of Coccidae (Homoptera) from South Africa. Proceedings of the
Linnean Society of London, 177, 143–149.
Hall, W.J. (1935) Observations on the Coccidae of southern Rhodesia. - VI. Stylops, 4, 73–84.
Heidel, W. & Kohler, G. (1979) Toumeyella cubensis sp. n. (Hemiptera: Coccinea - Coccidae) - eine Schildlaus in kuba-
nischen Zitruskulturen. (In German; summary in English). Zoologischer Anzeiger, Jena, 202, 132–144.
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