Table Of ContentPROC. ENTOMOL. SOC. WASH.
101(3). 1999, pp. 522-539
TWO NEW GENERA AND SPECIES OF MEALYBUGS
(HEMIPTERA: COCCOIDEA: PSEUDOCOCCIDAE) THAT PRODUCE
PLANT GALLS
Douglas J. Williams and Douglass R. Miller
(DJW) Department of Entomology, The Natural History Museum, Cromwell Road,
London SW7 5BD, U.K.; (DRM) Systematic Entomology Laboratory, PSI, Agricultural
Research Service, U.S. Department of Agriculture, Bldg. 046, BARC-West, Beltsville,
MD
20705-2350, U.S.A. (e-mail: [email protected]).
—
Abstract. In the mealybug family Pseudococcidae, only 23 species in 14 genera are
known to be gall formers. The purpose of this paper is to describe two new South Amer-
ican genera and species that cause galls; namely Miconicoccus ruebsaameni and Quad-
rigallicoccus lauracearum. These mealybugs are legless and apparently are related to
Antonina Signoret. They are the first gall-forming mealybugs known from the continental
land mass of the New World. A review of gall-forming mealybugs from other parts of
the world is also presented.
Key Words: gall-forming mealybugs, galls, Coccoidea, Pseudococcidae, new genera,
new species, Miconicoccus ruebsaameni, Quadrigallicoccus lauracearum,
South America, review
Gall producing scale insects have long outside of Hawaii only 5 other species of
been a subject ofgeneral interest (Beardsley mealybugs are known to form definite galls,
1984), particularly in Australia where a sig- i.e., Cataenococcus gallicolus (Mamet),
nificant proportion of the Coccoidea biota Eurycoccus stenilichti Williams, Grewia-
produces galls (Gullan 1984). Scale insects coccus gregalis Brain, Lantanacoccus sau-
in 10 families are gall formers, including roides Williams and Granara de Willink,
the Margarodidae (2 species in 2 genera), and Pseudoripersia turgipes (Maskell). In
Pseudococcidae (23 species in 14 genera), Hawaii, 1 1 species in 5 genera are known
Eriococcidae (more than 100 species in at to produce galls (Beardsley 1984) (actually
least 21 genera), Kermesidae (4 species in Beardsley did not mention Pseudococcus
3 genera), Coccidae (1 species in 1 genus), antricolens Ferris and Gallulococcus teno-
Asterolecaniidae, Cerococcidae, and Lecan- riori Beardsley but included 2 species in his
odiaspididae (most species in these families figures that do not produce galls).
form at least a pit or depression on their Since information on gall-forming
host, some form galls that enclose the body mealybugs has not been reviewed since
of the female), Beesoniidae (all species of Beardsley (1984), we felt that it would be
this family form galls some of which are useful to provide a summary as part of the
colonial), and Diaspididae (16 species in 12 introduction. While writing this section and
genera). examining illustrations of each species, we
There is a surprising paucity of true gall were struck by the fact that many ofthe true
formers within the Pseudococcidae; in fact. gall-forming species have one or more mor-
VOLUME NUMBER
101, 3 523
phological modifications that are shared by produces galls on Casiiarina (Casuarina-
other unrelated gall-forming mealybugs and ceae) by causing branchlets to become dis-
apparently relate to the gall-forming habit. torted and curl around the body of the in-
These apparent, gall-mediated, morpholog- sect; it occurs in Australia (Williams 1985).
ical characteristics include: a series oflarge Phyllococcus oahuensis (Ehrhorn) has a
conical setae on at least the posterior apex heavily sclerotized and flattened posterior
of the abdomen, often these are modified portion of the abdomen that is specialized
anal-lobe cerarii; a pear-shaped body with for sealing the entrance to the gall; it forms
the posterior apex narrowest; hind legs that erect galls on the leaves of Urera (Urtica-
are enlarged or modified in some manner; ceae) (Ferris 1948).
the posterior apex of the body that is scler- We were surprised to find that several
otized and may be modified. Species that true gall-forming species seem to have no
fall in this category are as follows: Gallu- unusual morphological modifications. With-
lococcus tenorioi has the body pear shaped; in this category ofmealybug gall formerwe
it forms shallow pocket galls on the leaves found species that have gall-forming con-
of its Metrosideros (Myrtaceae) host geners with morphological modifications.
(Beardsley 1971). Grewiacoccus gregalis Species without obvious modifications in-
has enlarged hind coxae and the posterior clude: Cataenococcus gallicolus which
apex of the abdomen is covered with en- forms galls on the twigs of an unidentified
larged conical setae; it forms a blunt leaf creeper in Madagascar (Mamet 1954). Eu-
gall on the undersides of its Grewia (Tili- rycoccus sternlichti Williams forms gallson
aceae) host in South Africa (Brain 1918). the twigs of Quercus ithuburensis (Faga-
Lantauacoccus sauroides has numerous ceae) that are up to 8 cm long and 3 mm
dorsal enlarged setae; it forms deep, hard- thick in Israel (Williams 1958). Nesoped-
ened gall-like depressions on the leaves of ronia cibotii (Beardsley) does not always
Lantana (Verbenaceae) in Haiti and Jamai- cause host deformation; when galling oc-
ca (Williams and Granara de Willink 1992). curs it is limited to the edges ofthe pinnules
Nesopedronio acanthocauda (Beardsley) which are curled on its Cibotium (Dickson-
has the anal-lobe cerarii so enlarged that iaceae) host (Beardsley 1957). Nesopedron-
they form a continuous row of setae across ia hawaiiensis (Ferris) forms a rosette-like
the apex of the abdomen and has only the gall of fern pinnules (Beardsley 1959) on
hind two pair of cerarii present; the gall is its Dicranopteris (Gleicheniaceae) host.
formed from a rolled fern pinnule on its Di- Pseudococcus antricolens produces finger-
cranopteris (Gleicheniaceae) host (Beards- like galls on the upper surface ofthe leaves
ley 1957). Nesopedronia crypta (Beardsley) of Santalum freycinetianum (Santalaceae)
has the anal-lobe cerarii so enlarged that in Hawaii (Ferris 1948). Pseudococcus gal-
they form a continuous row of setae across licola Ehrhorn produces pocket galls on the
the apex of the abdomen; the gall is formed upper surface of Santalum (Santalaceae)
from a rolled fern pinnule on its Dicran- leaves (Ferris 1948). Pseudotrionymus mul-
opteris (Gleicheniaceae) host (Beardsley tiductus Beardsley forms a gall by rolling
1957). Nesopedronia dura (Beardsley) has the entire leaf near the midrib of its Syzy-
the posterior apex of the abdomen sclero- gium (Myrtaceae) host (Beardsley 1959).
tized and developed into a small pygidial- Pseudotrionymus refertus (Ferris) forms a
like flap; it forms a hard, gall-like roll at gall by rolling the leaf margins of its Eu-
the apex of each affected pinnule on its Di- genia (Myrtaceae) host (Beardsley 1959).
cranopteris (Gleicheniaceae) host (Beards- Ohiacoccus cryptus Beardsley was re-
ley 1957). Pseudoripersia turgipes has dor- ported as forming galls (Beardsley 1984)
sal conical setae, large robust legs, and a but as far as we can determine it does not
large anal-lobe cerarius with many setae; it cause any host deformation. According to
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524 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
the original description ofBeardsley (1971) abdomen or at base ofanal tube, always with
"All the specimens were found at the bases setae at least as long as diameter ofring . . .
Antoniiui Signoret
ofleafpetioles where these were tightly ap- - Anal ring on surface, situated about twice its
pressed to the twigs and where the insects diameter from posteriorend ofbody, with se-
[were] imbeded in thick tomentum which is tae shorter than diameter of ring
characteristic ofthe typica variety ofohia." Paludicocciis Ferris
There are several species of mealybugs 3c. Posterior end of body pointed. Anal ring at
endofanaltube,with6setae. Multiplecirculi
that cause host deformation but are not ob-
present. Cluster of minute pores absent from
ligate gall formers and are not specialized each side of vulva
morphologically for gall habitation. They .... Miconicoccus Williams and Miller, n. gen.
include: Hypogeococciis festehamis (Lizer Posterior end of body rounded. Anal ring on
y Trelles) on Cereiis (Cactaceae) andEchin- body surface, not at end ofanal tube, without
setae. Circuli absent. Cluster ofminute pores
opsis (Cactaceae) in Argentina (Williams
present on each side of vulva
and Granara de Willink 1992); H. pimgens Qiiadrif^alliroccu.s Williams and Miller, n. gen.
Granara de Willink mainly on Cereus and
Eriocerius (Cactaceae) in Argentina and Methods
Brazil (Williams and Granara de Willink
Terminology in the descriptions follows
1992); Maconellicoccus hirsutus (Green) that of Williams and Watson (1988) and
on many hosts in many parts of the world Gimpel and Miller (1996). Sclerotized slits
(Williams 1996); Nipaecoccus viridis
are present laterad of the hind legs of the
(Newstead) on several hosts in Jordan
first instar and laterad of the middle legs of
(Sharaf and Meyerdirk 1987); Phenacoccus
the second-instar male of Quadrigallicoc-
herreni Cox and Williams on Manihot es- cus lauracearum. We have not observed
culenta (Euphorbiaceae) in South America these structures before and are uncertain of
(Cox and Williams 1981); Phenacoccus
their function. Measurements and numbers
manihoti Matile-Ferrero on Manihot escu- are from 10 specimens when available, and
lenta and several other hosts in South are given as an average followed by the
America (Cox and Williams 1981); Phen-
range in parentheses. Depositories of spec-
acoccus pan'us Morrison on many hosts imens are: The Natural History Museum,
throughout the world (Williams, personal London (BMNH); National Museum of
observations). Natural History, Beltsville, MD (USNM);
In this paper we describe two genera and R. M. Bohart Museum, Davis, CA (UCD);
two species from Central and South Amer- Museum National d'Histoire Naturelle, Par-
ica that are the first gall-producing mealy- is (MNHN). The coauthors are equally re-
bugs to be described from continental land sponsible for the research and production of
masses in the New World. These genera can
this paper.
be distinguished from others in the area by
the following key modified from couplets 2 Results
and 3 of the key to genera in Williams and
Miconicoccus Williams and Miller,
Granara de Willink (1992):
new genus
2. Minute ducts present ventrally in distinctoval Type species. Miconicoccus ruebsaa-
to round areas posterior of hind spiracles
Chaetococciis Ma.s.kell nieni Williams and Miller.
- Minute ducts absent ventrally posterior of Diagnosis. Adultfemale: Body rotund;
hind spiracles; sometimes with minute pores posterior apex sclerotized; segmental line
in this area 3a between segments VII and VIII heavily
3a. Antennae each with 2 or 3 segments 3b
- Antennae each represented by unsegmcnted sclerotized, with associated apophysis. Cer-
sclerotized area 3c arii represented by paired spine-like setae
3b. Anal ring eitheron surface at posteriorendof at apex of opening of anal tube. Trilocular
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VOLUME 101, NUMBER 3 525
pores uncommon; multilocular pores pre- Miller/Holotype/C.I.E./B.M.196" (USNM).
sent. Oral-collar tubular ducts of Antonina In addition there are 3 other adult female
type with partial external tube surrounding paratypes, 2 immature paratypes, and 4 first
internal vestibule; oral collars abundant. instar paratypes in USNM; there is 1 adult
Multiple circuli present. Anal ring with nu- female paratype and 1 immature paratype
merous pores, invaginated in long tube that in BMNH. All material is apparently from
has second ring near opening of invagina- the same location although one USNM
tion; anal-ring setae apically capitate. Legs slide indicates "On Miconia/Dr. Edw. Rub-
absent. Spiracles without pores in atrium. saamen, coll./ rec'd 1907" while the second
Antennae represented by unsegmented, says "(larvae)/ on Miconia sp./ from Edw.
sclerotized area containing several setae. Rubsaanie—nI July 1, 08."
First instar: With 2 sizes of trilocular Notes. This species was first mentioned
pores. Without ostioles. Anal-ring setae en- by Riibsaamen (1907) for whom the species
larged, with blunt apices. Enlarged setae on is named; he described the gall and com-
dorsum of segment VII. Cerarii present on pared the appearance of the anal wax with
posterior 2 abdominal segments. With 2 or that of Xylococcus filiferus Low. His de-
3 circuli. Apex—of abdomen sclerotized. scription indicated that the gall is a knotty
Etymology. The name of this genus is swelling on the branch and frequently is
formed from the generic name of the plant formed at a branch node.
host Miconia (Melastomataceae) and the Description. Adult female (Fig. 1):
mm
Latin coccum meaning seed or scale insect. Slide-mounted holotype 1.2 long, 1.0
mm mm
Miconico—ccus is a masculine noun. wide; paratypes 1.3(0.8-1.6) long,
mm
Notes. This genus is remarkably similar 1.2(1.0-1.5) wide. Body nearly round,
to Antonina Signoret and undoubtedly is posterior apex sclerotized. Segments V and
closely related, but differs by having mul- VI fused laterally.
tiple circuli on the ventral abdomen, anten- Dorsum with trilocular pores present in
nae represented by an unsegmented sclero- small numbers in medial and mediolateral
tized area, and paired, conspicuous spine- areas from head to segment V (on some
like setae on either side of the opening to paratypes triloculars were not located). Dis-
the anal tube. Antonina lacks circuli or has coidal pores uncommon, in small numbers
only 1, has antennae that are 2- or 3-seg- on submarginal areas of segments VI and
mented, and lacks paired setae at apex of VII, smaller than diameter of trilocular
opening of anal tube. The first instars of pore. Oral-collar tubular ducts of 3 variable
these genera are virtually identical except sizes becoming smaller anteriorly, abundant
that Miconicoccus has 2 or more circuli and over surface except absent from segment
lacks ostioles. Antonina rarely has circuli, VIII. Setae on segment VIII abundant, pos-
but when present there is only 1, and al- terior setae enlarged and spine-like; 1 or 2
ways has at least the posterior pair of osti- setae on segment VII also enlarged; setae
oles (Yang and Kosztarab 1967). on remainder of dorsum very small and in-
conspicuous, becoming thinner anteriorly.
Miconicoccus riiebsaameni Williams and
Segments VII and VIII, and posterior part
Miller—, new species of VI sclerotized. Segmental line between
Type material. The holotype adult fe- segments VII and VIII heavily sclerotized
male is mounted singly on a slide with the with conspicuous internal apophysis; seg-
following information: Left label "Peru, mental line between segments VI and VII
Tarapoto/ on Miconia ibaguensis/ galls/ E. also heavily sclerotized, with smaller
H. Riibsaamen/ X. 1902 No 95/gall no 548/ apophysis.
see Marcellia 1907 6:164/ 165; right label Anal ring represented by heavy band of
"'Miconicoccus/ ruebsaamenil Williams & more than 200 pores; present at end of in-
526 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 1. Miconicocciis ruebsaaweiu. Adult female.
VOLUME 101. NUMBER 3 527
vaginated anal tube about 200 |x long; ring VIII abundant, posterior setae enlarged and
with 6 setae that protrude from end oftube, spine like; setae on remainder of dorsum
each seta with small distal club; anal tube very small and inconspicuous. Segments
with heavy band on venter anterior of exit VII and VIII, and posterior part ofVI scler-
to exterior. otized. Segmental line between segments
Venter with multilocular pores of2 sizes, VII and VIII heavily sclerotized with con-
smaller size on thorax and head and on seg- spicuous lateral apophysis; segmental line
ment VIII, larger size in medial and medio- between segments VI and VII also heavily
lateral areas from posterior thorax to seg- sclerotized, without apophyses.
ment VII, multilocular pores concentrated Anal ring represented by heavy band of
around spiracles, but not in atrium. Discoi- more than 200 pores; present at end of in-
dal pores of 2 sizes, smaller size in small vaginated anal tube 105(99-111) |x long;
numbers in mediolateral areas of segments ring with 6 setae that protrude from end of
II to VI, with clear center, larger size in tube; anal tube without heavy band anterior
medial and mediolateral areas of metatho- of exit to exterior.
rax to segment IV (some paratypes with Venter with multilocular pores of2 sizes,
pores absent on segment IV) with heavy smaller size near spiracles and on segment
rim and granular center. Trilocularpores ab- VIII, larger size in medial and mediolateral
sent. Oral-collar tubular ducts of same 3 areas from just anterior of mouthparts pos-
sizes as on dorsum, abundant over surface terior to segment VII; concentrated around
except absent from segment VIII. Exclud- spiracles, but not in atrium. Discoidal pores
ing anal-lobe seta, longest setae on segment absent. Trilocular pores absent. Oral-collar
VIII; anal-lobe seta anterior of anal open- tubular ducts of same sizes as on dorsum,
ing, setae on rest ofsurface unusually small abundant over surface except absent from
except on segment VIII where posterior and segment VIII. Excluding anal-lobe seta,
lateral setae are slightly enlarged. Segment longest setae on segment VIII; anal-lobe
VIII heavily sclerotized. seta near edge of anal tube opening; setae
With 5 oval circuli, present on segments on rest of surface unusually small. Segment
II to VI; middle circulus largest (1 paratype VIII heavily sclerotized.
with middle circulus divided medially to With 4 or 5 oval circuli, present on seg-
form 2 lateral circuli). Labium 3-segment- ments II or III to VI; middle circulus larg-
ed, 84 |x long; paratypes 91(86-99) |jl long. est. Labium about 56 |jl long. Antennae rep-
Antennae represented by unsegmented resented by unsegmented sclerotized knob
sclerotized knob containing 7 or 8 setae. containing 7 or 8 setae. Legs absent; with
Legs absent; usually with small dermal small dermal pockets in position of legs.
pockets in position of legs. Notes: The above description is based on
Notes: The above description is based on 3 specimens.
5 specimens. First instar (gender not determined) (Fig.
Immature female (probably third in- 3): Slide-mounted paratypes 0.5(0.4-0.5)
star)(Fig. 2): Slide-mounted paratypes mm long, 0.3(0.2-0.3) mm wide. Body
mm mm
0.9(0.7-1.1) long, 0.7(0.5-0.9) elongate oval.
wide. Body nearly round, posterior apex Dorsum with conspicuous cerarii on seg-
sclerotized. ments VII and VIII; those on segment VII
Dorsum with anal-lobe cerarii composed each with 2 conical setae, 1 large trilocular
of 1 pair of conical setae. Trilocular and pore, 1 small trilocularpore, and 1 discoidal
discoidal pores absent. Oral-collar tubular pore; those on segment VIII each with 2
ducts of 3 variable sizes, becoming smaller conical setae, 1 small-sized trilocular pore,
anteriorly, abundant over surface except ab- and 1 discoidal pore; 2 thin setae and 1 or
sent from segment VIII. Setae on segment 2 associated trilocular pores on margin of
528 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
^--©^
Fig. 2. Micouicoccus niehsaameiu. Third-instar female
VOLUME NUMBER
101, 3 529
o
.----O
o
o-
Fig. 3. Micoiiicocciis niebsaaineni. First instar.
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530 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
most Other abdominal segments forming in- claw digitules clubbed apically; claw with-
conspicuous cerarii. Trilocular pores of 2 out denticle.
sizes; larger size present in 2 pairs of inter- Notes: The above description is based on
rupted longitudinal lines, 1 along body mar- 4 specimens.
gin and associated with 2 setae on each seg-
Quadrigallicoccus Williams and Miller,
ment except absent from segment VIII, 1 new genus
pair present in mediolateral lines, incom-
plete, present on head to abdominal seg- Type species. Quadrigallicoccus laur-
ment I, and present on any or all of seg- acearum Williams and Miller.
ments VI, VII, or VIII; smaller-sized trilo- Diagnosis. Adultfemale: Body rotund;
culars arranged in 3 pairs ofinterrupted lon- vulva sometimes located at posterior apex
gitudinal lines, 1 along entire body margin, of body on mounted specimens. Cerarii ab-
1 present in mediolateral line on segments sent. Dorsum covered with trilocular pores
where larger size is absent, and 1 present in similar to those of other pseudococcids;
submedial line on head to abdominal seg- venter covered primarily with quadrilocu-
ment I and 1 or 2 on abdomen. Discoidal lars that have same structure as triloculars,
pores absent except in posterior cerarii. but with 4 loculi; quinqueloculars most
Oral-collar tubular ducts absent. Setae ab- abundant near spiracles also similar to tril-
sent from segment VIII except in cerarii; 1 oculars but with 5 loculi. Oral-collartubular
pair ofenlarged setae in medial area ofseg- ducts present. Circulus absent. Anal ring ei-
ment VII; setae arranged in 3 pairs of lon- ther without pores or these very small and
gitudinal lines; marginal line composed of few; anal-ring setae absent. Legs absent.
2 setae which are homologous to cerarii in Spiracles without pores in atrium. Antennae
represented by unsegmented, sclerotized
other mealybugs; mediolateral and subme-
area containing several setae.
dial lines complete except absent from seg-
Description. Adult male: Body elon-
ment VIII; dispersed on head and thorax;
with elongate anal-lobe seta. Segment VIII gate. Penial sheath about Va length of body
with sclerotization on both surfaces form-
sometimes sclerotized.
ing complete capsule. Ventral eyes set on
Anal ring with 15(13-32) pores on each
protuberance; dorsal and ventral eyes each
side of ring; with 6 enlarged, apically blunt
with depression around perimeter and small
setae. Anal tube absent.
sclerotized dimples in lateral area of de-
Venter with trilocular pores present in pression. Body with bristle-shaped setae
mediolateral areas of any or all ofsegments only. Antennae primarily with elongated
IV, V, and VI; also present near spiracles. fleshy setae; capitate setae on 10th antennal
Discoidal pores present in lateral line with segment absent or with very small club.
1 on each side of each abdominal segment, Claw with digitules less than Va length of
also present laterad of each spiracle. Oral- claw; with minute denticle near tip ofclaw.
collar tubular ducts absent. Longest setae Abdominal tergites and sternites heavily
on segment VIII; setae arranged in 3 pairs sclerotized. With unusual minute tubular
of longitudinal lines, submarginal, medio- ducts.
lateral, and submedial. Second-instarfemale: Ostioles restricted
With 2 or 3 oval circuli, present on seg- to anterior pair. Trilocular, quadrilocular,
ments IV and V, sometimes present on seg- and quinquelocular pores with swirled pat-
ment III. Labium 62(57-67) |x long. Anten- tern. Antenna 6-segmented. Oral collar tu-
nae 6-segmented, 128(124-136) |jl long. bular ducts present. Anal ring on dorsal sur-
Legs well developed, hind femur 49 |x long; face removed from apex. Sclerotized slit
hind tibia 35 |x long; hind tarsus 42 |ji long; present laterad of mid pair of legs.
tibia/tarsus 0.8; femur/tibia 1.4; tarsal and Second-instar male: Trilocular, quadril-
VOLUME 101, NUMBER 3 531
Fig. 4. Qiiculrii^allicoccus lauraceanini. Adult female.
