A new genus and species of ant-associated coccid (Hemiptera: Coccidae: Myzolecaniinae) from Canthium Lam. (Rubiaceae)

8 Pages·1996·3.1 MB·English

A NEW GENUS AND SPECIES OFANT-ASSOCIATED COCCID (HEMIPTERA:
COCCIDAE: MYZOLECANIINAE) FROM CANTH1UMLAM. (RUBIACEAE)
PENNYJ. GULLAN ANDAIMORNC. STEWART
Gullan,PJ. & Stewart,A.C, 199607 20: A newgenus and speciesofant-associatedcoccid
(Hemiptera: Coccidac; Myzolecaniinae) from CanthiumLam. (Rubiaceae). Memoirsofthe
QueenslandMuseum39(2): 307-314. Brisbane. ISSN 0079-8835.
The adult female and first-instar nymph of Torarchus endocanthium gen. et sp. nov.
(Hemiptera: Coccidae: Myzolecaniinae) from Queensland, is described. This coccid is
knownonlyfrominsidehollow,swollenstems(antdomatia)ofplantsofthegenusCanthium
Lam.(Rubiaceae),whereit livesasatrophobiontinthenestsofantsofaPodomyrmaspecies
(Formicidae: Myrmicinae).The first-instarnymph hastypicalcoccid features, buttheadult
femaleisunusual,beingdistinguishedwithintheCoccidaebyitsmultiloculardisc-poresand
microducts resembling bilocularpores on the dorsum, and rounded, projecting anal plates
on the anterior edge of a setose bulge. Coccidae, Formicidae, Rubiaceae, ant-plant
association.
Penny J. Gullan & Aimorn C. Stewart, Division of Botany & Zoology, The Australian
National University, Canberra, ACT0200, Australia; received 15December 1995.
A new ant-plant association (Monteith, 1989, insects belong to an undescribed species ofCoc-
1990)frommarginalrainforestareasofcoastal to cidaewith unusual morphology, suggestiveofan
central Queensland involves trees of the Can- obligate relationship with the ants. Individual
thium odoratum-C. buxifolium complex adult coccids have their convex venter closely
(Rubiaceae) and ants of an undescribed fitted into pits gnawed by the ants into the inner
Podomyrma species (Formicidae: Myrmicinae). surface ofthe domatium. Theirdorsum is ridged
The ant colonies occur within specialised swell-
and covered in setae, and theiranal area appears
ings in the living stems (ant domatia) (Fig. 1), modifed to facilitate honeydew removalby ants.
whichtheantshollowoutby perforatingthe wall
andremovingthepith.Theantsneveroccuraway Theeversibleanaltubeisstronglydevelopedand
surroundedbyapairofrounded,dorsallyproject-
fromtheirhosttrees; likewise,thetreesare rarely
found without these ants. Monteith and co- ing anal plates so that the whole complex forms
worker Paul Flowerestablished that the ants ob- a prominent mound. These coccids have been
tained their nutrition largely from scale insects, collectedonlyfromantchambersinsidethestems
also called coccoids (Hemiptera: Coccoidea), ofCanthium odoratum (Forsterf.) Seemann and
livingwithintheantdomatia(Fig. 1).Thefeeding may be dependent on this ant-plant association,
stylets of these coccoids presumably tap the
phloem of the host plant and the coccoids* ex-
creta, called honeydew, is consumed by the ants.
It is not known whether the ants ever eat the
coccoids.
These coccoids of Canthium belong to two
families - the Pseudococcidae (mealybugs) and
the Coccidac (soft scale insects or coccids). The
mealybugs are Pseudococcus longispinus (Tar-
gioni Tozzetti) and two undescribed, closely re-
lated species, probably belonging to Crisicoccus
Ferris or Paracoccus Ezzat & McConncll. The
undescribed mealybug species might be specific
to the Podomyrma-Canthium association al-
though they belong to a group with no known
association with ants, whereas P. longispinus is FIG. Sectionedhollow stemofCanthiumodoratum
cosmopolitan, polyphagous, and one ofthe most show1i.ngantworkersand larvae(Podomyrmasp.)on
p1e9s8t5i;feWriolulsiammesal&ybWuagtssoinn, A1u9s8t8r)a.liTahe(Wsiofltlisacmasl,e tthheiurmigohtnatnhde laenft.adult female ofTorarchusendocan-
308 MEMOIRS OFTHEQUEENSLAND MUSEUM
although they are not found in all plants that METHODS ANDABBREVIATIONS
housecolonies ofPodomyrma. Todate, the coc-
cids have been collected only in SEQ coastal Terminology follows Hodgson (1994, 1995).
areas and one more inland site, although the ant- To prepare adult females and nymphs as micro-
plantassociationismorewidespread.Wherecoc- scope slide-mounts, body contents were cleared
cids are absent, the domatia house mealybugs, in cold 10% w/v potassium hydroxide (KOH)
often in large numbers and sometimes of more solutionovernight,thecuticlewasstainedinacid
than oneofthe above species. Inolderstems, the fuchsin in acid alcohol, dehydrated in 3 changes
mealybugs mostly reside in the ant-gnawed pits ofabsoluteethanoland 1 ofabsolutepropan-2-ol
but may feed anywhere within younger stems. and then placed in 3 changes of xylene prior to
Sometimes domatia house both coccids and mounting in Canada balsam. Scale insects were
mealybugs. prepared for scanning electron microscopy
Scale insects have been reported previously (SEM) after preservation and storage in 80%
from inside the swollen stems of Rubiaceae in ethanol. Each specimen was dehydrated in a
Africa (Bequaert, 1922). Those collected with graded ethanol series, dewaxed in xylene,
mCirceimnaateo)gainsstiedrelhaoulrleonwtisFtoermeslo(fFoPrsmyidcriadxaes:ubMcyorr-- drieshtyidlrleadtewdatetrh,ropuogsth-faixgerdadined1%etahaqnuoelousseroisesmiinutmo
ciata (DC.) Bridson (formerly Piectronia tetroxide, washed indistilledwaterandsonicated
laurentiiDeWild.) fromZaire weredescribed as briefly lo remove any black precipitate, critical
Hemilecanium recurvatum by Newstead (1910). point dried, glued onto a metal stub with nail
This species is morphologically very different varnish and coated with gold palladium under
from the coccid from Canthium in Queensland vacuum. Specimens were then examined and
(types of//, recurvatum (BMNH) have been ex- photographed usinga Cambridge S360 SEM.
amined) and belongs to a different subfamily. In Abbreviations used for the depositories are:
Australia, no coccids or mealybugs have been ANIC, Australian National Insect Collection,
reported previously from Canthium, although CSIRO, Canberra; BMNH, The Natural History
species of Myzolecanium Beccari (formerly Museum, London; QM, Queensland Museum,
placed in Cryptostigma Ferris; see Qin and Gul- Brisbane. Each listed scale insect is mounted on
lan, 1989; Gullan, Buckley & Ward, 1993) and a separate microscope slide, unless otherwise
Alecanopsis Cockerell (Green, 1924) have been specified.
described from ant nests in living hollow stems
SYSTEMATICS
of other plants. Alecanopsis and Myzolecanium
belongtothesamesubfamily asthecoccids from Torarchus gen. nov.
Canthiumbutareverydifferentmorphologically.
TYPE SPECIES. Torarchusendocanthium sp. nov.
This paper describes the coccid from Can-
thium. Featuresofthe adult female place it in the DIAGNOSIS. Adult female with rounded-elon-
aMyczooclceicdaniniihnaaveinHgodbgostohn,mu1l9t9i4l.ocIutliasratdyipsicc-aplorfeosr gate elevations on dorsum in a definite arrange-
ment; dorsal cuticle with both multilocular
and microductsresembling bilocularporesonthe
dorsum, and in having anal plates which are very disc-pores and microducts resembling bilocular
pores; anal plates rounded and projecting,
rounded and dorsally projecting, together form-
winhgeanmthoeupnldatewshaernectlhoeseadn)a.lItnumboesitsortehterracctoecdci(di.se,. Fsiirtsuta-tiendstoanranntyemrpiohreodfgetyopficraalisceodcacriedaofforcmut,icblue.t
each anal plate is triangular, posteriorly directed with single stout seta per stigmatic cleft. All
and lies level with the dorsal surface when the stages living in hollow stems of host plant at-
anusisretracted. Furthermore,inthisnewspecies tended by ants.
the anal area is located on the anterior edge ofa
prominentbulge whichbears very long setae in a DESCRIPTION. Features believed to be
centraldepression.Thefirst-instarnymphismore taxonomically significant at generic level are
typically coccid-like (Miller, 1991), except that highlighted. Thespeciesdescriptionprovidesthe
each stigmatic cleft has only a single stout stig- best summary ofthis monotypic taxon.
matic seta. It differs in this regard from known Adult female broader than long, with a row of
first-instarnymphsofotherMyzolecaniinae(Ray rounded-elongateelevationsorridgesdorsallyon
& Williams, 1980; Qin & Gullan, 1989; Sheffer each side ofmidline, these become less apparent
&
Williams, 1990). after slide-mounting; setae flagellate, clustered
NEWANT-ASSOCIATEDCOCCIDIN CANTHWM 309
FIG. 2. SEMs ofcuticularfeatures ofTorarchusendocanthium: A, anal area ofadult 9, showingroundedanal
platesandlongsetaeonraisedareabehindanus(scale50 p,m); B,enlargementofanalareaofanotheradult 9,
showing anal plates and partially everted anus (scale 50 p,m); C, multilocular disc-pores and openings of
microducts on dorsolateral surfaceofadult 9(scale 1 u-m); D,dorsal view ofanal areaoffirst-instarnymph,
showing anal platesand rugosecuticle (scale 25 p,m).
dorsally on ridges, in irregular double row mar- tered ventrally; eyespots absent; legs and anten-
ginally and sparsely scattered ventrally, but with nae reduced.
agroupofvery long (1.5-2.5 times length ofanal First-instar nymph oval; derm with rugose
plates), robust setae posterior to anal plates on sculpturing; dorsum lacking setae, pores, ducts
raisedarea; dorsal poresof3 kinds: 1,disc-pores andtubercles;analplateselongatetriangularwith
with 4-8 loculi, 2, microducts which resemble roundedangles;marginwithrowofslendersetae,
bilocular pores when viewed end-on in light each stigmatic cleft with single stout stigmatic
microscope (Fig. 3h) and 3, simple pores; dorsal seta; venter with few setae on head and thorax,
tubular ducts and dorsal tubercles absent; anal short setae in 2 submarginal rows on abdomen,
plates projecting and lobe-like with cluster of longersetae in 2 submedial longitudinalrowson
apical setae; spiracles much larger than legs; abdomen; antennae 6-segmented; legs well
spiracular disc-pores present in bands between developed, fore legs each with single digitule.
body margin and each spiracle, with 5 loculi;
pregenital disc-pores with 5 or 6 loculi; ventral COMMENTS. This genus shares several fea-
tubular ducts absent but small microducts scat- tures with others ofthe Myzolecaniinae; for ex-
310 MEMOIRS OFTHEQUEENSLANDMUSEUM
ample, Cyclolecanium Morrison, Megasaissetia DESCRIPTION. Adult 9 (10 specimens
Cockerell, Neolecanium Parrott and Pseudo- measured). Live material. Body ofyoung 9 yel-
philippia Cockerell also possess some form of low, with sparse coating ofwhite, powdery wax.
dorsal pore or microduct with a bilocular ap-
pearance and that ofN. imbricatum (Cockerell) Mounted material. Body transversely oval,
even has an inner duct (Hodgson, 1994) similar rounded on each side, 1.3-1.8 times wider than
to that of T. endocanthium. The dorsal ridges of long, rather flattened but with elevated areas on
T.endocanthiumareofsimilarshapebutdifferent dorsum;stigmaticcleftsshallow;analcleftfused.
number and arrangement from those of Coccus Length 1.5-2.6 mm, width 2.3-4.4 mm.
tumuliferusMorrison (Morrison 1921), which is
notatruespeciesofCoccusLinnaeus (Coccinae) Dorsum. Derm membranous except for narrow
but undoubtedly a member of the crescentofsclerotisation around anteriormargin
Myzolecaniinae. Furthermore, the anal plates of ofanalplatesand lightsclerotisationatmargin in
C. tumuliferus are ofsomewhat similar shape to stigmatic cleft; with 7 radial ridges dorsally on
those of T. endocanthium but with different ar- eachsideofbody,forminganelevatedsubmedial
rangement and number of setae; however, the area following curve of margin, with anterior-
dorsal pores and setae and ventral ducts of C. most and posteriormost ridges smallest; a
tumuliferus are very different from those of T. rounded raised area lies posterior to anal plates
endocanthium. The presence of multilocular
(Fig. 2A) with central depression marking posi-
disc-pores and microducts resembling bilocular tion ofanal cleft. Dorsal setae flagellate, 15-100
pores on the dorsum of Torarchus easily distin- |xm long in submarginal areas and around anal
guishes it fromAkermes Cockerell, Alecanopsis plates, longer (70-150 u.m) medially and
andMyzolecanium, whichare theothergeneraof clustered on ridges; a group of 13-26 very long
the Myzolecaniinae found in Australia; further- (230-500 u-m), robust setae posterior to anal
vmeorrye,shiatldliofwfesrtsigfmraotmicMcylzefotlseacnadnifurmominAkhearvmiensg pmlualtteisloocnulraarisdeidsca-repao.reDsor5s.a5l-7p.o5reus-mofin3 dkiinadmse:te1r,
and Alecanopsis in having no stigmatic setae. with 4-8 (mostly 6, Figs 2C, 3i) loculi, scattered
Torarchuskcysoutin Hodgson(1994,pp.91-92) in marginal and submarginal area; 2, microducts
tothecoupletcontainingAkermesandAlecanop- resemblingbilocularpores(Figs2C,3h), 3-4 u-m
sis, but does not fitthe description ofeither. in greatest dimension, densely distributed over
ETYMOLOGY.Latintorus,roundelevationorbulge, entire dorsum; 3, simple pores (Fig. 30 3-4 u-m
Greek archos, anus; refers to the shape of the plates in diameter, scatteredoverdorsum. Preopercular
surroundingthe anusand locationoftheanal areaona poresabsent. Dorsal tubularductsabsent. Dorsal
cuticularbulge. tubercles absent. Anal plates lobe-iike, rounded
apically (Figs 2A, 2B, 3d), each 150-210 fim
Torarchusendocanthium sp. nov. long, 1 10-150u-mwidewhenmeasuredinnatural
(Figs 1-4) position (distortion during mounting common);
with 5-6 small setae apically on each plate. Ano-
MATERIAL EXAMINED. HOLOTYPE, genital fold with 2 pairs of setae in hypopygial
QMT13986, adult 9,Qld, Mt Crosby, offCrosby Rd position, a pair of larger setae at each corner of
&BunyaSt.,27°32'S, 152°48'E,9.iv.1989,P.Flower. anterior margin and 1 pair laterally. Anal ring
PARATYPESadult 9 9 only,QLD:7adult 9 9,same 58-68 u-m in diameter, probably with 6 pairs of
data as holotype; 1 adult 9, 2 slides of first-instar setae, 65-1 15 u.m long, ratherflattened (Fig. 3e),
nymphs, Mt Crosby, 18.viii.1988, P. Rower; 1 adult difficult to see when retracted inside anal tube
9, 1 slideoffirst-instarnymphs,MtMoffattNat. Park, (Fig. 2B).
Kenniffs Lookout, 24° 55'S, 147° 59'E, 13.xii.1987,
G. Monteith,G. Thompson & D. Ycates; 7 adult 9 9,
3 immature 9 9, 6 slides of first-instar nymphs. Margin. Marginal setae flagellate (Fig. 3g), 25-
AuburnGorge,SW Mundubbera,25°43'S, 151°03'E, 150 u-m long, in irregular double row around
lateMarch 1989,G. Monteith; 3adult 9 9, Keysland, entire margin except absent in stigmatic clefts
20km NWofWondai, lateMarch 1989,G. Monteith. and posteriorly where fused anal cleftjoins mar-
Locationofparatypes: 4adult 9 9 and 2 slides gin. Stigmaticcleftsveryshallowwith smallarea
of first-instar nymphs in ANIC, 1 adult 9 in of light sclerotisation; lacking stigmatic setae.
BMNH, remainder in QM (QMT26022-26035). Eyespots apparently absent.
DIAGNOSIS. As forgenus. Venter. Dermmembranous;onlyabdominal seg
NEW ANT-ASSOCIATEDCOCCID IN CANTHIUM 311
FIG.3.Adult 9 ofTorarchuscndocanthtum. Enlargements: A,quinqueloculardisc-porefromstigmaticfurrow.
B,ventralmicroduct.C,pregenitaldisc-pore with6loculi. D,anal lobesvieweddorsally(onleft)withanalring
and anal ring setae indicated by dashed lines, and ventrally (on right) showing setae and supporting bar of
ano-genital fold. E, anal ring seta. F, simple pore. G. marginal setae. H, lateral and end-on views ofdorsal
microduct. I, multiloculardisc-pore.
mentation discernible. Ventral setae flagellate, conspicuous innermost end, each microduct ap-
15-75 u.m long, sparsely scattered. Pregenital pearingasasmallslightlyovalporewhenviewed
disc-pores(Fig. 3c)8-1 u.mindiameterwith5-7 end-on, scattered throughout venter. Ventral
(mostly 5-6) loculi, distributed around vulva on tubular ducts absent. Spiracles well developed,
posteriorsegmentsandin an irregularlinereach- withconspicuousmuscleplatetoeachperitreme;
ing to metathoracic spiracle of each side. Stig- anterior spiracle plus peritreme 150-200 u-m
matic furrows each with quinquelocular long, 1 10-135 u-m wide; posterior spiracle plus
disc-pores(Fig.3a),5-6|xmindiameter,inaband peritreme 150-200 u-m long, 120-140 u-m wide.
from margin to spiracle. No preantennal pores Legs reduced, each 90-130 u,m long, trochanter
present. Ventral microducts (Fig. 3b) each with fused withfemur, fusionoftibiaandtarsuspartial
outerductule4-5 u.m long and innerductule with to complete; each claw small, without denticle;
312 MEMOIRS OFTHEQUEENSLANDMUSEUM
FIG. 4. First-instar nymph of Torarchus endocanthium, with anus everted. Enlargements: A, quinquelocular
disc-pore from stigmatic furrows. B, stigmatic area with stigmatic seta in cleft, 2 marginal setae and quin-
queloculardisc-pores. C, apex ofhind leg showingdigitules. D,dorsal (on left)andventral (onright)viewsof
anal plates. E, anal ring with setal positions indicated by blackened spots; F, dorsal view of anal lobes and
evertedanal ring; G, marginal seta.
claw digitules longer than claw but shorter and First-instar nymph (10 specimens measured).
broader than tarsal digitules; tarsal digitules 22- Mounted material. Bodyoval,463-560 jxmlong,
25 (xm long. Antennae reduced, with 5-6 seg- 270-315 fim wide. Derm membranous
mentsatmost,segmentationoftenindistinct;total throughout but with rugose sculpturing dorsally
length 95-150 |xm; with fleshy setae 13-25 jxm and ventrally, most clearly visible in SEM (Fig.
long on apical segments and flagellate setae 15- 2D). Segmentation not readily apparent (but ob-
65 (xm long on all segments. Clypeolabral shield vious in whole nymphs underSEM).
350-390|xm long, 270-320|xm wide. Labium 1-
segmented, 140-150 \xm long, c.150 \xm wide.
Dorsum. Setae,pores,ductsandtuberclesabsent.
NEW ANT-ASSOCIATEDCOCCID IN CANTHWM 313
Anal plates (Fig. 4d) elongate triangular with 30-36 \xm long, 45-50 fim wide, with 4 pairs of
roundedangles,40-48 \xm long, 15-24 \xm wide; setae; stylets looped, total length 500-660 jim.
anterolateral margin 22-26 jxm long,
posterolateral margin25-30 p,mlong; dorsal sur- COMMENTS. The first-instar nymphs of most
face with scattered microspines. Each plate with Coccidaepossess 3 stoutstigmatic setaeperstig-
4 dorsal setae, 3 on apex of plate, 1 on mesal matic cleft. The nymph of T. endocanthium has
margin; mediansetaonapexrobust, 190-225 p.m only one differentiated stigmatic seta per stig-
long, abouthalflength ofbody. Ano-genital fold matic cleft (presumably homologous with the
with 1 pairofanteriormargin setae and 1 pairof median stigmatic seta of other coccids). The 2
lateral margin setae. Anal ring (Fig. 4e) ap- slender setae that border each stigmatic seta in
proximatelycircular, 18-22 jim in diameter, with nymphs of T. endocanthium appear identical to
about 14 irregularly shaped pores and 6 setae, the remainder of the marginal setae but may be
45-68p,mlong,with2setaedistinctlyshorterand homologous with the 2 stout lateral stigmatic
more slender than other4 (Fig. 40- setaethatnormallyaccompanythelongermedian
stigmatic seta in othercoccids.
Margin. Marginal setae(Fig. 4g) 10-15 p.m long, ETYMOLOGY. Greek endon, within or inside, with
slender,taperingtoapoint,usuallycurved,point- the name ofthe host plant Canthium.
ing in posterior direction, distributed as follows:
8 around head between anterior stigmatic clefts, ACKNOWLEDGEMENTS
4betweeneachpairofstigmaticclefts,and 18on
abdomen. Stigmaticclefts moderatelydeveloped We thank Geoff Monteith, Queensland
(Fig. 4b), each with single stout stigmatic seta
10-16 y.m long, bordered by 2 slender marginal Museum for drawing our attention to this scale
sseettaa.e,Ey1easnptoetrsioprreasnedntIjpuossttaebrioovretloeevealchofsatnitgemnantailc iWnesecatc,kfnoorwlloeadngeofthtehefascpileictiiemseannsdatnedchfnoircaFligs.up1-.
port of the Electron Microscopy Unit, ANU.
scape.
Chris Hodgson, Geoff Monteith, Katie Strong
and Doug Williams made helpful comments on
Venter. Ventral body setae slender, of2 lengths: the manuscript. RobertHoareadvised onthe use
ofLatinandGreektoformthenewnames. Diane
submarginal setae short, 2-5 fim long, in 2 lon- Bridson provided the name of the African
gitudinalrowseachof7oneachsideofabdomen
Rubiaceae.
(2 pairs of setae per segment), 1 seta between
anterior and posterior stigmatic clefts and pair
1 LITERATURECITED
at apex ofhead; submedial setae 20-45 urn long,
in 2 longitudinal rows of6 each on abdomen (1
ppaaiirrbpeertwseeegmneanntt)e,nnseatea.eSlpoinrgaecrlepsosptleursioprelryi,tarenmdes1 BEQUthAeEpRlTan,twJ.or1l9d2.2.BuAlnlettsininotfhtehierAdimveerriscearnelMautisonesutmo
ofNatural History45: 333-584.
15-21 u-mlong,6-1 |xmwide.Stigmaticfurrows GREEN, E.E. 1924. On some newspeciesofCoccidae
each with quinquelocular disc-pores (Fig. 4a),
from various sources. Bulletin of Entomological
2-3 jim in diameter, in single row; anterior Research 15:41-48.
spiracular disc-pore bands each with 4-6 pores; GULLAN,P.J.,BUCKLEY,R.C.&WARD,P.S. 1993.
posterior spiracular disc-pore bands each with Ant-tended scale insects (Hemiptera: Coccidae:
4-5 pores. Other pores and ducts absent. Legs Myzolecanium)withinlowlandrainforesttreesin
well developed, 210-245 u-m long, without Papua New Guinea. Journal ofTropical Ecology
tibiotarsal sclerotisation or free articulation; 1 or 9:81-91.
afewflagellatesetaeoneachsegment;2knobbed HODGSON, C.J. 1994. 'TheScale InsectFamily Coc-
claw digitules (Fig. 4c) per leg; 2 knobbed tarsal cidae:AnIdentificationManualtoGenera.' (CAB
digitules (Fig. 4c) per leg, except fore legs each International: Wallingford)639p.
with only I digitule; tarsal claw with small den- 199o5f.aOdbusletrfveamtailoensCooncctihdeaset.ruIcstruareeloJfoutrhneaslpiorfacElne-s
ticle. Antennae well developed, 6-segmented, tomology 29: 47-55.
130-160u-mlong;segmentIIIlongest,33-45fjtm; MILLER, D.R. 1991. Superfamily Coccoidea. pp. 90-
5 fleshy setae (1 on IV, 1 on V, 3 on VI), 13-25 107. In Stehr, F.W. (ed.) 'Immature insects.
p,m long, and about 15 hair-like setae, 10-53 jxm Volume2/ (Kendall/Hunt: Iowa).
long. Mouthparts with clypeolabral shield 83-90 MONTEITH, G.B. 1989. Ant symbiosis in the plant
jim long, 63-77 \xm wide; labium 1-segmented, genusCanthium inQueensland.NewsBulletinof
314 MEMOIRS OFTHEQUEENSLANDMUSEUM
the Entomological Society ofQueensland 17(3): philippia quaintancii (Homoptera: Coccoidea:
31-32. Coccidae). Annals ofthe Entomological Society
1990. The plant-antconnection. Wildlife Australia ofAmerica73: 437-447.
27: 6. SHEFFER, B.J. & WILLIAMS, M.L. 1990. Descrip-
MORRISON, H. 1921. Some nondiaspine Coccidae tions, distribution, andhost-plantrecordsofeight
from the Malay Peninsula, with descriptions of firstinstarsin me s joumeyella(Homoptera:
apparentlynewspecies.ThePhilippineJournalof Coccidae). Proceedings of the Entomological
QNiINENtW.WSTSCMocT.ciKEtce.inAA&dcUDaeG,eU1.KRL8J:Lo6A1ui3r9Ny7n1i.-a0uPl67.oOu7Jfnn..Ettc1wwo9oon89onn.meeiCwwcryssBpppileeoccoliisoeetssgiygoomrf5a:AmfF1ner8rci-rc2aia2snn.: WILL(Sc.IBo_rAcliM.teiS.tsy,,h'ofMcD.u.W.J.as,.sehu.1i.9mn8g5t(,..oN..na't9nAu2our(/sa1Itl).Xr:aH.l4A.ii.4Asa-t_n5Co7rn.yM)e.:alL.yJobnudg,osP'n.).
acoccoidgenuswithastrikinglydisjunctdistribu- 431p.
tion (Homoptera: Coccidae) Systematic En- WILLIAMS,D.J. &WATSON,G.W. 1988. Thescale
tomology 14: 221-232. insectsofthetropical SouthPacificregion.Part2.
RAY,C.H.,Jr. &WILLIAMS,M.L. 1980. Description TheMealybugs(Pseudococcidae)'.(C.A.B.Inter-
ofthe immaturestages and adult maleofPseudo- national: Wallingford)260p.