A complex of cryptic species in the genus Coptodactyla Burmeister (Coleoptera: Scarabaeidae: Coprini)

21 Pages·2000·9.4 MB·English

A COMPLEX OF CRYPTIC SPECIES IN THE GENUS COPTODACTYLA BURMEISTER
(COLEOPTERA: SCARABAEIDAE: COPRINI)
C.A.M. REID
Reid, C.A.M. 2000 12 31: A complex of cryptic species in the genus Coptodactyla
Burmeister (Coleoptera: Scarabaeidae: Coprini). Memoirs ofthe Queensland Museum
46(1): 231-251. Brisbane. ISSN 0079-8835.
Thesubaeneaspecies-complexinthecoprinegenusCoptodactylaBurmeisterisdescribed,
comprising C. nitida Paulian, C papua Lansbergc, C. subaenea Harold and three new
species, C. matthewsi and C. storeyi, from North Queensland, and C. merdeka from New
Guinea. Coptodactylanitida Paulian isremoved from synonymywith C. subaenea. These
externally almost identical species are primarily discriminated using sexual characters,
namely the parameres, endophallus and vulvar sclerites. A phylogenetic analysis ofthe
speciescomplex isgiven. Fiveofthespeciesareallopatric withrespecttoeachother. Two
sisterspeciesarewidespread butallopatric intheWetTropicsofNorthQueensland,where
they meet at the Black Mountain Barrier, a known dry climate barrier at past periods of
maximum aridity. A key is given for the 17 species ofCoptodactyla and lectotypes are
designated for C. papua and C. subaenea. Coleoptera, Scarabaeidae. Coptodactyla.
Australia, New Guinea, taxonomy, biogeography.
ChristopherAthol McEwan Reid. Co-operative Research Centrefor Tropical Rainforest
Ecology & Management. James Cook University. Smithfield 4878. Australia (current
address:CentreforBiodiversityandConservationResearch,AustralianMuseum, 6College
Street. Sydney2000. Australia); received31 March2000.
This paper revises a species-complex in the comprehensively revised by Matthews, using
dung-beetle genus Coptodactyla Burmeister, malecharacters(Matthews, 1976). Hehadmuch
which is abundant in the closed forests ofnorth lessmaterialavailablethanthepresentauthorand
Queensland. The project is one of several did not examine the female genitaliaorthe male
initiatedbytheCo-operativeResearchCentrefor endophallus; examinationofthese structureshas
Tropical Rainforest Ecology and Management. allowed more detailed discrimination ofspecies
Cairns, whichaim toprovideasoundtaxonomic and populations. This study has revealed six
and phylogenetic framework for the more sibling species confused under the names C.
significantgeneraoftheQueenslandWetTropics papua Lansberge, C. subaenea Harold and C.
WorldHeritageArea.Thissurveyofscarabaeine nitida Paulian. No other taxonomic changes are
distributionandsystematicshas led tostudies of necessary in thegenus.
dwiivtehrisnittyhe(MWoentteTirtoh,pic19s9(5R)eaidn,dCarraeansrteolnat&ionRsehiidp,s MATERIALS AND METHODS
in press). Male genitalia were immersed in dilute KOH
Such a project is feasible primarily thanks to for several hours, then transferred to water. The
the sound taxonomic base provided by Eric parameres were removed from the basal piece,
Matthews 27 ago (1972, 1974, 1976) and the and the penis and endophallus extracted. The
subsequenteffortsofprojectcollaboratorsGeoff penis was separated from the endophallus by
Monteith(QueenslandMuseum,Brisbane),Ross cuttingthethinconnectingmembraneattheapex
Storey (Department of Primary Industries, ofthepenis. The endophallic lobes were everted
Mareeba) and Tom Weir (Australian National with fine forceps. The ejaculatory sac was not
Insect Collection, Canberra), who have collect- invertedasthescleritesweremoreclearlyvisible
ed,curatedandsorted40,000specimensofabout in the relaxed condition.
190 species (G. Monteith, pers. comm.) ofWet Female genitalia, including the vulvar
Tropics scarabaeines. Their collective resource sclerites, were removed from the apical
provides an excellent platform for revisionary abdominal ventrite.softenedindiluteKOH,then
taxonomic studies. water, and cleaned. The spermatheca was
CoptodactylaBurmeisterisagenusofcoprine examined in glycerol.
scarabs (Montreuil, 1998) confined to Australia Terminologylargelyfollows Mathews(1976),
and New Guinea. The Australian species were with the following additions. Scarabaeines
232 MEMOIRS OF THE QUEENSLAND MUSEUM
8255 18.SB CSENT
FIG. 1. CoptodactylasubaeneaHarold, 6,showingdistributionofannularpunctures.A,dorsum;B,venter;C,
detail ofannularpuncturesanteriortoeye. Scalebar=2mm.
commonly have patches of large annular which may be furthersubdivided, branching off
punctures(Krikken, 1977)andthedistributionof the base ofthe ejaculatory sac.
these is important for discriminating species in
Coptodactyla. These punctures are deep (Fig. The female genitalia of Coptodactyla are
1C), enclose setae, and may function as fungal unusual amongst Scarabaeinae in having
storagepitsforinoculatinglarvalfood.Anomen- extensive secondary sclerotisation around the
pcilrnaoatpvupirrdeoeopdfrbimyaatlGeeenefionerdropC(ho1a9pl9tl6oi)dc,asbccutltyerltihatiesssapipenpcCeioaeprsrs.itnToihbiees tvahuneldvvaau.slyvTmahmreestacrlreiercaiatileml(myFeidfgoi.la9dtBeed)l.,ywBaheritocwuhenedmnathytehbeveuvlluvolabveaidrs
egnednoeprhaaClolupstoodfaaclltyAlusatarnaldiTahnysrpeegciisesBolfacthkebucropnrhianse sscclleerriotteisaedndtrtahnesvaernsuesbatrh,erteheidsourssaulalslcyleraitse,trwohnigclhy
maybederivedfromfusionofthehemisternites.
been examined. There are always three basal
sclerites (Fig. 5B), in an ejaculatory sac, which Seventeenhundredand fifty specimens ofthe
form the sperm pump when everted: the subaenea-species complex were examined.
flagellum, relatively short and stout and without Abbreviations for depositories are as follows:
divergent apices (flagella of Genier); the basal ANIC, Australian National Insect Collection,
sclerite, afoldedplate adjacent tothe apical half Canberra; BMNH, Natural History Museum,
oftheflagellum;theringsclerite,atorbeyondthe London; CMN, Canadian Museum for Nature,
tip ofthe flagellum. Otherareasofsclerotisation Ottawa; MCG, Museo Civici, Genoa; MZB,
maybepresent(asinFig.5B),butwhenobserved Museum Zoologicum Bogoriense, Bogor, Indo-
through the invertedwall are poorlydefined and nesia; DPIM,QueenslandDepartmentofPrimary
generally only lightly sclerotised. In Copto- Industry, Mareeba; QM, Queensland Museum,
dactyla there is always a lateral sac (Fig. 5B), Brisbane. The latitude and longitude for
CRYPTIC SPECIES OF COPTODACTYLA 233
Australian localities included in thisrevision are distributed as follows (Fig. 1): sides of
available fromthe Collection Manager,ANIC. frontoclypeus anterior to eyes; inner margin of
Abbreviationsforcollectors'nameswithmore eyes; lateral pronotal pit; posterior margin of
than one entry are as follows: GB, G. Bornem- pronotum; posterior ofpresternum; posterior of
issza; EB, E.B. Britton; JGB, J.G. Brooks;DC, hypomeron internal to lateral groove; base of
D.J. Cook; JF, J. Feehan; PF, P. Ferrar; HAH, H. each stria; mesosternum; mesepimeron; sides
& A. Howden; SM. S. Misko; GM,G.Monteith; and anterior of metasternum; metepistemum;
GSM, G. & S. Monteith; CR, C.A.M. Reid; RS. hind margin ofprofemur; meso-and metacoxae;
R.I. Storey;GT,Gl.Thompson;AWH,A.Walford- row across base of each ventrite except last;
Huggins; TW, T.A. Weir; DY, D. Yeates; PZ, P. lateral angles ofpygidium.
Zborowski. Other abbreviations: Bch, Beach; C, Head{V\gs 1, 2A-B). Dorsum without ridges or
Cape; Ck, Creek; I., Island; Mt. Mount/Mountain; horns in either sex; fronto-clypeal margin with
NP,NationalPark;Pt,Point; Ra., Range; R.,River; slight notch at junction of gena, produced
Rd, Road; Tbld, Tableland. anteriorly as two upturned triangular lobes with
Descriptions are based on freshly emerged median notch; downturned apex of clypeus
specimens, if available. Older specimens may almost triangular, with marginal arcuate groove
have densely scratched and dull dorsal surfaces, formostofwidth(Fig. IB);lateral (genal)angles
and the oldest individuals have their protibiae blunt, sides contracted in straight line towards
reduced to pointed stumps and fronloclypeal eyes; eye divided by narrow canthus for halfof
anteriormargins eroded. lateral length; eyes separated by 4-6 dorsal eye
widths; most of anterior of frontoclypeus
Coptodactyla Burmeister, 1846 transversely rugose (wrinkled), head between
eyesvariablypunctured,convex;antenna:length
TYPE SPECIES. Copris glahricolUs Hope, 1842, by segment 1 at least equal to 2-6 combined, 2
memorypy(Matthews 1976). elongate globular, 3 triangular, 3-6 increasing in
DISTRIBUTION.Australia(coastalQueensland width, 4-6 decreasing in length, width 6 twice
and Northern Territory) andNew Guinea. length, 7-9 forming lamellate club, 7 shallowly
excavate at base to house 6, 7-9 decreasing in
REMARKS. The genus was comprehensively width; last segment ofmaxillary palp fusiform,
described and its synonymy was discussed by length 2-2.5x penultimate segment.
Matthews (1976). The newspecies added below Thorax (Figs 1, 2A-B). Pronotum: anterior
do not require any alteration to this description. evenly convex, without excavation or tubercles,
Hitherto two species of this genus have been base with faint median longitudinal impression;
distinguishedbythe incomplete border(beading sides of pronotum almost straight to shallowly
sensu Matthews) of the anterior edge of the curved; uppersurface almost entirely punctured,
pronotum. Here fouradditional specieswiththis more strongly at sides; anterior edge without
character-statearedescribedandthewholegroup border for middle eighth to third; two circular
is placed inthesubaenea species-complex. lateral median depressions present, connected to
anterior angles by shallowly convex and often
Coptodactyla subaenea species-complex
effacedridge;hypomeronwithlongsubmarginal
DIAGNOSIS. Anterior border of pronotum ridge c.0.75 length from base towards anterior
medially effaced(Figs 1A, 2A-B); striae 8 and 9 excavation, impunctate except annular punc-
complete to elytral base; males without tures; scutellum minute, size equal to pronotal
secondary sexual modifications of head, sensorypit;elytrawith9distinctdeeplygrooved,
pronotum or legs (Fig. 1); apices ofparameres but feebly punctured (slightly foveolate), striae
strongly deflexed (Fig. 2C-F); endophallus with complete to base, stria 10 along edge of
one lateral sac (Fig. 5B). epipleuron, stria 9 fused to 10 one third from
base; interstices flat except apical halfofsuture
DESCRIPTION. The following attributes are slightlyconvexandouteredgeofelytronconvex
sharedbyallspeciesofthiscomplexandwill not between striae 8 and 9, but without pseude-
be repeated in the species descriptions (but note pipleuron; meso- and metasterna completely
thatsomefeaturesmaybeobliteratedorobscured fused;middleofmetasternumshining,finelyand
in worn specimens). sparsely punctured; wings fully developed; mid
Length 8-13.5mm, but rarely <9.5mm. An- andhindtrochanterswithsmalltuftofdensestilf
nular punctures, either circular or semicircular. golden setae (may be abraded); male without
234 MEMOIRS OFTHE QUEENSLAND MUSEUM
V3MH^^^^^. ^^k.
'Us. '' '
15KU X65 826? lee.eu csent
FIG2.Coptodactylaspecies.A-B,SEMleftsideheadcapsule:A,matlhewsi(IronRange);B,nitida(Broadwater
Park);C-E,SEMapexofparameres:C,matthewsi(IronRange);D,nitida(CardwellRange);E,storeyi(Mount
Webb); F,subaenea(McllwraithRange).
secondarysexualcharactersonlegs,femorawith Male.Pygidiumlarge,lessthantwiceasbroadas
complete posterior marginal ridge, without long (Fig. 7A), and apex last ventrite anteriorly
median spuron foretibia. excavate (Fig. 1B); genitalia: penis reduced to a
deeply bilobed weakly sclerotised sheath (Fig.
CRYPTIC SPECIES OF COPTODACTYLA 235
FIG. 3. Coptodactyla species, parameres. lateral, and apical: A, matthewsi; B. merdeka; C, nitida (Baldy
Mountain); D. nitida(Paluma); E.papua(paralectotype). Alltosamescale.
5A); parameres (Figs 2C-F, 3.4) symmetrical or Female. Pygidiumsmall,morethan2X asbroad
almost so (rare individuals are asymmetric. Fig. as long(Fig. 7B), internalmarginofapexdeeply
3D),apicesreflexedand flattened, usuallywitha toshallowlyexcavate(Fig. 7C-II);apicalmargin
basal spur or tubercle; endophallus (Figs 5-6) oflast ventrite evenly convex; genitalia: vulvar
withasinglelarge lateral sacwithpatchofdense sclerites (Figs 8-9) either symmetrical or asym-
stiffspinules and usually a strongly sclerotised metric,usuallywithbroaddorsalplate:spermatheca
spine or bar; flagellum slightly curved; basal approximately C-shaped, with weakly
sclerite elongate rectangular, with various folds sclerotised 'window' at innerangle (Fig. 10).
(almost identical in C. nitida, C. storeyi and C.
subaenea):ringscleritewithtwolobes,onemore REMARKS. Coptodactyla depressa Paulian. a
thantwice length ofother. speciesthatdoesnotbelongtothiscomplex,may
)
236 MEMOIRS OF THEQUEENSLAND MUSEUM
FIG. 4. Copiodactyla species, paramcres, lateral, and apical: A, stareyi (Smithfield); B, siunyi (19km NL
Mareeba);C,subaenea(BaldHill): D,subaenea(paralectotype, Somerset); E,subaenea(Iron Range).Allto
samescale.
occasionally have the anterior pronotal border than early season specimens. This seasonal
effaced in worn specimens, but can be distin- patternisalsoshownbythetwospeciesoccuring
guished by absence of striae 8 and 9 at elytral inareaswithcontinuousannualrainfall(C. nitida
base, male with incomplete posterior femoral Paulian, C, storeyi sp. nov.), but neither is
ridge, and female genitalia with dorsal sclcrite confined to such areas and both are found in a
reducedto an arched strut. range of forest types. Only one species, C.
mattkewsi sp. nov., appears to be restricted to
All species (for which there are biological
rainforest.
data) in thiscomplex arecatholic intheirchoice
of pabulum, being attracted to a range of Coptodactyla matthewsi sp. nov.
decayingmatter,notonlyfaeces.Nestingbiology (Figs2A,C, 3A, 7C, 8A, 9A, 10A-B, 11
hasnotbeenstudiedinthisgenus. Allspeciesare
found in areas with at least high monsoonal ETYMOLOGY.Named forEric Matthews, whose work
rainfall, and adult activity is restricted to the has resolved the major taxonomic problems in the
rainfall months, at most fromNovembertoJuly, AustralianScarabaeinae.
when there is softened soil. Thus late season MATERIAL.QUEENSLAND:Holotype, d: llkmNW
specimens are generally more worn and duller BaldHill, Mcllwraith Ra.. 13°44S I43°20E,searchparty
CRYPTIC SPECIES OF COPTODACTYLA 237
FIG. 5. Endophallus of Coptodtictylu species. A, before removal ofpenis, B-E. after removal ofpenis and
surroundingtissueandwithlateralsacevaginated:A-B,maitheusi(B-MHill);C,merafe/fca;D.m'tida(Cardwell
Range):E.nitida(Ravenshoe);b= basalsclerite;e=ejaculatorvsac: F= flagellum;I=lateralsac;p=penis;r=
ringsclerite. All tosamescale.
campsite, human dung trap, closed forest, 520m, DIAGNOSIS. Genal angles smooth orpunctate,
27.vi-12.vii.1989, TW (ANIQ. Paratypes (31) (* = muchmorefinelypuncturedthanrugoseanterior
specimen dissected): 2*, same data as holotype (AN1C); offrontoclypeus (Fig. 2A); pronotal hind angles
3*, Iron Ra., rainforest, human dungtrap, 15-21.iv.1977.
RS(ANIC);22*,Gordon'sMinearea.IronRa.,rainforest, and sides (lateral to pit and ridge) smooth and
12-18.ii.1976.GM(QM);2,LeoCkRd,McllwraithRa„ finely punctured or impunctate, without annular
30kmNECoen,500m,29.vM.vii.1976,GSM(QM); 1*. punctures; discofelytradenselymicroreticulate;
West Claudie R, Iron Ra, 24.V.I974, AWII (CMN): 1, lastventritewithconspicuousannularpunctures;
3W-e1s0t.xiCil.1a9u8d5i,eGMR..&IDroCn(RAaN.I.C)f.light trap, rainforest. apices ofparameres withelongate, narrow, vent-
rally directed lobe (Figs 2C, 3A); endophallus
238 MEMOIRS OF THE QUEENSLAND MUSEUM
FtG. 6. BndophallusofCoptodactylaspecies,afterremovalofpenisandsurroundingtissueandwith lateral sac
evaginated: A,papua; B. storeyi(Mareeba); C, storeyi(Mount Cook); D,subaenea (Bamaga); E, subacnea
'
(Bald Mill). Alltosame scale.
without apical spine on lateral sac (Fig. 5B); genal area finely punctured and smooth;
internal margin of female pygidium (Fig. 7C) posterior half of head increasingly finely and
with prominent inwardly directed lobes; ovi- sparselypuncturedtobase,puncturesmuch finer
positorwithoutdorsalsclerite; vuharsclerite Hut than pronotal disc; ventral clypeal groove
and symmetrical, without lobes (Figs 8A, 9A). shallow.
DESCRIPTION. Length: 10-13.5mm. Thorax. Pronotumalmostparallel-sidedformost
Head (Fig. 2A). Anterior of frontoclypeus of length; pronotal disc moderately strongly
wrinkledfrom linebetweenlateralangles,except punctured, interspaces 2-3 puncture diameters
CRYPTIC SPECIES OF COPTODACTYLA 239
FIG.7.Coptodactylambaenea,pygidium:A, 3;B, S;Coptoductylaspecies,apexof9pygidium:C,matthewsi;
D, menk-ka: E,pupiur. I , niiitkrX'.storeyi; II.subaenea. Ml to samescale.
and finely micropunctate with faint micro- Male, genitalia (Figs 2C, 3A, 5B). Apices of
SCulpture;hindanglesandsides(lateraltopitand parameresproducedasnarrowstraight lobeswith
ridge) smooth and finely punctured or impunc- bilobedtips: lateral sacwithapical patchofstout
tate, without annular punetures; anterior border scale-likespines,andmedian patchoflinespines
of pronotum lacking for 0.1-0.2 < width of at base of a large transverse sclerotised bar;
prothorax; elytra relatively convex in profile: flagellum relatively straight; basal sclerite
elytra dull,strongly and densely microretieulate, strongly folded, with lateral lobes; secondary
exceptshining(lessmicrosculptured),sutureand sclerite present in ejaculatory sac.
2-3 lateral intervals; elytral intervals minutely
and sparselypunctured, moreobviouslyat sides; Femalegenitalia (Figs SA, 9A, 10A-B). Dorsal
ventral surfaces ofmid and hind femora finely sclerite absent ornot strongly sclerotised; vulvar
andsparselypunctured,withoutannularpunctures. sclerite almost symmetrical, weakly sclerotised
above genital opening, without erect lobe;
Abdomen(Fig.7C).Pygidium:male:shining,not spermathecarelatively small.
microretieulate,minutelypunctured;female: dull,
strongly microretieulate, usually more evidently REMARKS.Thisspecies isknown fromisolated
punctate; internal margin of female pygidium localities on the Iron and MelIwraith Ranges,
with prominent inwardly directed lobes; last central Cape York Peninsula(Fig. 11). Thereare
ventrite with conspicuous annular punctures. no obvious morphological differences between
,
240 MEMOIRS OF THE QUEENSLAND MUSEUM
the populations, which may be continuous surfaces ofmid and hind femorawith numerous
through this rugged country. It is entirely sym- annularpunctures and large punctures.
patic with C. subaenea. All specimens were Abdomen (Fig. 7D). Pygidium: shining, not
collected in rainforest, at human dung and in microreticulate, disc finely punctured to im-
ilight traps. Collection dates include most punctate; last ventrite with conspicuous annular
months from December to July, which is the punctures; internal margin of female pygidial
monsoonal wet season. apex deeply excavate.
Malegenitalia(Figs3B,5C). Paramereswithout
Coptodactyla merdeka sp. nov.
elongate excavation, not laterally ridged, apices
(Figs 3B, 5C, 7D, 9B, IOC, 11) flat and strongly deflexed, with long, inwardly
directed, overlapping, slightly curved spurs;
ETYMOLOGY. The species name means freedom in lateral sac ofendophallus with a dense median
Indonesian.
brush of small stiff spines around small flat
MATERIAL. INDONES1A.WEST PAPUA. Holotype, sclerite; flagellum relatively straight; basal
6: New Guinea exped., Mamberamo R., Pionierbivak, sclerite quadrangular, with strong median fold
xii.1920, van Heurn (MZB). Paratypes (15): 2 ?: same and basal lobes.
dataasholotype(ANIC);13,Timika,FreeportConcession, Femalegenitalia(Figs 9B, IOC). Dorsal sclerite
IrianJaya, 1l.iii.1997,R.Ubaidillah(ANIC, MZB). sharplytransverselyridged,ridgeconvex;vulvar
DIAGNOSIS. Externally identical to C. papua plate almost symmetrical, generally weakly
Lansberge:pronotalhindanglesandsides(lateral sclerotised, producedabove genital openingas a
weak ridge; spermatheca relatively large,
topitandridge)puncturedandborderedby large
C-shaped.
annular punctures; disc of elytra not micro-
reticulate; lastventrite withconspicuous annular REMARKS. This species is known from 16
punctures; parameres(Fig. 3B) withoutelongate specimens collected in West New Guinea (Fig.
wlaittehrallaerxgceavaitniwoanr,dlnoytdliarteecratleldy, roidvgeerdl,apappiincge,s G1i1l)l,e3toifnwh1i92c5h.hTadhebesepneicdiemnetinfsietdaaksenC.aptaTpiumaibkya
slightly curved spurs; lateral sac ofendophallus were collected at 100m, in 'dry lowland forest'.
withminute subapical sclerite (Fig. 5C); ovipos- The habitat at the lowland riverplain ofPionier-
itor(Fig. 9B) dorsal scleritesharply transversely bivak is unknown.
ridged, theridge convex; vulvarsclerite approx-
imately symmetrical, without projecting lobe; Coptodactyla nitida Paulian, stat. rev.
spermatheca (Fig. IOC) significantly largerthan (Figs2B,D,3C-D,5D-E,7F,8B,9C-D, 10D-E, 12)
in otherspecies. Coptodactyla nitida Paulian 1933: 70; Matthews 1976: 32,
jun.syn.ofC. subaeneaHarold.
DESCRIPTION. Length 9.5-13mm.
MATERIAL. QUEENSLAND: Holotype, V:
Head. Anterior offrontoclypeus wrinkled from Nov.Holl.Queensld, Fry coll. 1905-100 (BMNH).
line between anterior margins of eyes, except OTHER MATERIAL. (951) (* = specimen dissected;
genal area strongly punctured, not wrinkled; label data abbreviated to site, altitude, date & collector):
posterior halfofhead moderately strongly and QUEENSLAND:88*.Atherton,7.V.1964,23-26.iii.1965.
closely punctured, punctures stronger than on 9-12.ii.1975,Bornemissza,HAH(ANICCMN); l,4mi.S
pronotal disc; ventral clypeal groove deep. Atherton, 18.ii.1975, HAH (CMN); 3, Ballv Knob,
fTihnoerlayx.anPdrosnpoatrusemlryoupnundcetduartedsi,deisn;tperrosnpoatcaelsd3i-s5c 3M1-t16.0.0iimR.,d1,69.9x98i,i6.0lGm99,M8-61.&1ii2.0D1-9C1919,3(0GQmMM,);&30.D1x,Ci.B(1aQ9lM9g)a7l-;192B5.c*ihi,.,B1a95l9md8,y,
puncture diameters, micropunctate but shining, 4-12.ii.1998, GM & DC (QM); 1*, Bilyana, 30m,
without microsculpture, punctures larger and 4-12.ii.1998, GM & DC (QM); 1*, Broadwater Park,
NW
cphliuonnsdcetrauntrgoelwdeasardnasdnsdbiodsreisdd,eersseedp(laabrtayetrealldarbtgyoe pa1i-nt2nuadlinaadrmerptiuednrgcse-;) C53ak5rmkdmweNllCaGradpIw,enlg1lh2a0Gmma,,p,42-21.12x.0iimii,..119499-88162,,.iGHi.Ma1m9l9&8e,tDGC(MQ(MQ)&M;);D11C7,,
0thu.yr2pesXo;mewairndtoetnrhiwooirfthbpororontdeheororraoxmf;oprbreaosnaenoontfuulmlaartleparuaclnkciptnaugrrtefsoo;rf IC11rQ00vMmms,),ta:l41-6I1C-20k2.*,5i,.ii.2C01va.9.ir91i89di.9w.9elG,l96lM5C,RR&aG.B,(DA2NC(.IAVCN.()1IQ9;CM6)4);,1;441G10*,B*,.FiG(DviAlealNmiIMepCise)lre;HwCC2ykk6,,,,
meliyctrroarerteilcatuilvaetley.Haitntienrvparlofsilfe;ineellyytraansdhinsipnagr,senloyt 3IlmeialeNsLHoerobkeorutton3.0.i1i4i..i1i9.6159,767,.VR.S196(9D.PGIBM),:PF1*(,A7NmIiC);SW"l
punctured, not more obviously at sides; ventral Herberton,6.xii.l968, EB(ANIC); 14*, Hinchinbrook I.,