Notes on the Rhinocypha cucullata Selys group from Borneo, with a description of R. viola spec. nov. (Zygoptera: Chlorocyphidae)

9 Pages·2002·2.7 MB·English

by  Orr, A G

Odonatologica31(3):287-295 September1,2002
Noteson theRhinocypha cucullata Selys group
from Borneo, witha descriptionof
R. viola spec. nov.
(Zygoptera: Chlorocyphidae)
A.G.Orr
CRC-TREM, GriffithUniversity,Nathan, Qld-4111, Australia
ReceivedOctober 16,2001/RevisedandAcceptedJanuary19, 2002
Thenewsp.fromthecentral KalimantanprovinceofBorneoisdescribedandfigured.
TheoriginaltypeseriesofR.cucullataSelyswasexaminedanda3specimenisdesignated
aslectotype.Thesingle 9syntypeisshowntoinfactbeR. humeralis Selys.Thetrue 9
RR. cucullatais described and figuredfor the first time. Sign.ificantcharacters of
aurofulgensLaidl.arefiguredforcomparativepurposes. Keysareprovidedtobothsexes
ofthe three species, comprisingthe extended cucullata group ofF.F. Laidlaw (1950,
Trans. R.ent. Soc.Land. 101:233-269).
INTRODUCTION
InhisrevisionoftheChlorocyphidae, LAIDLAW(1950)definedthecucullatagroup,
including Rhinochypha cucullataSelys, 1873andK. aurofulgensLaidlaw,1931,species
bothendemictoBorneo.Recently IexaminedmaterialfromcentralKalimantaninthe
collectionofDr AllenDavies,representing averydistinctnewspecies ofRhinocypha,
clearly belonging tothis group.Thematerialwas partof asmall series collectedby
Chris Jiggins, thenan undergraduate atCambridgeUniversity, andwas thesubjectof
asmallecologicalandbehaviouralstudy. A descriptionis provided below.
Inthecourseofstudyingcomparative materialIalsobecameawarethattheoriginal
description ofR. cucullata(SELYS, 1873)comprises acomposite species, maleand
femalesyntypesbeingdistinctspecies. Tostabilizethepresentnomenclaturalusage,a
malesyntype isdesignatedaslectotypeofR. cucullatahere.Thesinglefemalesyntype
isreidentifiedas R.humeralisSelys, 1873,aspecies whichwas describedinthesame
paper. Materialofboth species labelled “Labuan”(but in fact possibly originating
somewhereon mainlandBorneo)was availableto Selys and itappearshemadethe
wrong association, no recognized femaleofR. humeralisbeing recorded. As new
288 A.G.Orr
materialhas cometo my noticefromamongM.A.Lieftinck’scollections, housedin
theNationalMuseumofNaturalHistory,Leiden,Iamabletoprovideadescription of
whatisalmostcertainly thetrue femaleofR. cucullata.
Thefollowingabbreviationsare usedtoindicatedeposition oftypematerial:CUM,
CambridgeUniversityZoology Museum,England; — IRSN,InstitutroyaldesSciences
naturellesdeBelgique, Brussels, Belgium; — RMNH, NationalMuseumof Natural
History,Leiden,theNetherlands.
RHINOCYPHA VIOLA SP. NOV.
Figures la,b,c,2e,f,3a,4a
Material. — Holotype <J;central Kalimantan,SgMarung.(0°6'S, 114°19’E),7-VIII-1992,C.
Jigginsleg.(depositedinCUM). — Paratypes: 3 5,19,samedataasholotype; 1 6 coll.A.G. Orr;
remainder in CUM.
Etymology — viola, strictly afemininenouninapposition, suggestingthe violet colourofthe
hindmostsegments oftheabdomen.
maleholotype. — Head:dorsally mostlyblackwithbasesofmandiblesandgenae
light blue, and five small, yellowish marks on occiput as follows: a pairofspots
immediatelybehindposteriorocelli,apairofsmallpostocularspotsandashorttransverse
medianbaronthehindmarginoftheocciput. Antennaeblack. Ventrallyblackexcept
forbaseoflabiumwhichis well-definedcreamywhite.
Thorax. — Prothorax black marked withpale greenish-yellow; anterior lobe
entirelypaleabove;medianlobewithincompleteslightly roundeddorsolateralbands;
posteriorlobewithlateralangles pale.Synthorax darkabovewithveryfine palegreen
antehumeralbandsbrokenanteriorly toformasmall spot, continuing as afine linefor
about two thirds of the length of the humeralsuture, thenreappearing as a distinct
dorsal spot immediately in frontof theantealartriangle; antealartriangle traversed
lengthwise by two very fineparallel yellow lines; sideswithvery short, fine,dorsal,
greenish posthumeral stripe, asmall greenish dorsal streak immediately posteriorto
mesometapleural suture anda relatively narrow irregular pale blue transverse band
extending from themesinfraepistemum to thebaseoftheabdomen.Venterpalewith
extensiveblackmarking. Legs mostlyblack; coxaeandtrochantersyellowishbehind;
innersurfaceofmidandhindtibiaewithbright whitepruinescence; tibiaenotexpanded.
Wings (Fig. la) long and narrow, rounded apically; innerpart hyaline with heavy
brownish-yellow suffusion;apex withverybroadopaquepatches,only slightlyiridescent
onbothsurfaces; innermarginofapicalpatch fairlywelldefined;inforewing meeting
costaaroundPx7andstrongly slantedoutward;inhindwingmeeting costaaroundPx
5andonlyslightly slantedoutwards.Forewingopaquepatch withsmall, central, non-
iridescentwindowconsisting oftwo rows offive cellslevelwiththebasalendofthe
pterostigma (in the holotype both forewings are set on the left side of the body);
pterostigmablack, long,coveringabout8 cells,acutebasally andapically andexpanded
apically.
Rhinocyphacucullata-group 289
Abdomen. — Dorso-ventrally
flattenedwithstrongmediancarina;
subapical encircling dorsolateral
flange on segment2 very slightly
expanded (Fig. 3a); ground colour
mattblack,dorsumofsegments 1-
9 marked with blue and lilac as
showninFigurelbandasfollows:
segments 1-4 light cobalt blue;
segments 5-7 purplish blue;
segments 8-9 deep lilac; laterally
withbluemarking on segments 1-
5 only; segment 1a distinctlateral
spotjustreaching dorsum;segment
2,abroadapically expanded patch
continuouswith blue visible on
dorsum (Fig. lb) and forming
nearlyamirrorimageofit; segment
3asmallisolatedstreakbasallyand
aspotapically; segments4-5each
withatiny,obscure,isolated,basal
spot. Remainderofabdomenblack.
Anal appendages black, superiors
stronglyforcipateapically,inferiors
abouthalflength of superiors.
Measurements (mm):Abdomen+
appendages 18;hindwing22.5.
Fig. 1. Structural features: (a) Rhinocypha viola sp.n.
FEMALE paratype — Head holotype6fore-andhindwing;—(b)R. viola sp.n.holotype
(Fig. 2e): dorsally mostly black <Jabdomen,dorsal aspect;—(c)R.violasp.n.Sabdomen,
marked as follows: bases of lateralaspect; —(d)R.cucullata Selys $abdomen,lateral
mandiblesand genaepale yellow aspect (samescaleaslast); —(e) R.aurofulgensLaidlaw
butlabrumblackexcept forsmall (KualaBelalong,Brunei) 9 abdomen,lateral aspect(same
scale aslast).— [Scalebars =3mm]
lateralyellow spots (Fig. 2f); tiny
wellseparated paired orangespots mid-dorsally on postclypeus and frons; sides of
fronsandvertex withfine pale linebordering compound eye; apairof tiny orange
spots immediately behindposteriorocelli; occiput withtiny orangepostocular spots
connectedby ashortindistincttransversemedianbaronthehindmarginoftheocciput.
Antennaewithexternalsurfaceofsegment 1paleyellow,remainderblack. Ventrally
headblackexcept forbaseoflabiumwhichis extensively creamywhite.
Thorax. — Prothoraxblack, dorsallymarkedonthemedianandposteriorlobes
withpaleyellow lateralstreaks forming a discontinuousband oneithersideviewed
fromabove;anteriorlobealmostentirelypaleabove;sides dark.Synthorax darkabove
290 A.G. Orr
with very fine pale yellow antehumeral bands
brokenanteriorlytoformasmallspot,continuing
as afinelineforaboutthreequartersofthelength
ofthehumeralsuture,thenrecurringasatinydorsal
spotimmediately infrontoftheantealartriangle;
antealartriangletraversedlengthwise bytwo very
fine parallel yellow lines; faintyellowish dusting
onmediancarina; sideswithfine,dorsal,yellowish
posthumeral stripe, abouthalflength of humeral
suture,asmallyellowish dorsalstreakimmediately
posteriortomesometapleural suture,asmalldorsal
spotonthemetepistemum andarelativelynarrow
irregularcreamy transverse band extending from
themesinfraepisternumtothebaseoftheabdomen,
including almost all the metepimeron. Venter
(poststemum) groundcolourblackwith fourpale
marks as showninFigure 4a. Legs mostlyblack;
posterior halfof coxae, allofthe trochantersand
basesoffemorayellowishbehind.Wingslongand
Fig. 2.Dorsal viewofhead andanterior
view oflabrum infemales: (a-b)R. narrow, hyaline with heavy brownish-yellow
aurofulgens;— (c-d)R. cucullata; —(e- suffusion.Pterostigmablack, long,covering5-5.5
-f) R. viola sp. n.(paratype). — [Scale cells, acutebasally andapically.
bars= 1mm] • Abdomen (Fig. 1c). — Blackwithcreamyto
orangemarkings; segment1withlateralspot;segments2-7withdorsalstreakandspot
likeabackward directedexclamationmarkandventralstreak;dorsalandventralstreak
onposteriorsegmentsmuchreduced;mediancarinaondorsumofsegments2-7marked
withyellowish orange. Remainderofabdomenblack. Anal appendages and valves
black.
Measurements (mm). — Abdomen+appendages 17;hindwing23.5.
REMARKS.- This strikingspecies appearstobe closestto R. aurofulgens Laidlaw,
knownfromnorthernBorneo,andalsoshowsstrongaffinitieswithR. cucullataSelys,
widely distributedthroughout lowlandBorneo.ThemalediffersfromR. aurofulgens
inthedetailsofcolouronthehead,inthereducedcolouronthethorax,inthepresence
oftibialpruinescence, intheshape and colorationofthewings andintheabdominal
colorationwhichislessextensiveandofverydifferenthues.Theconspicuous subapical
flange onsegment2of maleR. cucullata(Fig. 3b)andR. aurofulgens (Fig.3c)isonly
very slightly developed inR. viola(Fig.3a). Thereislittlevariationinthetypeseries;
in two oftheparatypes the small windowin the forewing is entirely lacking. The
femaleis similar to that ofR. aurofulgens (Figs le, 2a,b, 4c), saving that the pale
colorationisconsiderably reduced,thelabrumis extensivelyblackinR. viola,creamy
in R. aurofulgens, andallothermarkings areconsiderably narrower. Ontheventerof
the synthorax (Figs 4a, c) the third marking fromthefront in R. viola is distinctly
Rhinocyphacucullata-group 291
smallerand emarginate on its
posteriorborder,whereasthatofR.
aurofulgens is mostlybroaderand
crown- shaped. On the abdomen,
besides an overall reduction in
extent of markings, pale lateral
markingspresentonsegment8and
segment 10 ofR. aurofulgens are
entirely lacking in R. viola.The
femaleof R. cucullata (Figs Id,
2d,e,4b) is very similar tothat of
R. viola. Apparent differencesare Fig.3.Dorsal view ofbaseofmaleabdomen: (a)R. viola
discussedbelow. sp.n.(holotype); — (b)R. cucullata (lectotype); - (c) R.
HABITATANDBEHAVIOUR. - R. aurofulgens(KualaBelalong,Brunei).Noteexpandedflange
viola was the subject of an nearapexofsegment2,especiallyprominentinR.cucullata.
ecological and behavioralstudy — [Scalebar=1mm]
(JIGGINS, 1993) in which itis erroneously referredto as R. cucullata Selys. The
localityisalmostatthegeographical centre ofBorneo,nearthejunctionoftheMarung
andBaritoRiversat analtitudeof 150-200m.Thevegetation includeddisturbedand
primary lowlanddipterocarpforest. Itwas foundonsmall, clear,rocky foreststreams
branchingoffthemainriverinbothdisturbedandprimaryforestbutespecially inthe
latter.Itwas by farthemostfrequentlyrecordedof22odonatespecies whichincluded
Rhinocypha spiniferLaidlaw,Heliocyphabiseriata(Selys),Dysphaea dimidiataSelys,
Euphaea imparSelys, E. subcostalis(?) Selys, Neurobasislongipes Hagen, Vestalis
amoena Selys, Rhinagrion borneense(Selys), undeterminedCoeliccia and
Prodasineuraspp.,Orthetrumchrysis (Selys), O.pruinosumschneideri(Forster) and
Zygonyx iriserrans (?)Lieftinck(JIGGINS, 1993).During territorialdisputes males
ofR. violafacedeachotherhovering withtheirabdomenscockedupwards. Thisisthe
exactpostureadoptedbyR. aurofulgens andR. cucullata,as describedandfiguredby
ORR (1996), butnot otherRhinocypha. Courtship behaviourinvolvedthe display of
thepruinosed tibiaeasisusualinthefamily(butnotR.aurofulgens). Femaleslaidtheir
eggs inthe rotting woodoffallenlogs inthestream. Thespecies was evidently very
common where it occurred, but the extentofits distributionis uncertain. Central
Kalimantanis very poorly collectedbut itis surely an obviousinsectand wouldbe
expected tooccur eveninstudentcollections, whichare appearingsporadically.
RHINOCYPHA CUCULLATA SELYS, 1873
FiguresId,2c,d, 3b,4b
Rhinochyphacucullata,Selys 1873: 492-493[pp. 28-29inreprintedn]
Typematerial. — Lectotype(heredesignated)<3[abdomendetached],“LabuanBorneo”(yellow
292 A.G.On-
label inSelys’ hand), “Rhino,humeralis S.” (white label inSelys’ hand), “Collection Selys/cucullata/
Revision Martin 190” (grey label),“Coll R.I.Sc.N.B./Borneo/ excollection Selys” (yellowlabel). —
Paralectotypes36 allwith“CollR.I.Sc.N.B./Borneo/excollection Selys”(yellowlabel):1 [abdomen
missing],“Labuan,Borneo”(yellowlabelinSelys’hand),“RhicucullataSelys <JLabuan”(whitelabelin
Selys’hand); 1 [head and abdominal segments3-10 broken andinseparate capsules],“Labuan” (white
labelnotinSelys’hand), “Labuan Borneo”(yellowlabel inSelys’ hand),“114” (tworoughgrey labels
bearingthisnumber);l[abdominalsegments 4-10 missing], “Labuan Borneo” (yellowlabel inSelys’
hand),“Rh. Cucullata Selys SLabuan”(whitelabelinSelys’hand). Allabovespecimensareheldinthe
IRSN, Brussels,
Other material.— 1 <J,Sarawak(inSelys’hand),(coll.Selys,IRSN); — 1 S,1 2, SEBorneo,
S.Mentawirleg.,20-10-1950 (RMNH); — 36,KualaBelalong,Brunei,IX-1992,A.G.Orrleg.; 1S,Sg.
Benutan,Brunei,V-1995,A.G.Orrleg.(coll.A.G.Orr).
male. — The lectotype is designated onthe basis of being the most intact and
recognizable specimen oftheputative type series available.The determinationlabel
“R. humeralis”(in Selys’s handwriting)attachedtothepinofthelectotypeisconfusing.
Itmay havebecomeplacedthereduetoanerror bySelys himself (or by somelater
worker), sinceSelys inthesamepaperinwhichhedescribedR. cucullata, accurately
describedalsoR. humeralis.Itwouldscarcelybepossible foranexperienced worker
makingadirectcomparison toconfusethetwo speciesandindeedthespecimen matches
extremely well with Selys’ description ofcucullata, thatofhumeralis being very
different.Evidencethatthelabelhasbeenrepeatedlyremovedisprovided bythepresence
ofthreeseparatepin holeswhichlendscredencetothesuggestion thatthelabelwas
erroneously placed on the specimen well after the determinationwas made.Other
determinationlabelshaveonly onehole.
Sinceinhisdescription,Selys givesonly “Labuan”asthelocalityofthetypematerial,
the specimen labelled“Sarawak”inColl.Selysisnotconsideredtorepresentasyntype.
Inhis original description SELYS(1873) indicatesthat heexaminedan unspecified
numberofmalesandfemalesfromhisown collectionandfromthatofR. McLachlan
(nowheldintheBMNH). Itis probable thatonly onefemalewas available, since he
provides asinglesetofmeasurements,whereasforthemalesheprovides arange.The
seriesexaminedis thought toincludeallsyntype malessince according toMrDavid
GoodgertherearenomalesfromLabuanintheMcLachlancollection.Selys’ description
of the maleofR. cucullataremains adequate except for the colourof the dorsal
abdominalmarkingsonsegments 8-9whichare pale lilacratherthanblue,atleastin
thelivingspecimen. Othersalientfeaturesare listedinthekeybelowandthemodified
secondabdominalsegmentofthelectotype is figured (Fig. 3b).
FEMALE. — Thefemalesyntype ofcucullata from “Labuan”,heldin theNatural
HistoryMuseum,London,exMcLachlancollection,andaccurately describedby Selys,
isclearlythefemaleofR. humeraliswhichis wellknownfromcollectionsof mating
pairs.Itbearsconspicuous broadterminalmarkingsonallwings.Ihavecloselyobserved
(butnot collected)R. cucullatamating inBruneiandalthough Iamunabletoprovide
a description of the femaleon this basis Iam certainthe wings ofthefemalewere
withoutterminal markings. Recently Iwas ableto examine,preserved in the same
Rhinocyphacucullata-group 293
envelope, apairofR. cucullatafromS.E.
Borneo.Thesewereevidently collectedby
A.M.R.Wegner, whosematerialfromthe
samesiteanddateislistedinLBEFITNCK
(1953).Ontheenvelope inLieftinck’shand
is written“9Rhinocypha cucullata(?)”. It
seemsreasonabletosupposethatthis is a
genuine pair,especially asitisinagreement
with my ownfieldobservations.Itisalso
reasonableto suppose that any doubtin Fig.4.Venter offemalesynthorax(poststemum):
Lieftinck’s mind might havearisen from (a)R. violasp.n.(paratype); —(b)R. cucullata;
thecleardifferencebetweenthisspecimen — (cR. aurofulgens. — [Scalebar= 1mm]
andSelys’ description withwhichhewas almostcertainly familiar.Adescription of
thefemalecucullatafollows.
Head (Fig. 2c). — Dorsally mostly black markedas follows:basesofmandibles
andgenaepaleyellow butlabrumblackexceptformoderatesizedlateralyellowspots
(Fig. 2d); fronswithsmallpairedspots. Inallotherrespects similartoR. violafemale
(above) exceptforthemarkings onthepostclypeus whichisalmostentirelyblack with
tinypale spotsconfinedtoitsposteriorcomersandthepresenceoftinypairedelongate
spots inthe intra-ocellartriangle, lacking inR. viola.
Thorax. — Palemarkings similarinformtothosedescribedaboveforR. violabut
a littlereduced. Legs and wings as in R. viola. Venterof synthorax (poststemum)
groundcolourblackwithfourpalemarks asshowninFigure 4b; differsfromR. viola
especially intheformofthethirdmarking fromthefront,whichisroughly ‘V’shaped
andtheposteriorband whichis considerably broaderthaninR. viola.
Abdomen (Fig. Id). — Black with creamymarkings, very likeR. violaand
differingas follows:segments2-5withdorsalstreakandspotlikeabackwarddirected
exclamation markandventralstreak; dorsalstreakabsentonsegments6-7; posterior
spots on segments2-5 largerthaninR. viola.
Measurements (mm).— Abdomen +appendages16.3;hindwing22.8.
remarks. - ThefemaleofR. cucullatais verysimilartothatofR. violadescribed
aboveanddiffersfromR.aurofulgens inbeingconsiderably darker.Differencesfrom
R. violatendtoinvolveaslightlossofpale colorationbutonthelabrumpalespots are
welldevelopedas opposedtosmallandobscure inR. viola.Theposteriormarkings on
theventerofthesynthorax are quite differentandmay provediagnostic. Differences
indicated,ifconsistent, wouldbesufficienttoseparatethetwobutasbothdescriptions
areeachbasedonasingle specimen, theremustremainsomeuncertaintyregarding the
possibilityofseparatingfemalesofthetwo species. The femaleofR. aurofulgens, of
whichfivefemales were examined(all fromKuala Belalong, Brunei), shows some
variation, includinga considerablereductionin thepale colorationinonespecimen,
althoughinthisindividualthemarkingonthelabrumandbasalsegmentsoftheabdomen
are stilldefinitive.
294 A.G. Orr
DISCUSSION
ThereseemslittledoubtthatR. aurofulgens, R. cucullataandR. violaformanatural
grouping with R. violaperhaps being themost primitive.Inthe maleofR. viola the
general formoftheabdomenis likethatof theothertwo species butintheposterior
flange onsegment2,aprobable synapomorphy, ispoorly developed (Fig. 3a). Using
R.humeralisforoutgroupcomparisons, thelegsare unmodified(expanded incucullata,
colourreduced in aurofulgens) and the wing shape and wing marking is the least
modifiedofthethreespecies.
Akey tothemalesandfemalesof thethreespecies follows:
MALES
1 Innerborderofterminalpatchesonallwingsmeetingcostanearerpterostigmathan nodus;upperand
undersides ofpatchesentirelywith lightmetallic copperiridescence. Dorsumofabdomenextensively
brilliantlightblueonsegments 1-9withthinmedianblacklineondorsalcarina;segment2somewhat
expanded into flangeat posteriormargin. Legsentirely black, lacking pruinescence (Fig.
3c) aurofulgens
— Innerborderofterminalpatchesonallwingsmeetingcostanearernodusthanpterostigma;upperside
ofpatchesdark andonly slightlyreflective.Abdominalmarkingdifferentfromabove;segment2with
orwithoutflange.Legswith whiteorblue pruinescenceonmid andhindtibiae 2
2 Wingsbroad;underside ofapicalpatch with brilliant coppery-greeniridescence onbasal twothirds.
Dorsumofabdomenblackexceptforsegments 8-9whicharepalelilac;segment2withaverywell
developedflangeattheposteriormargin,blueonitsanteriorface. Midandhind tibiaeexpandedand
pminescentlightblueonthe flexor surface(Fig.3b) cucullata
— Wingsofnormal width;undersidelacking strong iridescence. Abdomendorsumextensively dark
■ blue gradingtoviolet posteriorlyonsegments 1-9;mediancarinawithbroad black line; segment2
scarcely expandedat posterior margin.Mid and hindtibiae withbright white pruinescence; little
expanded(Figsla,b,3a) viola
FEMALES
1 Labrumentirelypale;spotsonpostclypeusandIrons conspicuous.Abdomenwithbroadlateralbands
confluent withposteriorspotsatleastin segments 2-3;small lateral spots presentonsegments8 and
10(Figs le,2a,b,4c) aurofulgens
— Labrumatleastpartiallyblack;spotsonpostclypeusandfrons verysmallandinconspicuous.Abdomen
darkerwithlateralbandsonanteriorsegments notmeetingposteriorspotandnolateralspotsposterior
tosegment7 2
2 Labrum withlateralspots almostround andwell developed;spotsabsentfromcentreofpostclypeus.
Third pale markfrom frontonpoststemum roughly ‘V’ shaped, posteriormark relativelybroad.
Dorso-lateral bandslackingonabdominal segments 6-7(Figs Id,2c,d,4b) cucullata
— Labrum with smallernarrowerlateral spots; small butdistinctpale spots oncentre ofpostclypeus.
Third palemark fromfrontonpoststernum ‘beret’-shapedwith asmallanteriormedian projection,
posterior mark relativelynamv.Very weak dorso-lateralbandspresent onabdominal segments6-7
(Figs 1c,2e,f,4a) viola
Rhinocyphacucullata-group 295
ACKNOWLEDGEMENTS
IthankALLENDAVIESfortheloanandgenerous donationofspecimensandmuchpracticalassistance
andhospitality,DAVIDGOODGERforlocatingandprovidingaphotographofthesyntypefemaleofR.
cucullata,JACQUES COOLSandPATRICKGROOTAERT forarrangingaloanofthemaletypeseries
of R. cucullata,JAN VAN TOLforarranginga generalloan ofspecimensfromLeiden and MATTI
HÄMÄLÄINENforassistingwithliterature andforprovidingmanyhelpfulcommentsontheprojectand
manuscript.
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new genera,anddescriptionofanew geographicalsubspecies.Trans.R. ent.Soc.London 101(8):
257-280.
LIEFTINCK,M.A.,1953.Additions totheodonatefauna oftheIndo-Australianarchipelago.Treubia22
(1):233-269.
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rivers.In:Murung'92:TheCambridgezoologicalexpeditiontoBorneo,pp. 77-94.Unpublished
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ORR,A.G., 1966.Territorial andcourtshipdisplaysinBorneanChlorocyphidae(Zygoptera).Odonatologica
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SELYSLONGCHAMPS,E.de, 1873.Troisiemes additionsauSynopsis desCalopterygines.Bull.Acad.
Belg.(II)35: 492-493.