Evolution, taxonomy, and biogeography of ancient Gondwanian libelluloides, with comments on anisopteroid evolution and phylogenetic systematics (Anisoptera: Libelluloidea)

42 Pages·1995·3.2 MB·English

by  Carle, F L

Odonalologica24(4):383-424 December I. 1995
Evolution, taxonomy, andbiogeography
ofancient Gondwanianlibelluloides,
with commentsonanisopteroidevolution
andphylogeneticsystematics
(Anisoptera:Libelluloidea)*
F.L. Carle¹
DepartmentofEntomology,Cook College,New JerseyAgriculturalExperimentStation,
RutgersUniversity,New Brunswick, NJ 08903,United States
Received October 14, 1994/Reviewed andAcceptedApril4, 1995/Final additions and
modificatonsreceived May 17,1995
New phylogenetic systematicmethodologies are presented and the terms
‘neapomorphy’,‘coapomorphy’, ‘exapomorphy’and ‘apophyletic’introduced. Re-
sultsarepresentedinsortedcharacterstatematriceswhichshowthe outcomeofchar-
acterstate evaluation while depictingphylogeneticarrangement.Higherclassifica-
tion,phylogeny,andbiogeographyofAnisopteraarereviewed andkeysprovidedfor
superfamiliesand families. The pattern ofanisopteroidneapomorphysupports the
superfamilyarrangement proposedby F.L.CARLE (1986, Odonatologica15: 275-
-326),whileindicatingpolyphylyforCordulegasteroidea[sic]ofEC. FRASER(1957,
A reclassificationofthe order Odonata,R. Zool. Soc NSW,Sydney) and both
“Neanisoptera”and“Petaluroidea”ofH.-K.PFAU(1991,Adv.Odonalol. 5:109-141).
Paraphyleticgroupingsinclude Aeschnidae [sic] ofR.J. TILLYARD(1917,Thebiol-
ogyofdragonflies,CambridgeUniv. Press),andCordulegastroideaofD.A.L.DAY1ES
(1981,Soc.int. odonatol. rapidComm. 3: 1-60).Nothomacromianom.n.isproposed
asareplacementforPseudomacromiaCarle &Wighton, 1990,nec.Pseudomacromia
Kirby, 1890.Congruencebetweenphylogeneticandbiogeographicpatternsindicates
twoormore distinct mesozoic utilizations ofatrans-pangaeian montanedispersal
route. Incorrectassociation ofNeopetaliawithaustropetaliidsforthe past 137years
has obscuredthe gondwanianoriginand subsequentradiation ofnon-cordulegastrid
Libelluloidea,aprocesswhichbeganatleast 140million yearsago onthenowfrozen
continent ofAntarctica. - Chlorogomphidevolution waslargelyinfluenced by un-
known vicarianteventsrelatedtothe combined forcesofmountainbuildingandrapid
driftofIndia toward theequator,coupledwith a40-60 million yearinsularisolation.
*Dedicated tothe memoryofJ.A.L.Watson
1 Mailingaddress: 146Mountain ViewRd, Warren,NJ 07059, United States
384 F.L.Carle
insular isolation. Classification isrevised as follows: Chlorogomphidae:Chloro-
petaliinaesubfam.n.: (Chloropetaliinitrib. n.):Chloropetaliagen.n. [type C. selysi
Fraser];- Chlorogomphinaecomb,n.: (Eorogomphinitrib. n.):Eorogomphusgen.n.
[type O.preciosus Fraser];(Sinorogomphinitrib.n.);Sinorogomphusgen.n.[type C.
nasutusNeedham];(Chlorogomphinicomb,n.):Neorogomphusgen.n.[typeC. fraseri
St.Quentin],Indorogomphusgen.n.[type O. xanthopteraFraser],OrogomphusSelys,
Aurorachlorus gen.n.[typeC.papilioRis], ChlorogomphusSelys.- Earlysynthemistid
evolutionwaslargelyinfluenced byunknown vicarianteventsinAntarctica relatedto
thecombined forces ofmountainbuildingand driftofAntarctica-Australia towardthe
southpole,coupledwith a60 million year isolation. Resultingselective pressures
producedthe largestsetofcongruent neapomorphyandexapomorphyknown in the
Anisoptera.Classification is revised as follows: Synthemistidae:(Synthemiopsini
trib.n.):SynthemiopsisTillyard;(Palaeosynthemistinitrib.n.):PalaeosynthemisForster,
Archaeosynthemisgen.n.[type:S.leachii Selys];(Synthemistinicomb.n.):Synthemis
Selys, Parasynthemisgen.n. [type S. reginaSelys], Calesynthemisgen.n. [typeS.
miranda Selys];(Eusythemistinitrib.n.):Austrosynthemisgen.n.[type:S. cyanitincta
Tillyard], ChoristhemisTillyard, EusynthemisForster.
INTRODUCTION
Currently only amodicumofagreementexists concerning anisopteran classifi-
cation.Disagreement is not primarily betweenso called“splitters and lumpers”,
butisratheraresultofdifferentmethodologies basedon diverseperceptions con-
cerning thefunctionofclassifications.Phenetic andhybridphenetic-phylogenetic,
basedmethodologies such as that utilizedby TILLYARD&FRASER(1940) and
FRASER (1957) are inherently anthropogenic and allow paraphyletic and in the
formercase polyphyletic groupings. Theobjective ofphylogenetic systematics is
to determinephylogeny upon which taxonomy is based, a consequenceis that
phenotypically similargroupsmaynot begrouped togetherifobvious similarities
areduetoplesiomorphy. Becausephylogenetic keys, likephylogenetic classifica-
tionsareinherently morestableandconfermoreinformationthanartificialphenetic
constructs, thefollowing descriptive phylogenetic keys are presented in place of
isolateddescriptions.
Phylogenetic methodologies potentially offertremendouspredictive power,but
withoutcareful characterevaluationinformationcontentcan beseverely reduced.
Characterweighting is then thecrucialissue: inphenetic classificationcharacters
areeitherweighted 0ifnot recognized, and 1 ifrecognized, orgreater depending
on howfinely a“character” issplit;in Hennigian phylogenetic classificationchar-
actersare weighted0ifunrecognized orplesiomorphic, and 1ifapomorphic. How-
ever,inphylogenetic systematics as developedandutilized by myself, in addition
to plesiomorphy being weighted 0, apomorphy is consideredto beeither neapo-
morphic,coapomorphic, orexapomorphic, withrelativeweights 1,0,and maximally
Vi, respectively. Categorization ofapomorphy is required toeliminatefalsecon-
gruencebased on correlated characters (coapomorphy), and to minimize false
synapomorphy based on convergent losses(exapomorphy). Neapomorphy then,
Evolution ofGondwanian Libelluloidea 385
representsa verysmallproportionofthetotalcharacterstatesstudied,inparticular
itrepresentsa character(orcorrelatedcharacterset) representing a unique evolu-
tionarysolution to aspecific setofselectivepressures.
Theobjective ofthenewapproach is to discovercongruentneapomorphy upon
whichaclassificationcomprising monophyletic groupsisbased,withgroupsranked
according to relative geologic age as determinedfrom the geologic record and
biogeographic vicariance.Determinationofneapomorphy throughtheelimination
ofcoapomorphy necessitates a keen understanding of function, as seemingly
uncorrelatedcharacterstatesmay bepartofasingle functionalsystem. Forexam-
ple,lossofthemalehindwing angulation andanalbrace isassociatedwiththeloss
ofabdominalauricles,thishasoccurredthreeseparatetimesintheAnisoptera (Tab
I, apomorphy set4),andpresumably reflects achange inthemethodusedtoestab-
lishthecopulatory position. Genuinecongruent neapomorphy typically involves
differentsystems, sexes, or life stages.For example, gomphid neapomorphy in-
cludeslarvalantennalsegment3enlarged, larvalproventriculus withrasplike lobes,
adulthead withpostocellar ridge, adult palpal end hook apically spinelike, and
maleposterior hamuliengaging femalesternum 9 withananteriorstroke.
HENNIG (1966) introduced the term paraphyletic to describe the class of
nonmonophyletic taxabasedonsymplesiomorphy, butunfortunately hisdefinition
also applies to certainpolyphyletic groups.Amore directapproach is todescribe
the group whose elevation in rankresults in the creationofa polyphyletic or
paraphyletic group;hereparaphyletic is limited to a groupthatincludes a most
recent common ancestor but not all of its descendants.For groupsofrelatively
exaggerated taxonomicrankthetermapophyletic isintroduced.Apophyletic groups
are typically based on obvious phenetic differencesand ignorance ofcongruent
neapomorphy, andalthough they areoften monophyletic they areresponsible for
mostofthecontroversy concerning anisopteroid classification.Aclassicapophyletic
groupis the “class”Aves which is thought to be part ofthe Dinosaur suborder
Theropoda. Inthepresentstudy apophyletic groupsincludeChoristhemisTillyard
andChlorogomphus Selys (ifOrogomphus Selys isrecognized).
Thenew methodlogy is not rigid,butbecausealltree topologies can bereduced
toapectinateseriesofmonophyletic groups,atableofnestedcharacterstatepos-
sibilitiescanbeinitially constructed.Homologous characterstatedistributionsare
thentabulatedwiththeorderoftaxonomicgroupsrearranging asexamplesofeach
characterstatepattern arediscovered. Notethat some patterns maynot match the
initialpectinate approximation and can be added if itis not desiredto combine
groups as for characterset 9ofTable I.Tenor moretaxonomic groupsshouldbe
includedsothattheprobability ofareversedtopology is minimal;note thatifonly
threegroupsareconsideredpolarity is confounded.Polarity is not predetermined
withfossilevidence,ontogeny,oroutgroups,butis anautomaticoutcomeofnested
congruent neapomorphy as long as the numberofgroups is greater than 3, but
preferably greater than 10. Whileexamples for each character state pattern are
386 EL.Carle
discovered, apomophic character states areevaluated by determining ifthey are
neamorphic, exapomorphic, or coapomorphic.
Careful characterstateevaluationgenerally eliminatesthe need toutilizeparsi-
mony analyses to determinethe besttopology from severallikely incorrectpossi-
bilities, exceptions are situations involving excessive exapomorphy. Becausethe
statistical robustness ofnestedcongruent neapomorphy is so high, an estimated
phylogeny willgenerally stand or fall based on characterstateevaluation; thisis
bestpresented inasorteddatamatrixwhichnot only shows theresultsofcharacter
state evaluation, but also depicts the estimatedphylogeny. In the presentpaper
neapomorphy is denoted by N, plesiomorphy by exapomorphy by X, and
-,
coapomorphy byC; lowercaselettersareusedtoindicateconditionswithingroups.
Ineach apomorphy set neapomorphy is listed first with each neapomorphy fol-
lowedby respective coapomorphy; exapomorphy is listedlast.
Numericalevaluationcanbeachievedby simply adding neapomorphic charac-
tersets although theinformationcontent frompossibly convergentcharacterstates
is more difficultto discern.Apparent neapomorphic convergenceoften coincides
withthelettern inthedatamatricesandtypically involvespoorly understood(lOf,
Tab. I),superficial (16c,Tab. I),or obviously nonhomologous characters(4c, Tab.
I).Experienced systematists areactually quiteadept atdiscerning homologousand
nonhomologous similarity, butreinterpretation is always possible. Therefore, ap-
parentneapomorphic convergenceshouldbelistedeventhough itisgiven no weight
inananalysis. In contrast,theprobability ofexapomorphic convergenceis always
relatively high dueto difficultiesin distinguishing exapomorphic similarity,there-
fore,resulting noncongruentexapomorphic characterpatterns areexpected andof
littleconcern.However,theformula1/(X+1),whereXis thenumberoflossevents,
canbe utilizedto weightcongruentexapomorphy sets orcongruentneapomophy-
-exapomorphy sets;forexample itis 1/5 forexapomoprhy set 14dofTableI, and
1/2 for neapomorphy-exapomorphy set 12g ofTableI.The apomorphy score for
TableIis asfollows: apomorphy set 1 (3.0),2(7.0), 3(5.5), 4(4.3),5(6.0),6(1.8),
7(6.0), 8(4.8), 9(2.0), 10(6.5), 11 (7.3), 12(5.8), 13(5.3), 14(3.7), 15 (8.0), 16
(2.5), 17(4.0), 18 (9.7), 19(1.3),20(4.0), 21 (1.5),22 (5.0); total score= 107.0,
averagescore =4.9. Other arrangementsofTableIareofcourse possible, butthe
addedexapomorphy required tomakeobserved neapomorphic charactersets con-
gruentsignificantlyreduces thetotalscore. Notethatmuchcoapomorphy hasbeen
removed from the tables to save spaceand thatcoapomorphic exapomorphy is
sometimeslistedwithrespective neapomorphy (e.g. TableI, character5g).
KEYTO THESUPERFAMILIESANDFAMILIESOFANISOPTERA,
EXCLUDINGFAMILIES OFNON-CORDULEGASTRID LIBELLULOIDEA
1 Adult labium with palpal endhook aslongas ligula,ligulaentire; positronswith postocellar
ridge;supratrianglessimilarinshapeand withanteriorsideconcaveposteriorly;posteriorhamuli
Evolution ofGondwanianLibelluloidea 387
engagingfemale sternum 9, apicalhooks directed anteriorly; ovipositor reduced to fused
progonocoxae andprogonapophyses,metagonopoditesatmostrepresentedby smallsternalplates.
Larvalantennae3or4segmented,third segment morethan Viantennal length;second mandibn•
larsegmentmovable;raesotarsi 2-segmented;abdominal segments4or5to6with linear trans-
verse muscles and anterolateral sternal apodemes;sclerotized proventricularlobes elongate-
-rasplike,with 8-20 scatteredteeth Gomphoidea- Gomphidae
- Adultlabiumwithpalpalend hookshorterthanligula,ligulavariable;postfronswithoutpostocellar
ridge;supratriangleseither notsimilar in shapeorwith anterior side straight;posteriorhamuli
engaging femalesternum8,apicalhooksdirectedposteriorlyormedially;ovipositorcompleteor
reduced,metagonopoditesatleastrepresentedbysmallpeglikestructures.Larvalantennae6to8
segmented,thirdsegmentlessthanViantennallength;secondmandibularsegmentnotmovable;
mesotarsi 3-segmented;abdominal segments 4 and5with vestigialorphragmatictransverse ab-
dominal musclesand withoutanterolateral apodemes;sclerotizedproventricular lobes moundor
toothlike,with0-8posteriorlyclusteredteeth 2
2 Adultcompoundeyeswidelyseparateddorsally,anterodorsalsurfaceofocciputtrapezoidal;ptero-
stigmataconcaveposteriorly and longerthan distancebetween costal braces; ligulawith wide
medial cleft;maleepiprocttypically divaricate,andcerci stronglyexpandeddistally;ovipositor
complete and stronglyupturned.Larval tibiaewith apicalborrowinghooks;terminalia forming
dorsallydirectedvent; labial palpswithrobust dorsolateral spuratbase ofendhook;molarlobe
with teeth;transverse abdominal muscles 4and 5 vestigial;proventriculuswith 8sclerotized
lobes each with0-6 similar blunt teeth Petaluroidea - Petaluridae
- Adultcompoundeyes contiguousorapproximatedorsally.anterodorsalsurfaceofocciputtrian-
gular;pterostigmatanotconcaveposteriorly,shorterthandistance between costalbraces; ligula
withmedial cleftvariable;male epiprocttypicallyquadrateortriangular,and cerci notstrongly
expandeddistally;ovipositorvariable,notupturned.Larvaldbiaewithoutapicalburrowinghooks;
terminalia notformingdorsallydirectedvent;labialpalps withoutrobustdorsolateral spuratbase
ofendhook;molarlobewithout teeth;transverse abdominal muscles4or5 phragmaticorobso-
lete;proventriculuswith 4sclerotized lobes each with2-8 sharpteeth,apicaltooth largest 3
3 Adultpterostigmalbracethickened andoblique;anteriorlaminawith elongatemedial cleft,ante-
riorhamulidirectedmedially,posteriorhamuli vestigial;median processofmaleabdominal seg-
ment2shortL-shaped;ovipositorcomplete,suitedforendophyticoviposition.Larvalprementum
flat,dorsal surface oflabiumwithout longprementalandpalpalsetae,labrum notconcealedby
triangularlabialpalps,labial endhook distinctlylongerthanbaseofpalp;epiprocttypicallybi-
furcateapically;proventriculus radiallysymmetrical,sclerotized lobes small-lobelike with 8or
fewerclustered teeth Aeshnoidea - 4
- Adultpterostigmalbraceobsolete,(presentinNeopetalia);anteriorlaminawithoutelongateme-
dial cleft,anteriorhamuli directedposteroventrallyorabsent,posteriorhamuli well developed;
median process ofmale abdominal segment 2long J-shaped;ovipositor reduced, suited for
exophyticoviposition.Larval prementum scooplike,dorsal surfaceoflabiumtypicallywithlong
prementalandpalpalsetae,labrum concealedbytriangularlabial palps,labialendhook distinctly
shorter thanbaseofpalp; epiproctacuminate;proventriculusbilaterallysymmetrical,sclerotized
lobes laige-toothlike,edgedwith 2-8 teeth Libelluloidea - 5
4 Adultwingswithcostal seriesof5-8reddish blotches;compoundeyesapproximateormeetingat
pointdorsally; abdomen without dorsal orlateralcarinae,7 or8 often withlateral expansions;
wingswithout planates;fore wingtrianglewith proximalside morethan Vianteriorside; penis
laterallyexposed,prepucewell developed,segment4pendulouswith hugepairedflagellaedi-
rected posteroventrally.Larval abdominal segments 1-10with lateral lobes;prementum slightly
widened distally;labrumwideneddistallytocawidthofprementum;ventrolateraloccipitalridge
massive;paraprocts shorter than Viwidth ofabdominal segment9; femora dorsallyexcrescent;
transverse abdominal musclesobsolete;body surfaceextensively granulate...Austropetaliidae
- Adultwingswithout costal seriesof5-8reddish blotches;compoundeyesmeetingalongdorsal
388 F.L,Carle
seam;abdomen with dorsalandlateral carinae,withoutlateral expansions;wingswithplanates;
fore wingtrianglewith proximalsidelessthanVianteriorside;penislaterallyconcealed,prepuce
obsolete,segment4swablike withouthugepairedflagellaedirected posteroventrally. Larval ab-
dominal segments 3-9 atmost withlateral spines;prementum greatly widened distally;labrum
ca.Viwidth ofprementum;ventrolateral occipitalridgelow;paraproctslongerthanVi widthof
abdominal segment9; femoradorsallysmooth;transverse abdominal muscle 5 phragmatic, 6
linear; body surfacenotextensively granulate Aeshnidae
5 Adultwithsupplementarysectorarisingnearbridgecrossvein;paraglossalspinespresent; lateral
clypeallobes notinflated;occellar lobe vestigial;pterostigmataparallelsided,lengthtypically
ca.8 times width; maleprotibiaewith shortapical keels and meso- and metatibiae with outer
spinespeglike;anteriorhamuli largeerect-foliate;progonopoditesappressedintoelongategently
taperedspade. Larval cerci vestigial,less thanVimidventral lengthofabdominal segment 10;
vulvar laminamorethan 6/10lengthofsternum 9 Cordulegastridac
- Adult without supplementary sectorarisingnearbridgecrossvein;paraglossalspinesobsolete;
lateralclypeallobes inflated;ocellarlobe present;pterostigmatanotparallelsidedwithlengthca
8timeswidth;malewithoutsmallprotibialpadlikekeels andmeso-and metatibial peglikespines;
anterior hamuli vestigial orelongate-triangularwith incurved end hook; progonopoditesnot
appressedintoelongategentlytaperedspade.Larvalcerciwell developed,morethanVimidventral
lengthofsegment 10;vulvarlaminaless thanVilengthofsternum9
Non-cordulegaslridLibelluloidea
KEYTOTHE GENERAOF ANCIENT GONDWANIAN LIBELLULOIDES
(NEOPETALIIDAE, CHLOROGOMPHIDAE,SYNTHEMISTIDAE
AND GOMPHOMACROMIIDAE)
1 Adultwithpterostigmalbracethickenedandoblique;wingswithcostal seriesof4reddishblotches,
apical blotchdivided byyellowishorangepterostigmata; abdominal terga5-8with ventroapical
tuftsoflongblack hair;male tibial keelsca 1/3lengthofprotibiaeand 1/5lengthofmeso- and
metatibiae;anterior hamuli contiguous and L-shaped; female sternum 10expanded into huge
circularsplashplate.Larvallabialpalpibilobate with5-6 irregularmedial teeth;prementumwith
2-6 vestigialsetae;palpiwith 1shortdorsomedialsetaenearbaseofpalpalendhook;antennae6-
-segmented,thirdsegmentmorethantwicelengthofsecondsegment;antefrons moundlike;male
epiproctaltubercleacuminate;vulvar laminaca 1/3lengthofsternum9
Neopetaliidae- NeopetaliaCowley
- Adult with pterostigmalbrace vestigial; wingswithout costal series of4reddishblotches; ab-
dominal terga5-8 withoutventroapicaltuftsoflongblack hair;maletibial keels typicallymore
than 1/3 lengthofprotibiae and 1/5lengthofmeso- andmetatibiae (obsoletein Libellulinae);
anteriorhamuli notcontiguousorL-shaped;female sternum 10withoutsplashplate.Larval la-
bialpalpinotbilobate with 5-6irregularmedial teeth;prementumwith4-30elongatesetae;palpi
with 2-8 elongatedorsomedial setae;antennaetypically 7 segmented,third segment less than
twice lengthofsecond segment;antefrons notmoundlike;maleepiproctal tuberclenot acumi-
nate; vulvar laminanottypically 1/3lengthofsternum9 2
2 Adultligulalongerthanwideandwithapicalcleft;labialpalpswithwell developedmovable end
hook and apical spine; sectors ofthe arculus separatedatbase; supratrianglesslightlyconvex
anteriorly;antenodal crossveins ofhind wingnotaligned,with two costalbraces;subtriangular
interspacedilated basally;anteriorhamulielongate,withinterhamularspur. Larval labial palpi
withelongateirregularmedialteeth;ligulawithglossalandparaglossallobes;mesostemumwithout
paracoxallobes;metastemumwithtransversesulcijoinedby medial sulcus;ventralproventricular
sclerotized lobes without largesubapicaltooth,dorsal lobeswithout posteromedialedgestrongly
Evolution ofGondwanian Libelluloidea 389
inclined laterally;sternum6without anterolateral apodemes;epiproctapicallyneedlelike
Chlorogomphidae- 5
- Adult ligulawider than longand without apicalcleft;labial palps with vestigialmovable end
hookandapicalspine;sectorsofthearculusfusedbasally;supratrianglesstronglyconvexanteriorly;
antenodal crossveins ofhindwingmostlyalignedorwith3 ormorecostalbraces; subtriangular
interspacenot dilatedbasally;anterior hamulishortorobsolete,withoutinterhamular spur.Lar-
val labialpalpiwithout elongateirregular medial teeth;ligulawithdistal margin entire;meso-
stemumwithparacoxallobes;metastemumwithtransversesulci contiguous,medial sulcusobso-
lete; ventral proventricular sclerotized lobes with large subapicaltooth, dorsal lobes with
posteromedialedgestrongly inclined laterally; sternum6 with anterolateralapodemes;epiproct
notapicallyneedlelike 3
3 Adultmedian spacewith I-5crossveins; hindwingswith3-5costalbracesalternatingwithpoorly
alignedantenodals;wingswith3-10 cubital-anal crossveins; abdomencylindrical, segments 3-6
typicallymorethan 5timesaslongaswide,malesegment 7cylindricalwithoutwelldeveloped
middorsal ridgeorlateralcarinae. Larval ligularepresentedby amedialtriangularlobe; meta-
stemumwith transverse sulci meetingatpoint; wingpads divergent;hind femur ca 1.5times
lengthoffrontfemur;abdomen typically2.0-2.5timesaslongaswide Synthemistidae- 12
- Adult median space without crossveins; hindwingswithantenodal crossveins alignedandpro-
gressively morebracelike proximally; wingswith 1-4cubital-anal crossveins; male abdomen
triquetrous,atleastonsegments7 and8,segments 3-6 less than 5timesaslongaswide,tergum
7with well developedmiddorsal ridge orlateral carinae. Larval metastemum with transverse
sulci fused;wingpads typicallyparallel;hindfemurtypicallymorethan 1.6timeslengthoffront
femur;abdomen lessthan2.1times aslongaswide 4
4 Analloopelongatewithgaffshorterthan base;sectorsofarculusdivergingbasally;male without
genitallobe, male tergum 1 with ventrolateral hamulelike spines(obsoleteinPseudocordulia).
Larval labial palpiwithoutsetaealongmedialmargin;metastemumwithtransversesulci fusedat
short seam; dorsal proventricularlobes not united by medial chutelike sclerotization; ventral
proventricularlobeswith posteriorfacecaaswideasthatofdorsal lobes; ventralproventricular
lobes with teeth separatedbycatwice widthofposteriorfaceofdorsallobes;lateral prothoracic
lobes shelflike Gomphomacrotniidae- 20
- Anal looprounded orelongatewithgafflongerthan base;sectors ofarculusoften stalked;male
with genitallobe and without hamulelike ventrolateral spinesonsegment 1.Larval labialpalpi
typicallywithsetaealongmedialmargin;metastemumwithtransverse sulcitypicallyfusedalong
longseam; dorsal proventricularlobes united by medial sclerotization; ventral proventricular
lobes with posterior faceVi, orless thanas wide asthat ofdorsal lobes;ventral proventricular
lobes typicallywith teethseparatedbyless thanwidthofposteriorface ofdorsal lobes;lateral
prothoracic lobes notshelflike macromiid-corduliid-libellulid complex
5 Dorsalmesanepistemalyellowstripeswedgeshaped,widerdorsally;lateralmesanepistemalstripes
absent;mesanepiraeralyellowstripeswell developed;metanepistemalstripes vestigialorabsent;
metanepimerawithwide yellowstripe;antefronsslightlyinflateddorsally,withdorsolateralcarinae
well developed;compoundeyesseparatedbyca 1/6widthofmedial ocellusinmale,andbyca 1/3
width infemale;male cerci dorsoventrallyflattened,ca3timesaslongaswide,withouteredge
evenlycurvedwithmidventral spine,andwithinnerapicaltoothacute anddirectedposteromedially;
anteriorhamuli directedposteroventrally,without well developedposterobasal lobe;ventroapical
penilelobes(cornua)elongate,morethan6timesaslongaswide;vulvarlaminaaslongaswide
withmidventral keel.Larvalligulaflattened withglossaandparaglossafusedintopairedrounded
medial lobes;palpal endhook withdorsal rowofsmall setae;ventral proventricularteeth with
scattered dorsal denticles;dorsal abdominal setaepeglike; abdominalsegment9without lateral
carinae Chloropetaliinaesubfam.n.,typegenusChloropetaliagen.n.,
typespecies ChlorogomphusselysiFraser; alsoincludes Orogomphusatkinsoni Selys, O.dyak
Laidlaw, and C.olympicusFraser
390 F.L.Carle
- Dorsal mesanepistemalyellowstripes linear;lateral mesanepistemalyellowstripeswell devel-
oped; mesanepimeralstripes narrow orabsent;metanepistemalpale stripes well developed;
metanepimeralstripes narroworabsent;antefrons stronglyinflated dorsally,withdorsolateral
carinae vestigialorabsent;compoundeyesseparatedby atleast 1/3 widthofmedial ocellus in
male, and by atleast 2/3width in female;male cerci notas above; anterior hamuli directed
ventrally, with well developedposterobasal lobe;ventroapical penilelobes stout, less than 4
times as longas wide, vulvar lamina wider than longwithout midventral keel. Larval ligula2
scooplikewithglossaandparaglossadistinct;ventral proventricularteeth withdenticles limited
totooth edges; dorsal abdominal setaehairlike;abdominal segment9with lateralcarinae
Chlorogomphinaecomb.n.,type genusChlorogomphusSelys- 6
6 Anterior halfofmesanepimeraand metanepiraerawith yellowstripe; labrum with submedial
paleareas; male compoundeyes separatedbyca2/5widthofmedianocellus;male hindwingas
longasabdominal segments 1-6,hind wingto abdominal lengthratio ofmalegreater than 1.40,
thatoffemalegreater than1.23;malecerci ca2.5timesaslongaswide,withouteredgeabruptly
curvedsubapicallywithout midventral spine, andwith inner apical toothblunt anddirected me-
dially Eorogomphinitrib.n.,type genusEorogomphusgen.n.,
typespeciesO.preciosusFraser; alsoincludes O.speciosus Selys, C.fernandiAsahina,and C.
schmidti Asahina
- Anterior half of mesanepimerawithout orwith a vestigial yellow stripe, anterior half of
metanepimerawithout yellowstripe;labrum predominantlybrown or black; malecompound
eyes separatedby atleast 2/3 width ofmedial ocellus; male hind wingaslong asabdominal
segments 1-8orlonger,hindwingtoabdominal lengthratio ofmale lessthan 1.32,that offemale
less than 1.22;male cerci notasabove 7
7 Dorsal mesanepistemalpale stripes dorsallyexpanded into transverse oval; meso-and meta-
katepistemawithyellowspots; abdominal tergum 3morethan twiceashighbasallyasapically;
femaleabdominal segments 3-7 subcylindrical,segment4more than 3times aslongas high;
femaleabdominal segment6 or7with pairedapicalyellowspots caaslongas wide;hindwing
trianglewithcostalsidelongerthanproximalside;hindwingtoabdominal lengthratiooffemale
more than 1.13; first penilesegment without medial groove; wingstypically withoutopaque
markings Sinorogomphinitrib.n.,typegenusSinorogomphusgen.n.,
typespecies C. nasutusNeedham;alsoincludes C. tuntiNeedham,C. infuscatusNeedham,O.
suzukii Oguma,C.kitawakii Karube,and C. urolobatus Chen
- Dorsal mesanepisternalpale stripes not dorsally expanded into transverse oval; meso-and
metakatepistemabrown orblack; abdominal tergum 3 less than 1.7times as highbasallyas
apically in male, less than 1.5times in female;female abdominal segments 3-7 laterally com-
pressed, segment4less than 3times aslongas high;female abdominal segment 6 and 7 with
pairedapicalyellowspotslessthan 1/2aslongaswideorobsolete;hind wingtrianglewithcostal
side shorter than proximalside orsubequalinmale; hind wingtoabdominallengthratio of
female less than 1.13; firstpenilesegment with medial groove; wings typically with apical or
basal opaquemarkings Chlorogomphinicomb.n. - 8
8 Labrum with U-shapedcentral greenishyellowspot; lateralmesanepisternalyellowstripesnar-
row, only slightly wider thandorsal mesanepisternalyellowstripes; basal abdominal segments
withbroadlateralobliqueyellowstripeextendingfromventraledgeofmetanepimerato antecostal
sutureofsegment3;abdominal tergum7 withpairedbasal yellowspotscaaslongaswide;male
hindwingto abdominal lengthratio 1.20-1.26;male withbases ofabdominal segments 8 and 9
constricted basally;cerci reduced, in female ca 1/2 length oftergum 10,in male with blunt
basolateral spine;maleepiproctstronglyupcurvedapically Neorogomphusgen.n.,
2 Larval characters basedonFRASER’s(1929)supposedlarvaofC. campioni.However, larvalchar-
acters described by MATSUKI, etal. (1995) forC. brunneus do not agree. IfFraser’s larvae are
cordulegastrid,listed characters arelikelydiagnosticforChlorogomphidae.
Evolution ofGondwanian Libelluloidea 391
typespecies C.fraseriSt.Quentin; also includes C. auratusMartin, C. arooniAsahina, and
vietnamensisAsahina
- Labrum black without U-shapedcentral greenishyellowspot; lateral mesanepistemalyellow
stripeswide, atleast 4timesaswideasdorsal mesanepistemalyellowstripesatmidlength;basal
abdominal segments with broad lateral obliqueyellowstripeextendingfrom ventral edge of
metanepimeratoapexofsegment2only;abdominal tergum7 withoutpairedbasal yellowspots
caaslongas wide; male hind wingto abdominal lengthratio 0.95-1.19;male withbases of
abdominal segments8 and 9constricted basally;cerciwell developed,infemale ca2/3lengthof
tergum10,in malewithoutblunt basolateral spine;maleepiproctnotstronglyupcurvedapical-
ly 9
9 Dorsal and lateral mesanepistemalpalestripes confluent dorsally; antefrons bilobate,without
well developedventrolateral carinae; male hind wingtrianglewith proximalside subequal to
costal side;male abdominal segment2longerthan high;abdomen tohind winglengthratio of
male ca 1.16,of female ca 0,97; anterior hamuli directed ventrally, end hooks directed
posteromedially,separatedapicallybycalengthofhamuli;fourthpenilesegment elongatewith
lateral keels formingacuteapex; anterior laminaslightlybilobate;male cerci withlateral spine
subequal toapicalspineand located justbeyondmidlength;male epiproctwith deepwide U-
-shapedapical notch Indorogomphusgen.n.,
typespecies O.xanthopteraFraser; alsoincludes O.campioniFraser
- Dorsal andlateral mesanepistemalpalestripes notconfluent dorsally;antefrons flattened,with
well developedventrolateral carinae; male hind wing trianglewith proximal side longerthan
costal sidein male;male abdominal segment2shorterthan high;abdomentohindwinglength
ratio ofmale and female atmost 12% different;anterior hamuli directed posterolaterally,end
hooksdirectedmedially,separatedapicallybymorethanlengthofhamuli;fourthpenilesegment
short withlateral keels formingobtuselyrounded apex; anterior laminastronglybilobate; male
cerciwithlateralspinelargerthanapicalspine andlocatedsubapically,spinesoccasionallyobso-
lete;male epiproctwithoutdeepwide U-shapedapicalnotch 10
10 Metastemum black medially;posterior third ofmetanepimeraabove ventrolateral carinae pre-
dominantlyyellow;submedial apicalabdominal spots increasingin size on segment7; male
abdominal segments 4-7 cylindrical;male hindwingto abdominal lengthratio morethan 1.08;
male epiproctwithsmall apical notchandapicallytruncaterami;female occiputnotdepressed;
female abdominal tergum 3with ventrolateral stripenotconfluent with apicalring;female ab-
dominal segments5and6morethantwotimesaslongashigh Orogomphus Selys,
typespecies O.splendidusSelys; alsoincludes C. brevistigmaOguma,C. okinawensisIshida,C.
brunneus Oguma,C. costalisAsahina,C.keramensis Asahina,C. iriomotensis Ishida,and C.risi
Chen
- Metastemum yellowmedially;posteriorthird ofmetanepimeraabove ventrolateral carinae nar-
rowly yellow;submedial apical abdominal spots decreasinginsizeonsegment7; maleabdomi-
nal segments 4-7 triquetrous;male hind wingto abdominal lengthratio less than 1.08; male
epiproctnotasabove; female occiputdepressed;female abdominal tergum 3with ventrolateral
stripe confluent withapicalring;female abdominal segments5 and6lessthantwotimesaslong
ashigh 11
11 Anal loopwith28-35cells;fore andhindwingswithextensivebasal opaquemarkings;malewith
posterobasalwingmarginslightlyangulate;maleand femaleabdomen tohind winglengthratio
dissimilar;female abdominal segment 4ca 1.5times aslong ashigh; lateral mesanepistemal
yellowstripesobsolete;male cercitriangular,basallyexpanded,withwell developedlateral and
subapicalspines;male epiproctdivaricate withshallow U-shapedapicalnotch
Aurorachlorus gen.n.,
typespeciesC.papilioRis
- Anal loopwith 8-17 cells; male wings and female fore wingswithout basal opaque markings;
male withposterobasalwingmarginevenlyrounded; maleand female abdomen tohind wing
392 F.L.Carle
lengthratio similar; female abdominal segment 4 ca 2.0 times aslong ashigh; lateral
mesanepistemalyellowstripeswell developed;male cercielongate, laterallycompressed, with-
out lateral orsubapicalspines; male epiproctnarrowed subapicallywith divaricate apices, and
with V-shapedapicalnotch Chlorogomphus,
typespeciesC. magnificusSelys; also includes C. mortoniFraser
12 Wingswith spots atnodus;fore wingarculus with posteriorportionca 1/3itslength;hindwing
triangle andanal loopelongate,base ofanal loop,distal side oftriangle,and gaffsubequal;
proximalsideofhind wingtriangleperpendiculartolongitudinalveins,distalside morethan 1.5
times lengthofproximalside;maleepiproctwithwideV-shapedapicalnotch;prepuce caaslong
asbase ofpenilesegment3;ovipositorknoblikeapically
Synthemiopsinitrib.n.,type genusSynthemiopsis,
monotypicS. gomphomacromioidesTillyard
- Wingswithout spots atnodus; fore wingarculuswithposteriorportionless than 1/3itslength;
hind wingtriangleandanalloopnotelongate,base ofanalloop,distalside oftriangle,andgaff
unequal;proximal sideofhind wingtriangleobliquetolongitudinalveins, distal side less than
1.5timeslengthofproximalside;male epiproctwithoutwideV-shapedapicalnotch;prepuce at
most 1/3aslongasbase ofpenilesegment 3;ovipositornotknoblike apically 13
13 Midbasal spacetypicallywith 1 crossvein;posteriorhamuli ofboxingglovetypewith denticles
at apex ofthumb andalongknuckles; abdominal segment1 ofmale with spinoseventrolateral
spurs; penilesegment 2gentlycurvedbasallyandlongerthansegment 1;penilesegment3 at
least twiceaslongaswide;ovipositorextendedbeyondcerci;larvaeasfarasknown withtrilobate
labial palps Palaeosynthemistinitrib.n.,type genus PalaeosynthemisFörster- 14
- Midbasal space typically with 2-5 crossveins; posteriorhamuli not ofboxingglovetypewith
denticles at apex ofthumb andalongknuckles; abdominal segment 1 ofmale without spinose
ventrolateral spurs;penilesegment 2abruptlybent basallyand shorterthansegment I; penile
segmentthree lessthantwice aslongaswide;ovipositornot extendedbeyondcerci,reduced to
vulvar laminae;larval labial palps with medial marginentireorwith small rounded teeth 15
....
14 Pterostigmataaslongashind wingsupratriangle;fore wingtriangledistant from arculusbyless
than lengthofproximalside oftriangle; abdominal segments 3-8 with pairedsubmedian oval
spots subequal and dividedby antecostalsuture; penilefilament shorter than penilesegment
1 Archaeosynthemis gen.n.,
typespeciesS.leachiiSelys; alsoincludesS.macrostigmaSelys, andS.spinigerTillyard
- Pterostigmata shorterthanhindwingsupratriangle;fore wingtriangledistantfromarculus by
morethanlengthofproximalsideoftriangle;abdominalsegments 3-7 withtransverse dashes
alongantecostal suture;penile filament longerthanpenilesegmentI
PalaeosynthemisForster,
type speciesP.primigeniaForster;alsoincludes S.alectoLieftinck,S.cervulaLieftinck,S.feronia
Lieftinck,S.cyreneLieftinck,S.evelynaeLieftinck,S.gracilentaLieftinck, S.wollastoniCampion,
andS.kimminsi Lieftinck
15 Hindwingwith 3-5 midbasal crossveins; posteriorhamuli without posteromediallydirectedend
hook,hamuli claspingvulvar laminawithventro-dorsalaction;base ofpenilecornuaexpanded
into medial septumlike plate; penilefilament directed dorsally at base;male cerci without
basolateral spine;vulvar laminabilobate and atleast 1/2aslongassegment 10
Synthemistinicomb.n.,typegenusSynthemisSelys- 16
- Hind wingwith 1-2midbasal crossveins;posteriorhamuli with posteromedially directed end
hook, hamuli not claspingvulvar lamina with ventro-dorsal action;base ofpenilecornuanot
expandedinto medial septumlikeplate;penilefdament directed posteriorlyatbase; malecerci
withbasolateralspine;vulvar lamina vestigialorappressedandshorter than 1/2aslongasseg-
ment 10 Eusynthemistinitrib.n.,type genusEusynthemisForster — 18
16 Sectors ofthearculus arisingat 1/4-1/3lengthofarculus;posteriorhamuli chisellike withapical
transverse row ofdenticles;ventrolateral marginofmaleabdominal tergum2with coarsesetal