Morphology of female external genitalia inPhenes raptor(Odonata: Petaluridae)

2011·0.38 MB·

by  Natalia A. Matushkina

InternationalJournalofOdonatology
Vol.14,No.3,September2011,199–215
Morphology of female external genitalia in Phenes raptor
(Odonata: Petaluridae)
NataliaA.Matushkinaa*andKlaus-DieterKlassb
aDepartmentofZoology,BiologicalFaculty,KyivNationalUniversity,vul.Volodymirs’ka64,Kyiv,01033,
Ukraine;bMuseumofZoology,SenckenbergNaturhistorischeSammlungenDresden,Königsbrücker
Landstrasse159,01109Dresden,Germany
(Received17March2010;finalversionreceived4February2011)
TheexoskeletonofthefemalegenitalicregioninPhenesraptorisdescribedbasedonlightmicroscopy
andscanningelectronmicroscopy.Itisshownthatinthisspeciesthepatternofsclerites,articulations,
processes,andapodemesisoverallthesameasinotherovipositor-bearingOdonata,i.e.Zygoptera,the
anisozygopteranEpiophlebia,andtheanisopteranAeshnidae.However,manymorphologicaldetailsdiffer
amongallthesetaxa.Fifty-fourcharacterswerescoredforP.raptorinordertobeincludedinapreviously
compileddatasetforphylogeneticanalysisofovipositor-bearingOdonata.Thesecharactersincludeonly
fewspecificsimilaritiesbetweenP.raptorandeitherAeshnidaeorEpiophlebia.Instead,P.raptorshows
a number of features that are unique among ovipositor-bearing Odonata.Absence of serration on the
ovipositorinP.raptorandreductionoftheinterlockingmechanismconnectingthetwofirstvalvesmedially
isprobablycorrelatedwiththeendosubstraticegg-layingofthefemale.Theovipositorbearsnumerous
sensillaofdifferentshape,whichprobablydetectsuitableplacesforoviposition.
Keywords: Odonata;dragonfly;morphology;ovipositor;Phenesraptor;Petaluridae;Anisoptera
Introduction
Extant Odonata are generally subdivided into three groups: the species-rich Zygoptera and
Anisoptera,andtheAnisozygoptera,withthesoleextantgenusEpiophlebia.Zygopteraareviewed
either as paraphyletic with respect toAnisoptera andEpiophlebia (e.g. Pfau, 2002; Saux et al.,
2003;Trueman, 1996), or as a monophyletic group (e.g. Bybee et al., 2008; Carle et al., 2008;
Dumontetal.,2010;Flecketal.,2008;Rehn,2003).Asister-grouprelationshipofEpiophlebia
and Anisoptera (together called Epiprocta) is recently strongly favoured (e.g. Dumont et al.,
2010;Flecketal.,2008;Rehn,2003)butnotuniversallyaccepted(e.g.Carleetal.,2008).The
monophylyofAnisopteraisbeyonddoubt,beingsupportedbothbymorphologicalandmolecular
data (e.g. Dumont et al., 2010; Rehn, 2003). Phylogenetic relationships among families within
Anisopteraremaindisputed(e.g.Bechly,1996;Bybeeetal.,2008;Carle,1982,1995;Fleck&
Nel,2003;Lohmann,1996;Misofetal.,2001;Pfau,1991,2005;Trueman,1996).
*Correspondingauthor.Email:[email protected]
ISSN1388-7890print/ISSN2159-6719online
©2011WorldwideDragonflyAssociation
http://dx.doi.org/10.1080/13887890.2011.607735
http://www.tandfonline.com
200 N.A.MatushkinaandK.-D.Klass
Odonatashowawiderangeinthemorphologyoftheirfemalegenitalia.Ahighlycomplicated
female genitalic region that includes a well-developed ovipositor is found in all Zygoptera, in
Epiophlebia, and in the Aeshnidae, Petaluridae (see St. Quentin, 1962), and Austropetaliidae
among theAnisoptera (“ovipositor-bearing Odonata”).While most species from these taxa lay
theireggsintolivingplanttissue(endophyticoviposition),thePetaluridaeandsomeAeshnidae
useanotheregg-layingstrategy(seebelow).InformationontheovipositionoftheAustropetaliidae
is unavailable.Anisopteran taxa apart from the aforementioned ones have the female genitalia
stronglymodified(suchasCordulegastridae)orgreatlysimplified(suchasLibellulidae)(seee.g.
Asahina,1954).
Despitethepotentiallygreatphylogeneticinformationcontentofthefemalegenitalicregion,
phylogeneticreconstructionsofOdonatahavenotyetconsideredthischaractersysteminmuch
detail.Therefore,Klass(2008)compiledadatasetwhere79charactersofthefemaleabdomen,
mostlythegenitalicregion,arescoredforasampleofovipositor-bearingOdonata:13zygopter-
ans,twoaeshnids,andEpiophlebiasuperstes.Thepossiblephylogeneticevidencefrommanual
evaluation of these characters was discussed, but there was much incongruence. Matushkina
(2008a)performedaverypreliminaryphylogeneticanalysisof70oviposition-relatedcharacters
in 33 species of Odonata, including 28 zygopterans, four aeshnids, and Epiophlebia superstes;
theresultssuggestthatoviposition-relatedcharacterssupportsomeofthetraditionalgroupings
ofOdonata.Duringthelastfiveyearstheauthorsofthisworkstartedtocollaborateinthefield
of genital morphology of insects; one of their goals is to establish a comprehensive dataset on
femalegenitaliainOdonata.ThePetaluridaeisoneoftheovipositor-bearingodonatanfamilies
stillmissinginthisattempt.
Petaluridsarelargeanisopterandragonflies,distributedinJapan,NewZealand,Australia,Chile,
SWArgentina,andNorthAmerica(Garrisonetal.,2006;Steinmann,1997).Adultsinhabitbog
andwoodlands,wheretheyperchontreesorflyoverstreams(Garrisonetal.,2006).Thefemales
lay eggs into fairly loose substrate such as moss, among grass roots, into decaying vegetable
matter,orintowetground(“endosubstraticdeposition”;Garrisonetal.,2006;Winstanley,1981).
Thelarvaeshowanobligatorysemi-terrestriallifestyle,whichisuniqueamongextantdragonflies.
Theyliveinburrowsinstreambanksorindepressionsunderwetleavesanddamppiecesofwood
(Garrisonetal.,2006;Winstanley,1981;Winstanley&Rowe,1980).
ThefamilyPetaluridaeiscomposedof11extantspeciesclassifiedintofivegenera.Theknown
fossilrecordstartsc.160millionyearsagointheJurassic(Neletal.,1998).Petaluridaeisgenerally
recognisedasamonophyleticgroup,butitsplacementwithintheAnisopteraisquitecontrover-
sial. Several authors considered Petaluridae as the sister group to the rest of Anisoptera (e.g.
Bechly,1996;Rehn,2003;Trueman,1996),whileotherauthorsplacedpetaluridswithindiffer-
entanisopteransubgroups(e.g.Bybeeetal.,2008;Carleetal.,2008;Lohmann,1996;Pfau,1991,
2005).Indeed,thepetaluridsarecharacterisedbyacomplexmosaicofplesiomorphicandapo-
morphicfeatures.St.Quentin(1962)regardedtheovipositorofPetaluridaeasthemostprimitive
in Odonata. However, Pfau (1991) emphasised that the ovipositor of Petalura sp. is only “rel-
atively primitive”, being intermediate between the endophytic zygopteroid-aeshnoid ovipositor
andthemoderatelyreducedanddistinctlymodifiedcordulegasteroidovipositor.
This study aims to thoroughly describe the general construction and the microsculpture of
theovipositorinthepetaluridPhenesraptor Rambur,1842,andtocomparethisdatawithcur-
rentinformationonotherovipositor-bearingOdonata(Klass,2008;Matushkina,2004,2008b,c;
Matushkina & Gorb, 1997, 2002). Characters related to the ovipositor and some other parts
of the female postabdomen will be scored for Phenes raptor in order to be included into
the dataset for phylogenetic analysis compiled recently by Klass (2008). The results of this
study will fill the most striking gap in Klass’s and Matushkina’s datasets and represent an
important step towards establishing a female genitalia character matrix for analysing odonatan
relationships.
FemalegenitaliaofPhenesraptor 201
Materialandmethods
TerminalabdominalsegmentsoftwofemalesofPhenesraptorwereexamined.Driedpostabdom-
inaweresoakedwithBouin’sfluidforafewmonthsforsofteningandelastificationofthecuticula.
Thenpostabdominawerewashedindistilledwateranddissectedinthemedianplane.Thetwo
halvesweremaceratedin10%KOH,andexaminedinglycerineunderastereomicroscope.For
the scanning electron microscope (SEM) study the cuticular parts of one female were washed
indistilledwater,dehydratedinagradedethanolseries,air-dried,sputteredwithgold-palladium
andexaminedwiththescanningelectronmicroscopesLEO1530VP,HitachiS-4800andZeiss
EVO-50SEM.
Morphologicaltermsandabbreviationsfollowtheinsect-wideapproachinKlass(2008)(see
also e.g. Klass, 2003; Klass & Ulbricht, 2009), with few additions marked with an asterisk in
Abbreviations.TermsbyKlass(2008)aregiveninstandardtype;correspondingtermsusedby
NMinpreviouspublicationsaregiveninparentheses.
Abbreviations
ac9,ac10 antecostaofS9(anteriorcarinaof9thtergite,partly),andantecostaofS10,
respectively
ag accessorygland
al aulax (ventral part of tongue-and-groove interlocking mechanism
on gonapophyses of S8 and S9 called the olistheter): groove on
gonapophysisofS8
b farposteriorlateralexpansionoftergite8
ba8 posteriorlateralapodemeoflaterocoxosternumofS8
ca,CA centralapodemebetweencoxaeofS9anditssclerotisedpart,respectively
(internalsclerite,orposteriorintervalvula)
CX8 coxaofS8(medialpartofbasalplateofovipositor)
CX9 coxaofS9(sclerotisationofpairedthird,orlateralvalveofovipositor)
da anterodorsalapodemeoftergumofS9,onanteriorsegmentalborder
dl dorsallobeofgonoplacofS9
dt10 dorsaltendonofantecostaofS10
et10 ventrolateraltendonofantecostaofS10
fa anterolateralapodemeofcoxaofS9(anteriorapophysisofthirdvalvesof
ovipositor)
fcs (*) field, or cluster of campaniform sensilla on ventromesal surface of
stylusofS9
fhs (*)fieldofshorthairsensillaonexpandedbaseofgonapophysisofS9
ft tendonsflankingcentralapodemeca,betweencoxaeofS9
g anteroventralextensionofscleriteofgonapophysisofS9,bearinganterior
endofrhachis
ga anteriorapodemeofcoxaofS8
gl9 gonoplacofS9(thirdvalveofovipositor)
gp8,gp9 gonapophysesofS8andS9(firstandsecondvalvesofovipositor)
GP8,GP9 scleritesofgonapophysesofS8andS9
gt anteriortendonofcoxaofS8,mesaltoapodemega
j (*) ventral extension of antecosta of S9, connecting tergite TG9 with
anteriorpartoflaterocoxaofS9
la posterolateralapodemeofcoxaofS9(posteriorapophysisofthirdvalves
ofovipositor)
202 N.A.MatushkinaandK.-D.Klass
LCa9 anteriorpartoflaterocoxaofS9(anteriorpartofgonangulum,lateralpart
ofbasalplateofovipositor)
LCp9 posteriorpartoflaterocoxaofS9(posteriorpartofgonangulum,elongated
sclerite)
LS8 laterocoxosternumofS8(sterniteofS8)
pa8 anteriorlateralapodemeoflaterocoxosternumofS8(‘sternal’apophysis)
PS9 poststernumonventralhindmarginofS9
rh rhachis (dorsal part of tongue-and-groove interlocking mechanism on
gonapophysesofS8andS9calledtheolistheter):ridgeongonapophysis
ofS9
sl9,SL9 stylusofS9anditssclerotisation,respectively(stylus)
ta (*)tendonuponcentralapodemeca
TG8,TG9 tergaofS8andS9,respectively
TG10+LP tergumofS10fusedwithsurmisedanterolateralscleritesof“paraproct”
(S10)
ty7,ty8,ty9 transverse external ridge on posterior part of tergum of S7, S8 and S9,
respectively
Results
DescriptionoffemalegenitalicregionofPhenes
Theanteriorpartoftergite8(TG8)isnormallysclerotised.Theshortposteriorpartbehindthe
transverse ridge with a row of denticles ty8, however, is much more weakly sclerotised – very
weakinthemedianportionbutmoderatelystronginthelateralportions.Theheavierlateralparts
aresomewhatexpandedposteroventrally(i.e.aweaklydevelopedextensionbispresent)butthey
do not reach the articulations between the tergite of S9 (TG9) and sclerites LCa9 (Figures 1a,
2a,c).
On the ventral side the long anterior part of S8 bears a large uniform sclerite, the laterocox-
osterniteLS8;thisisusuallycalledthesterniteofS8,butmostofitconsistsofthelaterocoxal
sclerotisations of this segment, while a true sternal component may be included but cannot be
demonstrated (Klass, 2008, section 6.4.1, figure 63). The anterior apodeme pa8 as well as the
postspiracularapodemeba8arestronglydeveloped(Figure1b).
TheventralborderareabetweenS8andS9bearstwopairedscleriteswhichareseparatedby
anarrowribbonofmembrane,withoutanyarticulation,andconnectthebasesofgonapophyses
of S8 with laterocoxosternite LS8 and tergite TG9 (Figures 1, 2a). The plate-like anteromesal
sclerites are the coxae CX8. Left and right CX8 are fully separated.Along its anterior margin
each CX8 forms a shallow transverse apodeme ga. The mesal part of apodeme ga, which is
demarcatedbyasmallnotch,bearsatendongt;thetendonissclerotisedbasally(partofCX8)
andmembraneousapically(Figure2a,b).Theposteriormarginandespeciallytheposteromesal
cornerofscleriteCX8isinclosecontactwiththesclerotisationGP8ofgonapophysisgp8.The
posterolateralscleritesaretheheavyantelaterocoxaeLCa9,whichbelongtoS9.Dorsolaterally
eachLCa9splitsintoalevelanteriorextensionandintoanarrow,thickenedposteriorextension
(j,thethickeninglikelyrepresentsaventralpartofantecostaac9);thelatterissomewhatmovable
relative to the remainder of LCa9 and eventually continues into the tergal part of the antecosta
of S9 (Figure 2a). The mesoventral tip of LCa9 forms a slightly movable articulation with the
sclerotisationofgonapophysisgp8(ateinFigure2a).
The gonapophyses of S8, the gp8 (first valves), are curved dorsad, with sub-acute tips and
slightly expanded bases (Figures 1, 3a, c, d). Basally, the anterior end of the cleft between the
gp8ofthetwosidesisroughlyatthelevelofthearticulationbetweenscleritesGP8andLCa9
FemalegenitaliaofPhenesraptor 203
Figure 1. FemalepostabdomenofPhenesraptor:(a)lateralview;(b)medialview.Greyscalecorrespondstodegree
ofsclerotisation.Arrowheadin(a)marksventralbulgeonposteroventralwallofS10.
(at e in Figure 2a). The sclerotisations GP8 of the left and right gonapophyses 8 are entirely
separatedfromeachotherattheirbases,bothdorsally(Figure2b)andventrally(Figure2a);there
isalsonomediansclerite(cf.MSinKlass,2008,e.g.figure32)betweenthemattheventralbase.
At its sclerotised mesal base, each gonapophysis gp8 is folded dorsally and laterally (part x in
Figures1b,2a,b,d).Thispartofthegonapophysisissclerotised(partofGP8)andhasasmooth
mesal surface, i.e. interdigitating ribs for keeping the two gp8 together (cf. gy in Klass, 2008,
e.g.figures29,36)areabsent.ThedorsalbasalmarginofsclerotisationGP8isnotstraight,but
thereisadeepmembraneousnotch(inFigure2bthismembraneisretainedontherightsidebut
cutoutontheleft).Theentireexposed(i.e.ventralandlateral)surfaceofgp8isalsosmooth,i.e.
anycuticularridgesorteeth(suchastm8andgz8inKlass,2008)areabsent;however,thereare
numerouscoeloconicandcampaniformsensillascatteredovertheentiresurface(morefrequent
apically;Figure4).Theventromesaledgeofthebasalpartofgonapophysisgp8showsasparse
vestitureoflonghairs.Thedorsolateraledgeofeachgp8formsalongitudinalgroove(aulaxal)
thatprovidesaslidinginterlockwiththerhachisofthegonapophysisofS9,thegp9(Figure2b).
204 N.A.MatushkinaandK.-D.Klass
Figure 2. DetailsoffemalepostabdomenofPhenesraptor,diagrammatically:(a)segmentalborderareaS8–9ofleft
side,ventralview,ladder-likeribbonsrepresentinternalridgesbasedonthickeningofcuticle;(b)coxaeCX8andbases
ofgonapophysesgp8,dorsalview;(c)rightgonoplacgl9andelementsarounditsbase,dorsalviewwithgonoplacforced
dorsolaterad,blackspotindicatesorificeofcommonductofaccessoryglands;(d)basalpartofleftgonapophysealsclerite
GP8,ventralview,surroundingmembranesnotshown.Greyscalein(c)correspondstodegreeofsclerotisation.
Theanteriorendoftheaulaxislocatedonashortbasalextension(f)oftheGP8sclerotisation,
andimmediatelylateraltothispoint,scleriteGP8formsalateralbasalextension(e)bearingthe
GP8-LCa9articulation(Figure2a,b,d).
Tergite9bearsaheavyridgenearitsanteriormargin,thedorsalantecostaac9;uponac9there
isananterodorsalapodemedaandatendonet9(bothveryweaklydeveloped),andmostventrally
ac9 forms an anteriorly directed extension c.When the transition area between antelaterocoxa
LCa9andTG9isartificiallybroken,athinbutfirmcuticularfibreisdetachedatthelateralend
of extension j; this probably represents the strip called h in Klass (2008, e.g. figures 26, 27)
(Figure2a),whichmarkstheborderbetweenLCa9andTG9.TheshortpartofTG9behindthe
transverserowofdenticlesty9isaltogetherweaklysclerotised,butthesclerotisationissomewhat
heavier in the lateral portions of this area (Figures 1, 2c). The posteroventral corners of these
sclerotisationsaresomewhatexpandedtowardstheposteriorbasesofthegonoplacs(sheathing
valves;seethepartofTG9showninFigure2c).
ThegonapophysesofS9(gp9,secondvalves)arecurveddorsad–incontactwithgonapophyses
gp8–andhaveratherblunttips(Figures1,3a,c,d);thetwogp9areseparatedfromeachother
down to their very base, and there is also no connection between the sclerotisations (GP9) of
the two gp9.The ventrolateral edges of the gp9 form a longitudinal rail, the rhachis rh, for the
slidinginterlockwiththeaulaxalofgonapophysesgp8.Theanteriorendoftherhachis–placed
onasmallextensiongofscleriteGP9–islocateddistinctlyposteriortotheanteriorendofthe
gonapophyseal sclerotisation GP9 (Figure 2c). The pair of accessory gland (ag) ducts is, near
the opening, fused into a common duct which opens at midline at the level of the base of gp9
(Figures 1b, 2c) – a condition so far not reported for ovipositor-bearing Odonata.The exposed
(i.e. dorsal and lateral) surfaces of gonapophyses gp9 are smooth, with numerous sensilla of
different shapes, the density of which increases apically (Figure 4). The base of each gp9 is
FemalegenitaliaofPhenesraptor 205
Figure 3. ScanningelectronmicrographsoftheovipositorofPhenesraptor:(a)lateralviewofgonapophysesS8and
S9–insetshowsfieldofshorthairsensilla;(b)stylusofS9–insetshowssinglecampaniformsensillum;(c)mesalview
ofgonapophysesinapicalarea;(d)ventralviewofdistalpartsofgonapophysesandgonoplacsofS9;(e)lateralview
ofventraledgeofgonoplacofS9;(f)ventrolateralviewofventralbulgeofS10–insetsshowcampaniformsensillaof
differentshape.d:distaldirection.Arrowheadin(b)markssensillumenlargedininset.Scalebarsofinsets:(a)8μm,(b,
fbelow)2μm,(fabove)3μm.
slightlyexpanded,andontheproximalhalfoftheexpansionbearsafieldfhsofshortsetaeupon
itsexposedsurface(Figure3a).
Thegonoplacsgl9(sheathingorthirdvalvesofS9)haveatransversedepressiondemarcating
theproximalfromthedistalparts,whicharethusbothabitconvex(Figures1,2c).Thedistalparts
of the gonoplacs cover the tips of both pairs of gonapophyses laterally in the resting position.
All the exposed externally visible parts of the gonoplacs gl9 are sclerotised by the large main
scleriteofcoxa9,CXa9(Figure2c),butthecentrallateralandthedistalportionsaredistinctly
weakened(seelighterareasinFigure 2d).Thetipofthegonoplacformsaslightlyset-offlobe
(dl in Figure 2c), which is weakly sclerotised dorsally; this sclerotisation is fully continuous
with sclerite CXa9, while part of it likely corresponds to sclerite CXb9 of other Odonata (cf.
Klass,2008,figures7,39,46–61).Themesalfaceofthegonoplac,whichisincontactwiththe
gonapophyses, is membraneous. From the sclerotised lateral base of each gonoplac arise two
conspicuousapodemes,theweaklydevelopedfaanteriorlyandthetriangularla,withawidened
internal end, posteriorly (Figures 1b, 2c).The ventral edge of the gonoplac is densely covered
withrobust,sharpsetaedirectedposteriorly;theseareshortontheproximalpartoftheedgebut
206 N.A.MatushkinaandK.-D.Klass
Figure 4. ScanningelectronmicrographsofsensillaongonapophysesofPhenesraptor:(a)lateralviewofgonapophysis
ofS9;(b)lateralviewofgonapophysisofS8;(c)–(e),sensillaofgonapophysisofS8;(f)–(h)sensillaofgonapophysisof
S9.d:distaldirection.
longeronitsdistalpart(Figure3d,e).Solitarycampaniformsensillaarescatteredoverthelateral
(external)gonoplacwalldorsaltothesetae.
The membraneous mesal walls of the gonoplacs continue, towards the midline, into a wide
membranelyingabovethegonapophyses.Faranteriorlyinthismembraneliesamedianinfolding
(apodemeca)thatisweaklysclerotised(scleriteCA)andbearsapairoflobe-likemembraneous
tendons(ta)(Figures1b,2c).
Afirmtransversesclerite,thepoststernumofS9,PS9,islocatedbetweentheposteriorgono-
placbasesandisdividedintotwohalvesbyamedianstripofmembrane(Figures1b,2c).The
anteroventralmarginofscleritePS9isbilobate,anditsposterodorsalmarginlacksatransverse
fold,whichinsomeotherOdonataprovidestransversestiffeningbetweentheposteriorgonoplac
bases of the two sides. The lateral ends of the two PS9 halves are quite narrowly but almost
immovablyfusedwiththemaingonoplacscleritesCXa9.
The paired postlaterocoxa LCp9 is a ribbon-like sclerite extending lengthwise from the
anteroventral corner of tergite TG9 to the margin of sclerite CXa9 at the lateral base of the
gonoplac (Figure 1b); an internal ridge runs along it (anterior part seen in Figure 2a). Sclerite
LCp9reachesfaranteriorlyalongsideTG9,totheareawhereTG9isalsoincontactwithextension
jofscleriteLCa9(Figure2a).WhereLCp9andTG9meettheyarefusedandalmostimmovable
relativetoeachother.However,thereisnodirectcontactbetweenLCp9andLCa9.Theposterior
endofscleriteLCp9ismovablyarticulateduponasmallnotchinthemarginofCXa9–quitefar
anteriortothebaseofapodemela(Figure2a,c).
Thestylussl9isveryshort,essentiallycylindrical,butwithanobliquelytruncatedtip(Figure3b,
d).ItssclerotisationSL9ishardlymovablerelativetotheCX9sclerotisationssurroundingitsbase.
Theventralsurfaceofsl9iscoveredwithrobust,apicallydirectedsetaeofvariouslengths,and
theapexofsl9bearsatuftofverylongsetae.Campaniformsensillaarescatteredovertheventral
surfaceofthestylus(moredenselynearthebase).Theclusterfcsofcampaniformsensillaisnot
clearlybordered(Figure3b).
FemalegenitaliaofPhenesraptor 207
S10isentirelysclerotisedbyaringTG10+LP,butthesclerotisationoftheventralwallincluding
themedianprojectionisweaker(Figures1,2c,3f).TheventromedianprojectionofS10ispoorly
developed, represented only by a low bulge (Figure 3f). The antecosta of S10, ac10, is absent
ventraltotheshorttendonet10uponthetergalpartofTG10+LP.Theventralmembranebetween
sclerites PS9 and TG10+LP is rugose and bears neither sclerotisations nor melanisations (no
sclerite IT9, cf. Klass, 2008, figures 46–61). The dorsal tendon dt10 on the anterior margin of
scleriteTG10+LPishugeandentirelysclerotised,thoughcorrespondingtendonsarelackingin
S8andS9.TheventrolateralregionofTG10+LPbearsrobustsharpsetaedirectedposteriorlyand
campaniformsensillaoftwodifferentshapesscatteredamongthesetae.
CharactersofKlass(2008)scoredforPhenes
Characters are numbered and defined following Klass (2008). From the previously published
datasetcharacterswereselectedthatarerelatedtotheovipositor(12–24and38–64),andmostof
thosethatrefertothestructureofthesegmentwallofS8,S9andS10(9–11,65–68).Ifnecessary,
supplementary explanations are provided for these characters – especially if a new state has to
bedefinedforPhenes.Inaddition,somenewcharactersarehereindefined(80–86);fortheseall
definedstatesareexplained,andscoringforalltaxastudiedinKlass(2008)isgiven.AsinKlass
(2008),weareusinginstructiveabbreviationsforcharacterstates,whichforanalysishavetobe
replacedby0,1,2etc.
Char.09. Presence of extension b on posteroventral corner of tergum of S8: [sho] short. Note:
The posterolateral margin ofTG8 has a very weak but broad sclerotisation, which is
situatedposteriortotherowofdenticlesty8;thisisclearlyexpandedventrallytowards
articulationTG9-LCa9,butitdoesnotreachthearticulation.
Char.10. Presenceofpost-spiracularapodemeba8onlateralmarginoflaterocoxosternumLS8:
[dst]distinct.
Char.11. Presenceofmedianprocesspp8onposteriormarginoflaterocoxosternumLS8:[abs]
absent.
Char.12. Location of origin of tendon gt: [sct] arising from distinct sclerotisation on anterior
marginoranteromesalcornerofcoxaCX8.
Char.13. ExtensionofbaseofapodemegaofCX8towardsmidline:[lon]reachingandcontinuing
throughareaoforiginoftendongt,whichthusoriginatesfrommesalpartofapodeme
ga.
Char.14. Interrelation between coxae CX8 of the two sides: [sep] entirely separated by
membrane.
Char.15. Interrelation between coxae CX8 and antelaterocoxae LCa9: [sep] separated by
membrane,notarticulateduponeachother.
Char.16. Extension to the anterior of cleft between gonapophyses gp8 of the two sides: [ime]
anteriorendofcleftroughlyatthelevelofarticulationGP8-LCa9.
Char.17. Size of sclerite MS at ventral gp8 bases (i.e. at anterior end of cleft between
gonapophysesofS8):[abs]absent.
Char.18. InterrelationbetweenscleritesMSandGP8atventralgp8bases:inapplicable.
Char.19. InterrelationbetweenscleritesGP8ofthetwosidesatdorsalgp8bases:[sep]separated
bymembrane.
Char.20. ConditionofGP8sclerotisationatdorsalgp8bases:[mes]withdeep,broadmembra-
neousnotch.
Char.21. Presenceofobliqueridgesgyonmesalfaceofgp8:[abs]absent.
Char.22. Conditionofridgesgyonbasalmesalfaceofgp8:inapplicable.
Char.23. Presenceofsaw-teethtm8distolaterallyongp8:[abs]absent.
208 N.A.MatushkinaandK.-D.Klass
Char.24. Presenceofridgesgz8distolaterallyongp8:[abs]absent.
Char.38. Presence of spine sa on antelaterocoxa LCa9 near its articulation with sclerite GP8:
[abs]absent.
Char.39. AnteriorextensionofpostlaterocoxaLCp9:[lon]reachingarticulationbetweentergum
TG9 and antelaterocoxa LCa9 (Figure 2a). Note: LCa9 is in contact with TG9 only
intheareaofh(uponantecostaac9,viaextensionj).LCp9doesnotreachthispoint
directly,butreachestheposteriorflankofthebulgeonac9fromwhichextensionsjand
cdiverge.
Char.40. Interrelation between tergum TG9 and anterior part of postlaterocoxa LCp9: [cnt]
connectedbysclerotisationthatisasheavyasthatofthetwoscleritesinthisarea.
Char.41. Interrelation between ante- and postlaterocoxa, LCa9 and LCp9: [sep] separated by
membrane,two-piece“gonangulum”.Note:Thecriteriaforconnection(=one-piece
gonangulum)orseparationofscleritesLCa9andLCp9arediscussedinKlass(2008,
pp.118–120);onlyaconnectionofLCa9andLCp9mesaltostriph(Figure2a)rep-
resentsatrueconnectionbetweenthesetwoscleritesthatisnotmediatedbypartsof
tergiteTG9.
Char.42. Interrelation between gonapophyses gp9 of the two sides: [sep] free from each other
downtotheverybase.
Char.43. Extensionofrowofsaw-teethtm9alongdorsaledgeofgonapophysisgp9:inapplicable
(seebelowChar.81).
Char.44. LocationongonapophysealscleriteGP9ofextensiongbearinganteriorendofrhachis
rh:[pos]distinctlyfartherposteriorlythananteriortipofGP9.
Char.45. Presence of tendon ft flanking apodeme ca: [abs] absent. The tendons ta that in
Phenes are placed on apodeme ca (see Char. 82) are not homologous with the ca-
flankingtendonsftofZygoptera,sincetheyprobablyprovideattachmenttodifferent
muscles.
Char.46. Presenceofposteriad-directedextensioniatbaseofscleriteGP9:[abs]absent.Note:
extensionsofaronlyfoundinEpiophlebia(Klass,2008,iinfigure41).
Char.47. Locationofopening(s)ofaccessoryglandsagofthetwosides:Newstate[mid]right
andleftductsofaccessoryglandsuniteintoanunpairedcommonductshortlyinternal
totheorifice(Figure1b),whichisthuslocatedatmidline(atthesameanteroposterior
levelasthepairedopeningsinotherovipositor-bearingOdonata).
Char.48. LocationofarticulationbetweenscleritesCXa9andLCp9:[ant]distinctlyanteriorto
lateralbaseofapodemela.
Char.49. PresenceofdorsalgonoplacscleriteCXb9:Newstate[ews]embeddedinweakscle-
rotisation.Note:intheentiredistalpartofthegonoplac–posterodorsaltothestylus
base–thecuticleisuniformlyweaklysclerotised(strongerthan“typical”membrane
but weaker than “normal” sclerites) (Figure 2c); this sclerotisation is confluent with
scleriteCXa9,butpartofitrepresents(byposition)aweaklysclerotisedCXb9.
Char.50. Interrelation between distal parts of sclerites CXa9 and CXb9: inapplicable due to
lackingdifferentiationofscleriteCXb9(seeChar.49).
Char.51. PresenceofscleriteCXc9besidebaseofstylussl9:inapplicable.Note:scleritesCXc9
are, when present, tiny in other Odonata. Since in Phenes the sclerotisations of the
gonoplac and of the stylus (CXa9 and SL9) are largely confluent, it is impossible to
scorethepresenceorabsenceofaCXc9.
Char.52. Shapeofstylussl9:[cyl]straight,cylindrical,tipnotwidened.
Char.53. Presenceoftubercles/projectionsondistalventraledgeofgonoplacgl9:[abs]absent.
Char.54. Conditionoftubercles/projectionsondistalventraledgeofgonoplacgl9:inapplicable.
Char.55. Presenceofsetaltuftontipofstylussl9:[dst]distinct.
Char.56. PresenceofscleritePS9:[dst]distinct.