Fossil Zygoptera and Anisoptera from Upper Miocene of Monte Castellaro (Pesaro, Marches, Central Italy). (Insecta Odonata)

2004·2.4 MB·

by  G. Gentilini

Quoderno di Studi e Nofizie di Storio Nqturole dellq Romogno
Qttutl.S ndi Nat.R onrctgnu1,9 : 17-41.g iugno2 004 lssN 1123-6787
GiuseppGee ntilin&i LorisB agli
FOSSIL ZYGOPTERA AND ANISOPTERA FROM THE
UPPER MIOCENE OF MONTE CASTELLARO
(PESAROOM ARCHES' CENTRAL ITALY)
(I nsectOa donatCa oenagrionidLaees, tidaeS,i eblosiidae,
CalopterygidLaieb,e lluild ae)
Riassun to
lZigoilari c Ani.softari. /os.rili dal Miocane.sultariora di MonÍe Cu.slallurrt, Pa.strrrt,M ur-
gha, Itttliu c'qttt'ulc (ln,sat'lu Oclonutu Coutugrittnicluc. Le.;Íidue, Siehlo'siitlac,
(' u I oy scl t '.t'gi t l u a, L i h aI I u I i d u e) I
Vengono cJiscr-rsese l ìgr-rrates ctte ali fbssili di Odonata Zygoptera e descritte due nuove
spccicc li OclonataA nisopteraa ppartencntai lla lami-uliaL ibellLrlidaes, coperten el giacirnen-
t6 rtriocenicod i Montc Castellaro.P esaro.Vicncit ioltrc esatttinatau n'ala postcriored i libel-
lirla attribLritaa lla sottofirmiglia Pantalirliìec al -uencreT rupa:tt.stigmup, Íovenientc dalla
stessal ocalitrìf i rssilil-cra.
Abstract
Scvcn fitssil wings of OdonataZ ygopteraf t'onrt he Upper Miocene of Montc Castellaro,
Pcsalt, are cliscr-rsseacnld lìgured, and two nerv speciest t1'OdonataA rlisopterab elonging
to thc lhnrily LibcllLrlidaca, re dcscribed.A dragonfìy hindu,ing attribr.rtedto thc sr-rbfarnily
Pantalinaea nd genus Tt'upe:ostigrraf ior.nt he sarnef ìlssililèrousl ocality, is also exarrined.
Key wgrds: Insecta,O donata,C oenagrionidaeL. estidae,S ieblosiidaeC, ìalopterygidae,
LibellLrliclacT,a xotromy,U pper Miocene, Motrte Castellaro.P esaro,M arches,C entral ltaly.
lntroduction
New fbssild an-rself'liaensd d ragonfìiefsì 'or-trh-re U pperM ioceneo f MonteC astellaro,
haveb eene xallined.T he Odonataf àunac onsistso f separatew ing relîains,
preserveda s colllpl'essionins fìne-grainedb,i turninottsm alls, exposedi n twcl
f-ossilifèrosuesc tionds evelopeadl ongt heA driaticc oasti,n thel owerp arto f a cliff
about3 kpr north-wesot f Pesaro( MarchesC, entrall taly) and includedi n the
Gessoso-solfìfèFrao nnationo f Lower Messiniana ge.M ost of the dragonflies
"strato
haveb eend iscoverebdy thes eniora uthori n the deglii nsetti"( Gt'NltltNt,
I 9tì9.1 992"I 993:B ,qc;L&r Gr,NrrL-r2N0r0, 3) , a well celîerìtedin secbt eda bout1 40
17
mm thick, locatedn eart he top of the Formationa nd consistingo f very thin, dark-
greya ndl ight-greye vaporiticl aminaep robablyr elatedt o a seasonadl epositionI.n
this layer severali nsecto rdersh ave beend iscoveredo, ften associatedw ith fish
and leaf remains.T hesef ossil assemblagems ay indicateb oth deepo ff-shorea nd
shallow,n ear-shorep aleoenvironmentst:h e presenceo f varved shalesn ormally
occurringi n quiet basins,t he predominanceo f articulatedf ish remainsa nd com-
plete insectsp reserveda s adults( especiallyH emiptera-HomopteraD,i ptera
BibionidaeM, ycetophilidaeL, imoniidaea nd insectsw ith strongf lying ability as
Diptera Syrphidae,H ymenopteraa nd Odonata),m ay indicatea relatively deep-
water/ofÈshorep aleoenvironmentw, hile abundantr emainso f evergreenn eedles
and taxodiaceousle avesa nd the presenceo f insectsw ith weak flying ability
(Coleoptera,T richoptera,t errestrialH emiptera),m ay representa near-shore
paleoenvironmen(Wt rsoN, 1980).M oreover,t he scarcityo f marinet axaa mong
"strato
thei chthyofaunao f the deglii nsetti"(o nly few specimenos f Latess p.a nd
Harengula sp. have been found) and the predominanceo f the hypereuryhaline
speciesA phaniusc rassicaudus(A gassiz,l832),i ndicateh ighly salinew ater
conditions( SonsrNr,l988w),i th a normals alinitym arinee nvironmentp, resenta t
leasti n cerlainp eriods.Them ajorityo f the terrestriapl lanta ssemblagecso llected
in this insectb ed, suggestv egetationabl elts characterizedbya greatd iversity of
habitatsw ith the presenceo f Thxodiumin the warm, humid and swampyc oastal
plains,a belt of thermophylep lantsw ith prevailing Quercus,A cer, Populus and
probablem ountains lopesw ith predominancoef xerophilef orestso f Pinus,S equoia
and Thuja. However,t he occurrenceo f DipteraT richoceridaein the insect
assemblageasn dt heh igh proportiono f Nearctica ndP alearcticg eneraa mongt he
Odonatafa unar, nights uggesat lsoc limaticf luctuationisn cludinga possiblec ooling
phasec, orrelatedw ith the salinityc risisi n the MediterraneaSn ead uringt he Late
Miocene,w ith a vegetationabl elt of conifersl oweredt owardst he depositional
basin( BenrolaNrM nncHErr&r MnnrorrrL rrer,1 989);a nyway,f urtherc ollecting
and comparatives tudy will increaseo ur knowledgeo f this Lower Messinian
paleoenvironment.
F-ossilAnisopte(raad ultsa ndl arvae)f rom MonteC astellaroa rer elativelyc ommon,
while fossil Zygopteraaree xtremelyr are:o nly eights pecimens( about3 .5oAo f
the total numbero f Odonata)h aveb eenf ound up to now. It is interestingt o note
that speciesb elongingt o the subfamilyI schnurinae( OdonataC oenagrionidae)
and the family Calopterygidaea rei nfrequenti n Tertiaryi nsectl ocalitiesa nd only
one recordo f the genusI schnura is known from the Eocene/Miocenea mber
inclusionso f the DominicanR epublic;o n the contraryt,h e familiesL estidaea nd
Sieblosiidaea rew ell known from Cenozoicd epositsM. oreover,t he finding of the
generaD eielia and Trapezostigm(aO donataL ibellulidae)i s also of considerable
interest:t he wing of Deielia discovereda t Monte Castellaro,r epresentsth e first
fossil recordo f this genus,w hile well preserveds peciesb elongingt o the genus
Trapezostigmaa re poorly known from EuropeanN eogened eposits.
Fortunately,a Nature Reserve( ParcoN aturaled i Monte San Bartolo, including
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the Monte Castellaroo utcrop,c onsidereda s insectsd eposito f exceptional
preservation)w, as establishedin 1994a nd now the protecteda rea- uy b. used
for researche, ducationa nd orgamzedv isits. The most extensivef ossil insect
collectionsf rom the Monte Castellaros ite, includedt he specimensd escribedi n
this paper,a ted epositedin the Museod el Territorioo f Riócione( Rimini, Emilia
Romagna,N orthernl taly).
The venationa bbreviationsa re basedo n Rrpr & KuralovA-pEcr (19g4) with
modificationsb y NEr-e t al. (1993)a nd BEcHlv( 1996).Thec lassificationo f fossil
ande xtantO donataf,o llows BEcHI-(v1 996,2000)andti e principleso fphylogenetic
systematic(ss ensuH nNNrc1 966,19 69).
Systematicp alaeontology
Order Odonata Fabricius,lT93
Suborder Zygoptera Selys1, 854
Superfamily Coenagrionoidea Kirby,1890
Family Coenagrionidae Kirby,1 890
Subfarnily Ischnurinae Fraser,1957
Genus Ischnura Charpentier1,8 40
Ischnura sp. (SpeciesA - Fig. i;
Material- Specimenn o.1289a nd 1289a( part and counterpart)c; oll. Gentilini;
Museod elTenitorioR, iccione(R imini,E miliaR omagnaN, orthernl taly);G essoso-
solfiferaF ormation,M arne bituminoseu nit, "Stratód egli insetti,,l lvel; Upper
Miocene( LowerM essiniano) f MonteC astellaro(P esaroM, archesC, entrall tàiy)
Fig. I - Venationo f Ischnurtt sp. (SpeciesA ): right forewing of male. Scale:3 mm.
t9
Description
The spèciesis known frorn an isolatedr ight forewingo f male with part of the
posteiiorw ing marginr nissing.Wingm embraneh yaline;v enationn ot densew ith
ielativelyl argec el1sv; einsb rown'b asen arrowa ndw ell petiolatew; ing apexw ide
andr oundedl;o ngitudinavl einsl ong ands traightW. ing length,1 5.2r nn"rw; idth at
nodus,2 .4 nm; width at wing base,0 .5 rnrn,p etiolea bout2 mm long; distance
fiorn baset o nodus,5 mm; distancefi om baset o arculus2, .3 rnm; distancefi om
arculusto nodus,2 .2mm; distancefi om nodust o pterostiglna3,. 6r nrn,d istance
fiorn pterostigrnato apex, 1.8 mm; noduss ituateda t about3 2.8 % of the wing
lengtlt.P rirnarya ntenodacl rossveinAs xl andA x2 aligneda ndb racket-likeA, x2
in linew ith thea rculusa nds ituateda t 1. 1 mrnd istalo f Ax I ; absencoef secondary
antenodaaln da ntesubnodcarlo ssveinsA.r culusk inked;s ectorso f arculusd istinctly
separatefdio rnt heiro rigin;n odalc rossveinin linew ith subnodusn;i nep ostnodal
crossveinsb etweenc ostalr nargina nd RA all alignedw ith the postsubnodal
crossveinbse lowt heme xcludingt he mostd istali ncotnpleteP. terostigrnsah ort
with prargintsh ickenedb,i coloure(db rownt o whitey ellowish)a, sl onga sw idea nd
coveringónloy nec ell;pterostigrnbarla cev eino bliquea ndi n linew itht hep roximal
sideo f thep terostigl.ntah;e rea ret hrees inglec ellsb eyondt hep terostigrnbae tween
thea nteriowr ing margina ndv ein RA. Discoidacl ellt rapezoidainl shapec, losed
basallya 1dw itSd istala nglea cute:le ngtho f basasl ide,0 .3r nrn;l engtho f ventral
side,0 .9 rlpr; lengtho f dorsals ide,0 .5 rr-ur;le ngtho f distals ide,0 .5 urtn.V ein
MAb strolglyo bliques; ubdiscoidcael llr elativelyw ide( length,1 .3n -rmw; idth,
0.25 rnm),w ith distals idea ngr-rlatecde;l l sitr,ratebde low the subdiscoidacle ll,
unusuallyc rossedB. asaal reaf iee;c ubitacl ellw ith onlyC r"rP-crossisnitgu ated2 .3
pm distalo f'thew ingb aseb; asapl ortiono f CuA,s traightP. ostdiscoidfarle ldw ith
oner ow of cells;v ein MA andM P basallys traighwt ith oner ow of cellsb etween
therr-vre, inC uA straighat tb aseF. irstb ranchingo f RP,0.2r nrnb asaol f thes ubnodus;
"O",
baseo f IR2 alignedw ith subnodusle, stineo bliqr-rvee in absenta; reab etween
IR2 apclR p3-4,v ery narrowb asallyF. orko f RPI at aboutf burcellsa r-r4d. 3 rnrrr
distalo f the subnodusv;e in RPI kinkedb elowt he pterostign-rbarla cev cin and
slightly zigzaggitrgV. ein IR I separatefsio r-rR"rP I , 2.5 tlun distalo f the fbrk of RPI
andt wo cellsb asalo f thep terostigmav;e in RP2s eparatefsio m RPI threec ells
basalo f IRI andf -ourc ellsb eyondt he nodus.O ner ow of cellsd istallyb etween
RA arrdR Pl. betweenR PI and IRI ( only a dor-rblcee ll is presenot n the wing
rnargin)a, nd IRI andR P2.D istalp ortiono f IR2 ztgzaggedd;o ublec ellsb etween
vein.Rs P2a ndI R2.a ndI R2a ndR P3-4o nt heh indr nargint;h ev einR P3-4r eachcs
thep osteriowr ing borderw ell befbret hep roximals ideo f thep terostigma.
Discussion
The fbssild arnselflyw ing describeda bovec an be attributedto the extantg enus
Isc,hnurbue causeo f the followingv enationacl haracterss:r nalls izeo f thew ing,
distipctlyp etiolatev; enationr athero pen;w ing apexf airly roundedin f-orewing;
analv ei n separatinfgio rnt hep osteriowr ing borderb ef-orteh eC uP-crossinga;r culus
20
situateda t thel evelo f thed istala ntenodald; iscoidacl ellb asallyc losedw ith distal
anglea cute;a reab etweent he baseso f IR2 and RP3-4,n arrowed;p terostigma
bicolouredi n male specieso f the genus,e speciallyo n the fbrewing;f ork of RP
generallys ituateda boutf bur cellsd istalo f then odusi n forewingsa ndt hreec ellsi n
hindwingsM. oreovert,h isf ossils peciess howsa n unusuafle aturet:h ec ell situated
belowt hes ubdiscoidacle lli s crossedn earlyi n them iddle;t hisc haracteirs usually
not presenitn Coenagrionoideaan dp robablyr nayb e considereda s an aberration
rathert han a diagnosticf -eature.Thgee nusE nullugtl'ter esernblesIs c:hnurai n
lnanyc haractersb.u t it differsi n havingf ìve-eighct ellsi nsteado fthree-fourb, etween
subnodusa nd fbrk of RPI; moreovert,h e wing apexo f Enallagntuis slightly
pointeda nd more der-rselrye ticulatedT. he f-ossilI sc'hnttra( SpeciesA ) shares
synaponlorphiewsit h thef ollowingt axai ntroducedb y Bnc'rtv( 1996)E: uzygoptera
(longitudinavle inss traighat ndl ong;o ner ow of cellsb etweenC uA andp osterior
wing rnargino; nly primarya ntenodalAsx l andA x2 presenbt asalo f the nodus;
antesubnodaalr eaw itl-rouct rossveinsa s well as areab etweenR P and MA, from
arculusto rnidfork)C, oenagrionomorp(hpao stnodaaln dp ostsubnodcarl ossveins
"O"
in alignrnenwt,i tht her nosdt istasl ometimeisn cornpleteo;b liquev ein secondarily
abser-drti;s coidacl ellc loseda tb asein fbrewings)C; oenagrionifbrm(ipat erostigma
rathers hortw ith two crossveinbse lowi t; subnodusir -lri ne with theb aseo f IR2;
nurnbeor f exagonaal ndp entagonacle llss tronglyr educeda; reab etweenR PI and
RP2w ith two rowso f cellss eparatebdy the vein IRI;postnodacl rossveinisn
aligr-rmewnti th crossveinssi tuatedb elow;i ntercalarvye insl acking,e xceptI R1
and IR2); Coenagrionid(a RPI kinkedb elow the pterostigmabl racev ein);
Cenagrionode(a p roxirnal,ventraaln dd istalr narginso f pterostigmath, ickeneda nd
fonninga U-shapeds tructure)C; oenagriorrida(Kei rby, 1890),( vein MAb very
obliques ot hatt hed orsals ideo f thed iscoidacl elli s shortetrh ant hev entrasl ide).
Thef bssilr ecordf or theg enusIs c'hnurius p oor:o nlyo nes peciecse rtainlyb elonging
to this genus,h as beend escribedb y Br,c'rrlv(2 000a)f iorn Dominicana mber
inclusion(sE oceneto Miocene)w, hile specieso f Coenagrionidaoer Ischnurinae
incertaes edish, aveb eenr eportedb y Nr,r&- Pnpnzrn(N 1 990)a ndb y Nr,L,M nnríNr,z-
Dt'lc'LosP, nt'll,&tt Oult.ttto( 1 997)f,i ornt heU pperO ligoceneo fAix-en-Provence
and the Upper Mioceneo f Sainte-ReineF,r ance.I sc:hnura(S peciesA ) differs
fiom tlreD orninicana mbers pecieIss c'hnurave lÍeni(Bechly2,0 00),i n thel onger
andw iderw ing,t hel ongerp etiolea ndt hec ellb elowt hes ubquadranglcer,o ssed.
Moreoveri,t is distinguishablfeio rn theF renchs peciesb y thes horlera ndn arrower
wing ( thew ing of thes pecirnenno .lPM-R0710f6r omA ix- en ProvenceF, rance,
hast hes amel ength( 15.2r nm)b ut it is wider),b y thes hapeo f thep terostigrnnao t
rhornboidalt,h e crossedc ell below the subquadrangalen d the highero r lower
numbero f postsubnodcarl ossveins.
At tlrep resendt ay the genusI sc'hnttruis a cosmopolitanw, idespreadg enusa nd it
is representeidn continentaEl uropea nd westernM editerraneabna sinb y seven
species.
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Ischnurinae (SpeciesB - Fig. 2)
(Genusa nd speciesu ndetermined)
Material- Specimenn o. 1288a nd 1288a( parta nd counterpart)c; oll. Gentilini;
Museod elT enitorio,R iccione( Rimini,E milia RomagnaN, ortheml taly); Gessoso-
"Strato
solfiferaF ormation,M arne bituminoseu nit, degli insetti" level; Upper
Miocene (Lower Messinian)o f Monte Castellaro( Pesaro,M arches,C entral
Italy).
Description
The speciesis representebdy an isolatedl eft hindwing with a small portion of the
posteriorwingm argin,m issing.Wingh yalinew ith basen affowa ndd istinctlyp etiolate;
veinsb rown,w ing length,l 4 mm; width at nodus,2 mm; width at wing base,0 .4
mm; petiole,2 mm long;d istancefr om baset o nodus,5 .1m m; distancefr om base
to arculus,2.7m m, distancefr om arculust o nodus,2 .5 mm; distancefr om nodus
to pterostigma7, . 3 mm;distancefr om pterostigmato apex,1 .8m m; noduss ituated
at about 36.4% of the wing length.P rimarya ntenodalsA xl andA x2 aligneda nd
bracket-like;Ax2in linew ith thea rculusa nds ituatedI mm distalo fAxl, absence
of secondarya ntenodaal nd antesubnodaclr ossveinsA. rculus kinked and sectors
of arculuss eparatedfr om their origin; sevenp ostnodacl rossveinsin line with the
correspondingp ostsubnodacl rossveinse, xcludingt he most distal,i ncomplete.
Pterostigm(al ength,0 .5 mm; width, 0.3 mm) rhomboidailn shapew ith proximal
andv entrals idest hickened:s omet raceso f paleb rown colourp atterna rep reserved
on them embraneT. he pterostigmais well braced( itso bliqueb asals idei s aligned
with the brace)a ndc oversa boutt hreeq uarterso f a cell; therea ref our singlec ells
with the first well elongate,b eyond the pterostigmab etweent he anteriorw ing
margina nd vein RA. Discoidalc ell closedb asallya nd with distal anglea cute:
lengtho f basals ide,0.25m m; lengtho f ventrals ide,0 .8m m; lengtho f dorsals ide,
0.4m m; lengtho f distals ide. 0.4m m. Subdiscoidacle ll moderatelye longatew ith
distals idea ngulatedc; ell situatedb elowt hes ubdiscoidacle ll freeo f crossveins.Vein
MAb obliquea nda lignedw ith subdiscoidavle inlet;b asals pacef ree;c ubitalc ell
2
Fig. 2 - Venationo f Ischnurinae(S peciesB ): lefi hindwing' Scale:5 mm.
22
with CuP-crossings tronglyo bliquea nds ituateda t about2 .2 mmdistalo f thew ing
base;a nalv ein separatingfr om the wing margin,0 .25m m basalo f CuP-crossing.
Discoidalf ield naffow with only oner ow of cells;v ein MA andC uA straighti n the
first portiona nd zigzaggedd istally;o ner ow of cellsb asallyb etweenv einsM P and
CuA'''a''n d betweenC uA and posteriorw ing margin.F irst branchingo f RP at 0.2
mm basalo fthe subnodus;basoef IR2 in linew ith subnodusa ndb asala reab etween
IR2 and RP3-4,v ery narrow.V ein RP2 separatesfr om RPI three cells basalo f
IRI and two and a half cells beyondt he nodus.VeinR P1 kinked below the
pterostigmablr acev ein;o rigino f IRI, 3 mm distalo f theo rigino f RP2a ndo nec ell
basalo f the pterostigmat;h erea re doublec ellsb etweenR A and RPl, Rpl and
IRl,IRI andR P2,o n thew ing apex.L estineo bliquev ein "O" absentv; ein IR2
stronglyz igzaggeddistallwy ith doublec ellso n theh ind margin;v ein RP3-4s lightly
sinuousa nd reachingt he posteriorw ing bordero ppositet he distal side of the
pterostigma.
Discussion
The speciess howsv enationafle atureso f thee xtants ubfamilyI schnurinae(w ings
petiolatet o level of CuP-crossinga nd anal vein separatingfr om posteriorw ing
borderc loset o CuP-crossinga;r culuss ituateda t levelo fAx2; discoidacl ell closed
basalfy andw ith distala nglea cute;a reab etweenth eb aseso f IR2 and RP3-4 very
narrow),b ut the reliablea ttributiont o a genusi s difficult for the lack of body
charactersT. his specimeno f Ischnurinae(S peciesB ) differs from Ischnura
(SpeciesA ) describeda bove,i n the shortera nd narrowerw ing, the apexs maller
and pointeda nd the pterostigmas carcelyc oloureda nd rhomboidali n shape.
Moreover,t he cell situatedb elow the subquadranglies free of crossveinsa nd
therea res evenp ostnodalisn steado f nineb etweenth ec ostalm argina ndv ein RA;
thef ork of RP is at aboutt wo anda half cellsd istalo f thes ubnodusin steado f four
cellsa nd the origin of IRI is only one cell basalo f the pterostigmabl racev ein,
insteado f two cells.T he Ischnurinae(S peciesB ) differsf rom the ambers pecies
Ischnurav elteni( Bechly,2 000), in the longer and wider wing and the longer
petiolea ndw iderd iscoidacl ell;m oreoveri,t is quited istinctf rom Coenagrionidae
or Ischnurinaien certaes edisf rom Franced, iscusseidn thed escriptiono f Ischnura
(SpecieAs ), in the shortera ndn arrowerw ing, in the lessn umerousp ostnodaal nd
postsubnodaclr ossveinsa ndi n havingt wo anda half cellsi n theb asala reab etween
RPI andI R2, insteado f four or five cells.A ccordingt o NEr-& PnrcHElE(n1 993a),
fossilC oenagrionoideaar ed ifficult to recognizeo nly by wing remainsa nds everal
determni ationsa reu ncertain.
Family Lestidae Calvert,1 901
Subfamily Lestinae Calvert,l9 0l
Genus LesÍes Leach,1 815
ZJ
Lestes sp. (SPeciesC - Fig. 3)
Material- Specimenn o. lzgi and l2gia (part and counterparl)c; oll. Gentilini;
Museod elr érritorio,R iccione( Rimini,E miliaR omagnaN, orthernI taly);G essoso-
solfrferaF -ormationM, arne bituminoseu nit, "strato degli insetti" level; LJpper
Miocene( LowerM essiniano)f MonteC astellar(oP esaroM, archesC, entraIl taly)'
Description
The spéciesis representebdy a single,h yalinel eft wing with the distalh alf of the
membranes tronglyf olded io*ardi the costalm argin.Thew ing venationo f the
specirnenh asb eJnr econstructebdy parta ndc ounterparl.Veinbsr own;w ing length,
l7 rnrn;w idth at nodus,I .2 mm; width at base0, .5m rn;p etiolea boutI '9 mm long;
distancefi om baset o nodus,5 mrn;d istancefr om baset o arculus2, -4m m;d istance
fiorn baset o nidf-ork,2 .9 mrn; distancefr orna rculust o nodus,2 '5 mm; distance
fiorn' odus to pterostigma8,. 5m rn;d istancefr om nodust o apex,l 2'3 rnrn;d istance
fior' nodusto origino ÍRp2.2.6r nm;d istancfeio m nodusto origino f IR2' l.tì mrn,
oA
distancefi orn noAu,t o origin of Rp3-4, 1.9m rn; noduss ituateda t about2 9 of
thew ing lengthp. rimarya ntenodaAlsx I andA x2 aligneda ndb racket-like;Axi2n
line with the arculusa nd 1.1 rnrn distalo f Ax1. No secondarya ntenodaal nd
antesubnodaclr ossveinsb;a sala reab etweenR PI and IR2, lree of crossveins'
Sectorso f arculgss eparateda t the base;n ine postnodacl rossveinsm ore or less
alignedw ith the postsubnodaclr ossveinssi tuatedb etweenR A and RPI' except
fbr the lastt wo incompleteP. terostigmcao louredp aley ellowish,e longatea nd
bracedw. ith theo bliqueb asals idea lignedw ith theb race.Thpet erostigmias 1.4
mm long and 0.35 mrn wide; therea re two crossveinsp reservedb elow the
pterostigrnaan d six singlec ellsb eyondi t, betweent he anteriorw ing margina nd
veinR A.Discoidacl elln arrowe, longateb,a sallyc losedw ith distala ngles trongly
acute:le ngtho f basals ide,0.2r nm;l engtho f ventrals ide,I mm; lengtho f dorsal
side,0 .6r nn; lengtho f distals ide,0.4m rn.B asasl pacefi ee;c ubitacl ellwithC uP-
crossings ituateda t 1.7r nrnd istalo f thew ing basea; nalv eins eparatinfgio m ving
rnargina tt hel evelo f Cup-crossingo.n ly oner ow of cellsi n thec ubito-anaalr ea;
vein CuA srronglyz igzaggeddi stalìya ndv ein MP straightw ith distale ndd istinctly
bentt owardst hei lna màigin.B asala ndd istalp ortiono f vein MA zigzaggedo; ne
row of cellsb etweenM A and MP; vein RP3-4s traighta ndr eachingth e posterior
wing marginw ell beyondt he distals ideo f the pterostigrnaa;r eab etweenR P3-4
andM A widenedd istàlly;a longa nds traighitn tercalarylo ngitudinavl eins ubtending
two rows of cells,p resénbt etweenv einsR P3-4a ndM A. Spaceb etweenb aseso f
IR2 and Rp3-4 very narrow:f irst branchingo f RP situated1 .8m m basalo f the
subnodusd; istancete tween origino f IR2 ands ubnodusI, '7 mm; nodalc rossvein
"O"
ands ubnodusw ell obliquei n porition.L estineo bliquev ein betweenR P2a nd
IR2, situated4 .5 mm distalo f the subnodusV. ein RP2o riginatesfr om RPl,2.4
mrnd istalo f thes ubnodusc;o urseo f vein IRI strong ly zigzaggebda sally;m osto f
distala reab etweenR p1 andI R2 missing.o ne row of cellsb etweenv einsR A and
Rpl andt wo-threer ows of cellsb etweenl R2 and RP3-4t owardst he wing apex'
/-+
Discussion
This fossill estidc anb e attributedto thee xtantg enusL estesb ecauseo f thef ollowing
venationafl eaturesw: ing hyalinea nd well petiolateds, ectorso f arculusa rising
neart heu pperp arto f thea rculusd; iscoidacl ell with distala nglew ell acute,o rigin
of RP3-4n earert o arculust hant o subnodusp; terostigmae longater,e ctangulairn
"O"
shapea nd as long as the two cells underi t; obliquev ein situatedb etween
veinsR p2 and IR2; analv ein separatinfgio m posteriowr ing rnargina t levelo f
CuP-crossinga;r eab etweenR P3-4 andM A widenedd istally,w ith secondary
longitudinavle insp resentT. hes peciessh aress ynapornorphiwesit h thef bllowing
taxài ntroducedb y Br,c'rn-(v1 996):E ,uzygopte(rlao ngitudinavl einss traighta nd
long;e ndo f vein RP3-4s ituatedb eyondt he pterostignav;e in MA moreo r less
1..uò1,inthge p terostigrnav;e in CuA endingd istalo f ther nidwing;o ner ow of cells
betweenC uA and the posteriorw ing margin;o nly Axl andA x2 presentin the
antesubnodaalr ea);E ulestifonr-r(ivae in MP curveda fteri ts origin on the distal
angleo f t5ed iscoidacl ell);L estode(aa rcr.rlussh iftedto wardsth ew ing baseb, eneath
the-prirnaAryx 2); Lestinoide(aC alveftI, 9 01) , (rnidfbrkr ecessebda sallya nds ituated
ata bour2 0-26%oft hew ing length;v ein MA zigzaggeddi stally);L estidae( dorsal
arcularb rackerte ducedd; istald iscoidavl einM Ab veryo bliquea ndd iscoidacl ell
with distala nglew ell acr-rteth; reer ows of cellsb etweenI R2 and RP3-4o n the
hindm argin)L; estinae(l owerp ortiono f thea rculusf- onnebdy theb asasl ideo f the
discoidacl ell,s ot hati t is veryc loset o vein RA). Severasl peciebs elongingto the
genusL csÍe,hsa veb eend iscoveredin E,oceneO,l igoceneM, iocenea nd Pliocene
aepositso f E,uropet:h e speciesfì 'ornt he Monte Castellaros ite,r nayb e placedi n
tlref -ossiLl estagsr oupB (Nr,r&- Pnrc'rrr,Lr1,9n9. 4)a, grollpc haracterizebdy lxore
tnodernv enationacl haractersb,e causeo f tl"rea bsenceo f a sr-rppletnentarroyw ot
cellsb etweenv eil MP andt hep osteriowr ing margin.I n the Palaeogensep ecies
of Lcstc.gs roupA , fbundi n westernE ,uropelo calitiest,h ree-fburr owso f cellsa re
presenbt etweenv ein MP andt heh indr narginC. oncerninglt alianf bssill estidsa'
ibre*i1rga nd a hindwingb elongingt o the subfarnilyS yrnpecrnatinaaen d genus
(Selys,1 840),h aveb eenf ìguredb ut not describedb y Cìnvnllo&
$,rrlrocirrr
Gnr.ì-,,'r(t1' 987),f rom the UpperM iocene( Lower Messiniano) f the Scaparoni
andP iobessi itesn, earA lba (PiedmontN, orthernl taly).
Reconstrgclign of'lefi *i1,s gl' Le.sle.ss p. (Specics Cì) by part irtld coutrtcrpart. Scale: 3 tl'tt't.t'
25
At the presentd ay the genusL estesi s cosmopolitea nd occursi n vegetateda reas
with still, sluggisho r moving water and in cool temperateto tropicalc limate.S ix
specieso f Lestesa re representedin Europe.
Lestidue( SpeciesD - Fig. a)
(Genusa nd speciesu ndetermined)
Material- Specimenn o. 1290a nd 1290a( parta nd counterpart)c; oll. Gentilini;
Museod elTenitorioR, iccione( Rimini,E miliaR omagnaN, ortheml taly);G essoso-
"Strato
solfifera Formation,M arne bituminoseu nit, degli insetti" level; Upper
Miocene( LowerM essiniano) f MonteC astellaro( PesaroM, archesC, entralI taly).
Description
The speciesis known from head,t horaxa nd abdomenf ragmentsa nd from the basal
areaso f fore and hindwings.H ead in poor condition; thorax dark brown to black
colouredm: esepisternumab out4.3m m long and I mm wide; mesepimero3n. 5 mm
long and about0 .7 mm wide; preservedle ngtho f dark brown abdomen,1 3.Jm m;
4
Fig. 4 - Vcnationo f fore and hindwing baseso f Lestidae( SpeciesD ). Scale:2 mm
26