Quoderno di Studi e Nofizie di Storio Nqturole dellq Romogno Qttutl.Sndi Nat.Ronrctgnu, 19: 17-41.giugno 2004 lssN 1123-6787
GiuseppeGentilini & LorisBagli
FOSSIL ZYGOPTERA AND ANISOPTERA FROM THE UPPER MIOCENE OF MONTE CASTELLARO (PESAROOMARCHES' CENTRAL ITALY) (Insecta Odonata Coenagrionidae, Lestidae, Sieblosiidae, Calopterygidae,Libellul idae)
Riassunto Mur- lZigoilari c Ani.softari./os.rili dal Miocane.sultariora di MonÍe Cu.slallurrt, Pa.strrrt, gha, Itttliu c'qttt'ulc (ln,sat'lu Oclonutu Coutugrittnicluc. Le.;Íidue, Siehlo'siitlac, (' u I oysl c t'.t'g i tl u a, L i h aI I u I i d u e) I Vengono cJiscr-rssee lìgr-rrate sctte ali fbssili di Odonata Zygoptera e descritte due nuove spcciccli OclonataAnisoptera appartencnti alla lami-uliaLibellLrlidae, scoperte nel giacirnen- t6 rtriocenicodi Montc Castellaro.Pesaro.Vicnc itioltrc esatttinataun'ala postcrioredi libel- lirla attribLritaalla sottofirmiglia Pantalirliìec al -uencreTrupa:tt.stigmu, pÍovenientc dalla stessalocalitrì f irssilil-cra.
Abstract
Scvcn fitssil wings of OdonataZygoptera ft'onr the Upper Miocene of Montc Castellaro, Pcsalt, are cliscr-rssecland lìgured, and two nerv speciestt1'Odonata Arlisoptera belonging to thc lhnrily LibcllLrlidac,are dcscribed.A dragonfìy hindu,ing attribr.rtedto thc sr-rbfarnily Pantalinaeand genus Tt'upe:ostigrrafior.n the sarnefìlssililèrous locality, is also exarrined.
Key wgrds: Insecta,Odonata, Coenagrionidae. Lestidae, Sieblosiidae, Cìalopterygidae, LibellLrliclac,Taxotromy, Upper Miocene, Motrte Castellaro.Pesaro, Marches, Central ltaly. lntroduction
New fbssildan-rself'lies and dragonfìies fì'or-r-r the Upper Miocene of MonteCastellaro, havebeen exallined. The Odonatafàuna consists of separatewing relîains, preservedas colllpl'essionsin fìne-grained,biturninotts malls, exposedin twcl f-ossilifèroussections developed along the Adriatic coast, in thelower part of a cliff about3 kpr north-westof Pesaro(Marches, Central ltaly) and includedin the Gessoso-solfìfèraFonnation of Lower Messinianage. Most of the dragonflies "strato havebeen discovered by thesenior author in the degliinsetti" (Gt'NltltNt, I 9tì9.1992" I 993:B,qc;Lr & Gr,NrrL-rNr,2003 ), a well celîerìtedinsect bed about 140
17 mm thick, locatednear the top of the Formationand consistingof very thin, dark- greyand light-grey evaporitic laminae probably related to a seasonaldeposition. In this layer severalinsect orders have beendiscovered, often associatedwith fish and leaf remains.These fossil assemblagesmay indicateboth deepoff-shore and shallow,near-shore paleoenvironments: the presenceof varved shalesnormally occurringin quiet basins,the predominanceof articulatedfish remainsand com- plete insectspreserved as adults(especially Hemiptera-Homoptera, Diptera Bibionidae,Mycetophilidae, Limoniidae and insectswith strongflying ability as Diptera Syrphidae,Hymenoptera and Odonata),may indicatea relatively deep- water/ofÈshorepaleoenvironment, while abundantremains of evergreenneedles and taxodiaceousleaves and the presenceof insectswith weak flying ability (Coleoptera,Trichoptera, terrestrial Hemiptera), may representa near-shore paleoenvironment(WrsoN, 1980).Moreover, the scarcityof marinetaxa among "strato theichthyofauna of the degliinsetti"( only few specimensof Latessp. and Harengula sp. have been found) and the predominanceof the hypereuryhaline speciesAphanius crassicaudus (Agassiz,l832), indicate highly salinewater conditions(SonsrNr,l988), with a normalsalinity marine environment, present at leastin cerlainperiods.The majority of the terrestrialplant assemblages collected in this insectbed, suggestvegetational belts characterizedbya greatdiversity of habitatswith the presenceof Thxodiumin the warm, humid and swampycoastal plains,a belt of thermophyleplants with prevailing Quercus,Acer, Populus and probablemountain slopes with predominanceof xerophileforests of Pinus,Sequoia and Thuja. However,the occurrenceof DipteraTrichoceridae in the insect assemblagesand the high proportionof Nearcticand Palearctic genera among the Odonatafauna, rnight suggest also climatic fluctuations including a possiblecooling phase,correlated with the salinitycrisis in the MediterraneanSea during the Late Miocene,with a vegetationalbelt of coniferslowered towards the depositional basin(BenrolaNr MnncHErrr & MnnrorrrLrrer, 1989); anyway, further collecting and comparativestudy will increaseour knowledgeof this Lower Messinian paleoenvironment. F-ossilAnisoptera(adults and larvae) from MonteCastellaro are relatively common, while fossil Zygopteraareextremely rare: only eightspecimens ( about3.5oA of the total numberof Odonata)have been found up to now. It is interestingto note that speciesbelonging to the subfamilyIschnurinae ( OdonataCoenagrionidae) and the family Calopterygidaeare infrequent in Tertiaryinsect localities and only one recordof the genusIschnura is known from the Eocene/Mioceneamber inclusionsof the DominicanRepublic; on the contrary,the familiesLestidae and Sieblosiidaeare well known from Cenozoicdeposits. Moreover, the finding of the generaDeielia and Trapezostigma(Odonata Libellulidae) is also of considerable interest:the wing of Deielia discoveredat Monte Castellaro,represents the first fossil recordof this genus,while well preservedspecies belonging to the genus Trapezostigmaare poorly known from EuropeanNeogene deposits. Fortunately,a Nature Reserve(Parco Naturale di Monte San Bartolo, including
18 the Monte Castellarooutcrop, considered as insectsdeposit of exceptional preservation), was establishedin 1994and now the protectedarea -uy b. used for research, educationand orgamzedvisits. The most extensivefossil insect collectionsfrom the Monte Castellarosite, includedthe specimensdescribed in this paper, atedeposited in the Museodel Territorioof Riócione(Rimini, Emilia Romagna,Northern ltaly). The venation abbreviationsare basedon Rrpr & KuralovA-pEcr (19g4) with modifications by NEr-et al. (1993)and BEcHlv(1996).The classification of fossil andextant Odonata, follows BEcHI-v(1996,2000)and tie principlesofphylogenetic systematics(sensu HnNNrc 1966,1 969).
Systematicpalaeontology
Order Odonata Fabricius,lT93 Suborder Zygoptera Selys,1854 Superfamily Coenagrionoidea Kirby,1890 Family Coenagrionidae Kirby,1890 Subfarnily Ischnurinae Fraser,1957 Genus Ischnura Charpentier,1840
Ischnura sp. (SpeciesA - Fig. i; Material- Specimenno.1289 and 1289a(part and counterpart);coll. Gentilini; Museo delTenitorio,Riccione (Rimini, Emilia Romagna, Northern ltaly); Gessoso- solfiferaFormation, "Strató Marne bituminoseunit, degli insetti,,llvel; Upper Miocene(Lower Messinian)of MonteCastellaro (Pesaro, Marches, Central ltàiy)
Fig. I - Venation of Ischnurtt sp. (SpeciesA): right forewing of male. Scale:3 mm.
t9 Description The spèciesis known frorn an isolatedright forewingof male with part of the posteiiorwing marginrnissing.Wing membrane hyaline; venation not densewith ielativelylarge cel1s; veins brown' base narrow and well petiolate;wing apexwide androunded; longitudinal veins long and straight. Wing length,15.2 rnn"r; width at nodus,2.4 nm; width at wing base,0.5 rnrn,petiole about 2 mm long; distance fiorn baseto nodus,5 mm; distancefiom baseto arculus,2.3 rnm; distancefiom arculusto nodus,2.2mm; distancefiom nodusto pterostiglna,3.6 rnrn, distance fiorn pterostigrnato apex, 1.8 mm; nodussituated at about32.8 % of the wing lengtlt.Prirnary antenodal crossveins Axl andAx2 alignedand bracket-like, Ax2 in linewith thearculus and situated at 1.1 mrndistal of Ax I ; absenceof secondary antenodaland antesubnodal crossveins. Arculus kinked; sectors of arculusdistinctly separatedfiorn their origin; nodal crossvein in linewith subnodus;nine postnodal crossveinsbetween costal rnargin and RA all alignedwith the postsubnodal crossveinsbelow them excluding the mostdistal incotnplete. Pterostigrna short with prarginsthickened, bicoloured (brown to whiteyellowish), as long as wide and coveringónlyone cell;pterostigrnal brace vein oblique and in linewith the proximal sideof thepterostigl.na; there are three single cells beyond the pterostigrna between theanterior wing marginand vein RA. Discoidalcell trapezoidal in shape,closed basallya1d witS distal angle acute: length of basalside, 0.3 rnrn; length of ventral side,0.9 rlpr; lengthof dorsalside, 0.5 rr-ur;length of distalside, 0.5 urtn.Vein MAb strolglyoblique; subdiscoidal cell relatively wide (length, 1.3 n-rm; width, 0.25 rnm),with distalside angr-rlated; cell sitr,ratedbelow the subdiscoidalcell, unusuallycrossed. Basal area fiee; cubital cell with onlyCr"rP-crossing situated 2.3 pm distalof'the wing base; basal portion of CuA,straight. Postdiscoidal freld with onerow of cells;vein MA andMP basallystraight with onerow of cellsbetween therr-r,vein CuA straightat base. First branching of RP,0.2rnrn basal of thesubnodus; "O", baseof IR2 alignedwith subnodus,lestine obliqr-re vein absent;area between IR2 apclRp3-4, very narrowbasally. Fork of RPI at aboutfburcells ar-rd 4.3 rnrrr distalof the subnodus;vein RPI kinkedbelow the pterostign-ralbrace vcin and slightly zigzaggitrg.Vein IR I separatesfior-r"r RP I , 2.5 tlun distalof the fbrk of RPI andtwo cellsbasal of thepterostigma; vein RP2separates fiom RPI threecells basalof IRI andf-our cells beyond the nodus.One row of cellsdistally between RA arrdRPl. betweenRPI and IRI ( only a dor-rblecell is presenton the wing rnargin),and IRI andRP2. Distal portion of IR2 ztgzagged;double cells between vein.sRP2 and IR2. and IR2 and RP3-4 on the hind rnargin; the vein RP3-4 reachcs theposterior wing borderwell befbrethe proximal side of thepterostigma.
Discussion The fbssildarnselfly wing describedabove can be attributedto the extantgenus Isc,hnurubecause of the followingvenational characters: srnall size of thewing, distipctlypetiolate; venation rather open; wing apexfairly roundedin f-orewing; analve in separatingfiorn the posterior wing borderbef-ore the CuP-crossing; arculus
20 situatedat thelevel of thedistal antenodal; discoidal cell basally closed with distal angleacute; area between the basesof IR2 and RP3-4,narrowed; pterostigma bicolouredin male speciesof the genus,especially on the fbrewing;fork of RP generallysituated about fbur cellsdistal of thenodus in forewingsand three cells in hindwings.Moreover, this fossil species shows an unusualfeature: the cell situated belowthe subdiscoidal cell is crossednearly in themiddle; this character is usually not presentin Coenagrionoideaand probably rnay be consideredas an aberration ratherthan a diagnosticf-eature.The genus Enullugtl'te resernbles Isc:hnura in lnanycharacters. but it differsin havingfìve-eight cells instead ofthree-four, between subnodusand fbrk of RPI; moreover,the wing apexof Enallagntuis slightly pointedand more der-rselyreticulated. The f-ossilIsc'hnttra (Species A) shares synaponlorphieswith thefollowing taxa introduced by Bnc'rtv( 1996):Euzygoptera (longitudinalveins straight and long; one row of cellsbetween CuA andposterior wing rnargin;only primaryantenodals Axl andAx2 presentbasal of the nodus; antesubnodalarea witl-rout crossveins as well as areabetween RP and MA, from arculusto rnidfork),Coenagrionomorpha (postnodal and postsubnodal crossveins in alignrnent,with the rnost distal sometimes incornplete; oblique vein "O" secondarily abser-rt;discoidal cell closed at base in fbrewings);Coenagrionifbrmia (pterostigma rathershort with two crossveinsbelow it; subnodusir-r line with thebase of IR2; nurnberof exagonaland pentagonal cells strongly reduced; area between RPI and RP2with two rowsof cellsseparated by the vein IRI;postnodalcrossveins in aligr-rmentwith crossveinssituated below; intercalary veins lacking, except IR1 and IR2); Coenagrionida( RPI kinkedbelow the pterostigmalbrace vein); Cenagrionodea( proxirnal,ventral and distal rnargins of pterostigma,thickened and fonninga U-shapedstructure); Coenagriorridae (Kirby, 1890),(vein MAb very obliqueso that the dorsal side of thediscoidal cell is shorterthan the ventral side). Thefbssil record for thegenus Isc'hnuru is poor: only one species certainly belonging to this genus,has beendescribed by Br,c'rrlv(2000a) fiorn Dominicanamber inclusions(Eocene to Miocene),while speciesof Coenagrionidaeor Ischnurinae incertaesedis, have been reported by Nr,r-& PnpnzrnN( 1990) and by Nr,L,MnnríNr,z- Dt'lc'Los,Pnt'll,tt & Oult.ttto( 1997), fiorn the Upper Oligocene ofAix-en-Provence and the Upper Mioceneof Sainte-Reine,France. Isc:hnura (Species A) differs fiom tlreDorninican amber species Isc'hnura velÍeni(Bechly, 2000), in thelonger andwider wing, the longer petiole and the cell below the subquadrangle, crossed. Moreover,it is distinguishablefiorn theFrench species by theshorler and narrower wing ( thewing of thespecirnen no.lPM-R07106 from Aix- en Provence,France, hasthe same length ( 15.2rnm) but it is wider),by theshape of thepterostigrna not rhornboidal,the crossedcell below the subquadrangleand the higheror lower numberof postsubnodalcrossveins. At tlrepresent day the genusIsc'hnttru is a cosmopolitan,widespread genus and it is representedin continentalEurope and westernMediterranean basin by seven species.
21 Ischnurinae (SpeciesB - Fig. 2)
(Genusand speciesundetermined) Material- Specimenno. 1288and 1288a(part and counterpart);coll. Gentilini; Museodel Tenitorio, Riccione (Rimini, Emilia Romagna,Northem ltaly); Gessoso- "Strato solfiferaFormation, Marne bituminoseunit, degli insetti" level; Upper Miocene (Lower Messinian)of Monte Castellaro( Pesaro,Marches, Central Italy).
Description The speciesis representedby an isolatedleft hindwing with a small portion of the posteriorwingmargin, missing.Wing hyaline with basenaffow and distinctly petiolate; veinsbrown, wing length,l4 mm; width at nodus,2mm; width at wing base,0.4 mm; petiole,2 mm long;distance from baseto nodus,5.1 mm; distancefrom base to arculus,2.7mm, distancefrom arculusto nodus,2.5 mm; distancefrom nodus to pterostigma,7 .3 mm;distancefrom pterostigmato apex,1.8 mm; nodussituated at about 36.4% of the wing length.Primary antenodals Axl andAx2 alignedand bracket-like;Ax2in linewith thearculus and situated I mm distalofAxl, absence of secondaryantenodal and antesubnodalcrossveins. Arculus kinked and sectors of arculusseparated from their origin; sevenpostnodal crossveins in line with the correspondingpostsubnodal crossveins, excluding the most distal,incomplete. Pterostigma(length, 0.5 mm; width, 0.3 mm) rhomboidalin shapewith proximal andventral sides thickened: some traces of palebrown colourpattern are preserved on themembrane. The pterostigmais well braced(its oblique basal side is aligned with the brace)and covers about three quarters of a cell; thereare four singlecells with the first well elongate,beyond the pterostigmabetween the anteriorwing marginand vein RA. Discoidalcell closedbasally and with distal angleacute: lengthof basalside,0.25 mm; lengthof ventralside, 0.8 mm; lengthof dorsalside, 0.4mm; lengthof distalside .0.4 mm. Subdiscoidalcell moderatelyelongate with distalside angulated; cell situatedbelow the subdiscoidal cell freeof crossveins.Vein MAb obliqueand aligned with subdiscoidalveinlet; basal space free; cubital cell
2
Fig. 2 - Venationof Ischnurinae(Species B): lefi hindwing' Scale:5 mm.
22 with CuP-crossingstrongly oblique and situated at about2.2 mmdistalof thewing base;anal vein separatingfrom the wing margin,0.25 mm basalof CuP-crossing. Discoidalfield naffow with only onerow of cells;vein MA andCuA straightin the first portionand zigzaggeddistally; one row of cellsbasally between veins MP and CuA'''''and betweenCuA and posteriorwing margin.First branchingof RP at 0.2 mm basalofthe subnodus;baseof IR2 in linewith subnodusand basal area between IR2 and RP3-4,very narrow.Vein RP2 separatesfrom RPI three cells basalof IRI and two and a half cells beyondthe nodus.VeinRP1 kinked below the pterostigmalbrace vein; origin of IRI, 3 mm distalof theorigin of RP2and one cell basalof the pterostigma;there are doublecells between RA and RPl, Rpl and "O" IRl,IRI andRP2, on thewing apex.Lestine oblique vein absent;vein IR2 stronglyzigzaggeddistally with doublecells on thehind margin;vein RP3-4slightly sinuousand reachingthe posteriorwing borderopposite the distal side of the pterostigma.
Discussion The speciesshows venational features of theextant subfamily Ischnurinae (wings petiolateto level of CuP-crossingand analvein separatingfrom posteriorwing borderclose to CuP-crossing;arculus situated at levelofAx2; discoidalcell closed basafly andwith distalangle acute; area between the bases of IR2 and RP3-4 very narrow),but the reliableattribution to a genusis difficult for the lack of body characters.This specimenof Ischnurinae(Species B) differs from Ischnura (SpeciesA) describedabove, in the shorterand narrowerwing, the apexsmaller and pointedand the pterostigmascarcely coloured and rhomboidalin shape. Moreover,the cell situatedbelow the subquadrangleis free of crossveinsand thereare seven postnodals instead of ninebetween the costal margin and vein RA; thefork of RP is at abouttwo anda half cellsdistal of thesubnodus instead of four cellsand the origin of IRI is only one cell basalof the pterostigmalbrace vein, insteadof two cells.The Ischnurinae(Species B) differsfrom the amberspecies Ischnuravelteni (Bechly, 2000), in the longer and wider wing and the longer petioleand wider discoidal cell; moreover, it is quitedistinct from Coenagrionidae or Ischnurinaeincertae sedis from France,discussed in thedescription of Ischnura (SpeciesA), in the shorterand narrower wing, in the lessnumerous postnodal and postsubnodalcrossveins and in havingtwo anda half cellsin thebasal area between RPI andIR2, insteadof four or five cells.According to NEr-& PnrcHElEn(1993a), fossilCoenagrionoidea are difficult to recognizeonly by wing remainsand several determinations are uncertain.
Family Lestidae Calvert,1901 Subfamily Lestinae Calvert,l90l Genus LesÍes Leach,1815
ZJ Lestes sp. (SPeciesC - Fig. 3) Gentilini; Material- Specimenno. lzgi and l2gia (part and counterparl);coll. Gessoso- Museodel rérritorio, Riccione (Rimini, Emilia Romagna, Northern Italy); "strato level; LJpper solfrferaF-ormation, Marne bituminoseunit, degli insetti" Italy)' Miocene(Lower Messinian) of MonteCastellaro (Pesaro, Marches, Central
Description half of the The spéciesis representedby a single,hyaline left wing with the distal venationof the membranestrongly folded io*ardi the costalmargin.The wing wing length, specirnenhas beJn reconstructed by partand counterparl.Veins brown; I mm long; l7 rnrn;width at nodus,I .2 mm; width at base,0.5 mrn; petiole about '9 mm;distance distancefiom baseto nodus,5 mrn;distance from baseto arculus,2-4 distance fiorn baseto nidf-ork,2.9 mrn; distancefrorn arculus to nodus,2'5 mm; distance fiorn'odus to pterostigma,8.5 mrn; distance from nodusto apex,l2'3 rnrn; IR2' l.tì mrn, fior' nodusto originoÍRp2.2.6 rnm; distance fiom nodusto originof 29 oAof distancefiorn noAu,to origin of Rp3-4, 1.9mrn; nodussituated at about in thewing length.primary antenodals Ax I andAx2 alignedand bracket-like;Ax2 and line with the arculusand 1.1 rnrn distalof Ax1. No secondaryantenodal crossveins' antesubnodalcrossveins; basal area between RPI and IR2, lree of more or less Sectorsof arculgsseparated at the base;nine postnodalcrossveins RPI' except alignedwith the postsubnodalcrossveins situated between RA and and fbr the lasttwo incomplete.Pterostigma coloured pale yellowish, elongate is 1.4 braced.with theoblique basal side aligned with thebrace.The pterostigma below the mm long and 0.35 mrn wide; thereare two crossveinspreserved and pterostigrnaand six singlecells beyond it, betweenthe anteriorwing margin strongly veinRA.Discoidal cell narrow, elongate, basally closed with distalangle of dorsal acute:length of basalside,0.2 rnm; length of ventralside, I mm; length CuP- side,0.6 rnn; lengthof distalside,0.4 mrn. Basal space fiee; cubital cellwith fiom ving crossingsituated at 1.7rnrn distal of thewing base;anal vein separating area; rnarginat the level of Cup-crossing.only onerow of cellsin thecubito-anal enddistinctly vein CuA srronglyzigzagged distalìy and vein MP straightwith distal one benttowards the ilna màigin.Basal and distal portion of vein MA zigzagged; the posterior row of cellsbetween MA and MP; vein RP3-4straight and reaching RP3-4 wing marginwell beyondthe distalside of the pterostigrna;area between subtending andMA wideneddistàlly; a longand straight intercalary longitudinal vein basesof two rows of cells,presént between veins RP3-4 and MA. Spacebetween of the IR2 and Rp3-4 very narrow:first branchingof RP situated1.8 mm basal nodalcrossvein subnodus;distance tetween originof IR2 andsubnodus, I '7 mm; "O" RP2and andsubnodus well obliquein porition.Lestine oblique vein between RPl,2.4 IR2, situated4.5 mm distalof the subnodus.Vein RP2originates from mostof mrndistal of thesubnodus; course of vein IRI strongly zigzaggedbasally; RA and distalarea between Rp1 andIR2 missing.one row of cellsbetween veins wing apex' Rpl andtwo-three rows of cellsbetween lR2 and RP3-4towards the
/-+ Discussion This fossillestid can be attributedto theextant genus Lestes because of thefollowing venationalfeatures: wing hyalineand well petiolated,sectors of arculusarising nearthe upper part of thearculus; discoidal cell with distalangle well acute,origin nearerto arculusthan to subnodus;pterostigma elongate, rectangular in of RP3-4 "O" shapeand as long as the two cells underit; obliquevein situatedbetween veinsRp2 and IR2; analvein separatingfiom posteriorwing rnarginat levelof CuP-crossing;area between RP3- 4 andMA wideneddistally, with secondary longitudinalveins present. The species shares synapornorphies with thefbllowing taxàintroduced by Br,c'rn-v(1996): E,uzygoptera (longitudinal veins straight and long;end of vein RP3-4situated beyond the pterostigna;vein MA moreor less 1..uò1,ingthe pterostigrna; vein CuA endingdistal of thernidwing; one row of cells betweenCuA and the posteriorwing margin;only Axl andAx2 presentin the antesubnodalarea); Eulestifonr-ria (vein MP curvedafter its origin on the distal angleof t5ediscoidal cell); Lestodea (arcr.rlus shifted towards the wing base,beneath the-prirnaryAx2); Lestinoidea(Calveft, I 901 ), (rnidfbrkrecessed basally and situated atabour 20-26%of the wing length;vein MA zigzaggeddistally); Lestidae ( dorsal arcularbracket reduced; distal discoidal vein MAb veryoblique and discoidal cell with distalangle well acr-rte;three rows of cellsbetween IR2 and RP3-4on the hindmargin); Lestinae (lower portion of thearculus f-onned by thebasal side of the discoidalcell, so that it is veryclose to vein RA). Severalspecies belonging to the genusLcsÍe,s have been discovered in E,ocene,Oligocene, Miocene and Pliocene aepositsof E,urope:the speciesfì'orn the Monte Castellarosite, rnay be placedin tlref-ossil Lestas group B (Nr,r-& Pnrc'rrr,Lr,n.1994), a grollpcharacterized by lxore tnodernvenational characters, because of tl"reabsence of a sr-rppletnentaryrow ot cellsbetween veil MP andthe posterior wing margin.In the Palaeogenespecies of Lcstc.sgroup A, fbundin westernE,urope localities, three-fbur rows of cellsare presentbetween vein MP andthe hind rnargin. Concerning ltalian fbssil lestids' a ibre*i1rgand a hindwingbelonging to the subfarnilySyrnpecrnatinae and genus not describedby Cìnvnllo& $,rrlrocirrr(Selys, 1840), have been fìgured but Gnr.ì-,,'rt'(1987), from the UpperMiocene (Lower Messinian)of the Scaparoni andPiobesi sites, near Alba (Piedmont,Northern ltaly).
Scale: 3 tl'tt't.t' Reconstrgclign of'lefi *i1,s gl' Le.sle.ssp. (Specics Cì) by part irtld coutrtcrpart.
25 At the presentday the genusLestes is cosmopoliteand occursin vegetatedareas with still, sluggishor moving water and in cool temperateto tropicalclimate. Six speciesof Lestesare representedin Europe.
Lestidue(Species D - Fig. a)
(Genusand speciesundetermined) Material- Specimenno. 1290and 1290a(part and counterpart);coll. Gentilini; MuseodelTenitorio, Riccione (Rimini, Emilia Romagna, Northem ltaly); Gessoso- "Strato solfifera Formation,Marne bituminoseunit, degli insetti" level; Upper Miocene(Lower Messinian) of MonteCastellaro ( Pesaro,Marches, Central Italy).
Description The speciesis known from head,thorax and abdomenfragments and from the basal areasof fore and hindwings.Head in poor condition; thorax dark brown to black coloured:mesepisternum about4.3 mm long and I mm wide; mesepimeron3.5 mm long and about0.7 mm wide; preservedlength of dark brown abdomen,13.J mm;
4
Fig. 4 - Vcnationof fore and hindwing basesof Lestidae(Species D). Scale:2 mm
26 width about I mm. Left forewing:preserved length ofwing' 10 mm (right forewing, about9 mm); width at base0.7 mm; lengthof petiole3.1 mm; distancefrom baseto nodus,8 mm (right forewing, 8 mm); distancefrom baseto arculus,4.1 mm (right forewing,4.8mm); distancefrom arculusto nodus,3.7 mm(rightforewing, 3.5 mm); distancefrom baseto midfork, 5.2 mm (right forewing,5.5 mm). Primaryantenodals Ax 1 and Ax2 aligned;Ax2 in line with the arculusand situatedI .6 mm distalofAx I . Basalspace without crossveinsand cubital cell free,except for CuP-crossingsituated 3.3 mm distalof the wing base.Anal vein separatingfrom hind margin0.5 mm basal of CuP-crossing.Sectors of arculusseparated at origin; discoidalcell trapezoidalin shapewith distalangle not preserved.Discoidal cell of left forewing:length of basal side,0.4mm (rightforewing,0.3 mm);preserved length ofventral side,0.9 mm (right forewing,I mm); lengthof dorsalside,0.4 mm ( right forewing,0.5mm); preserved lengthof distalside, 0.4 mm ( rightforewing, 0.5 mm). Two singlecells present basally betweenveins RP and IR2, beforethe subnodus.Left hindwing in poor condition. Righthindwing: preserved length, 8.5 mm; distancefrom arculusto nodus,3.1 mm; Ax2 alignedwith arculus;cubital cell with only CuP-crossing,present; anal vein separatingfrom posteriorwing margin 0.1 mm basalof CuP-crossing;sectors of arculusseparated at origin.Discoidal celltrapezoidal in shapewith distalangle acute: lengthof basalside, 0.35 mm ( left hindwing,0.35 mm); lengthofventral side, I mm (ventralside ofleft hindwingincomplete); length ofdorsal side' 0.45 mm (lefthindwing, 0.35mm); length ofdistal side, 0.7 mm (distalside of lefthindwing not preserved).Vein MAb alignedwith subdiscoidalveinlet; basal portion of MA, zigzagged;one row of
cellsbetween veins RP3-4 and MA andbetween MA and MP"'Jbasallv.
Discussion The fossil wings describedabove belong to the family Lestidaebecause of the followingvenational characters: analvein separating from hind marginslightly basal of CuP-crossing,distal discoidal vein MAb stronglyoblique and distalangle of discoidalcell well acute;vein MA zigzagged;vein RP3-4with origin nearerto arculusthan to subnodus.Thesize of thediscoidal cell slightly smaller in forewings than in hindwingsis a venationalcharacter present in the genusLestes, but the areabetween nodus and apex is missingand it is very difficult to determinethe genusonly by thewing basevenation, because of the similaritiespresent in allied generaas Chalcolestes(Kennedy,l920), Paralesîes(Schmidt, l95l ) and Platyles tes (Selys,1862).
Lestidae(Species E - Fig. 5)
(Genusand speciesundetermined) Material- Specimenno. l29l;coll.Gentilini;Museo del Territorio, Riccione (Rimini, Emilia Romagna,Northern ltaly); Gessoso-solfiferaFormation, Marne bituminose "Strato unit, degli insetti" level; Upper Miocene (Lower Messinian)of Monte
21 Castellaro(Pesaro, Marches, Central ltaly).
Description The speciesis known only frorna right wing base.Veinsbrown; preserved length of wing,[ì mrn; width at base,l mm, petioleabout 1.9 mm long;distance from base to nodus,5.1 rnm;distance fiorn baseto arculus,2.9 rnrn;distance l'rom arculus to nodus,2.2 nm; distancefrom nodusto originof RP2,2.4 mn. Prirnaryantenodals Ax I andAx2 aligned:Ax2 in linewith thearculus and about 0.7 rnm distal ofAx 1; sectorsof arculusseparated at origin.Discoidal cell basally closed, trapezoidal in shape,narrow and elongate: length of basalside 0.2 mrn; length of ventralside 1.2 mrn;length of dorsalside 0.6 mr-n; length of distalside 0.7 rnm.Basal discoidal field with onerow of cells;basal space fiee, cr"rbital cell with onlyCuP-crossing situated 2.4rnm distalof thewins baseand nearer to Axl thanto Ax2.
Discussion The speciesis placedin the extantfarnily Lestidaebecause of the fbllowing venationalcharacters: wing well petiolated,arculus at levelofAx2; distaldiscoidal vein MAb very oblique;discoidal cell trapezoidalin shapewith well acutedistal angle.The lower portion of thearculus is fbnnedby thebasal side of thediscoidal cell, situatednear the baseof the vein RP.Unf-ortunately, the specirnenis too ir"rcornpletefbr genusplacing. Taxon Caloptera Belyshev& Haritonov,l9tì3 Farnily Sieblosiidae Handlirsch"1907 Subfàmily Ischnurinae Fraser.l95l Genus Itulole,ste.r('l) Nel, Petrulevicir-rs,Gentilini & Martínez- Delclos(in press)
Fig. 5 - Vcnationof'uing bascof'l,csticlac (Spccics E,). Scalc I rrnr.
28 Italolestes (?) sp. ( SpeciesF - Fig. 6)
(Species undetermined) Material- Specirnenno.1292a (counterparl); coll. Gentilini; Museo del Territorio, Riccione(Rimini, Ernilia Romagna, Northern ltaly); Gessoso-solfifèra Formation, "strato Marnebiturninose unit, degli insetti"level; Upper Miocene(Lower Messinian)of MonteCastellaro (Pesaro, Marches, Central ltaly).
Description The speciesis knownfiorn a hyaline,rounded wing apexin goodcondition; the venatior"risfàirly dense, consisting of nnmerousrectangular, pentagonal or hexagonal smallcells. Preserved length of wing, 11.2 rnrn; maxitnutn width, 7.3 mrn, distance fiorn proximalside of pterostigmato apex,6.5 mm. Thereare six postnodal crossveinspreserved bef'ore the pterostigma, not in linewith thefour postsubnodals situatedbetween veins RA andRPI;pterostigrna (length,3.8 rnrn; width,l rnrn) veryelongated, pale brown coloured arrd with upperand lower tnargins thickened. Pterostigrnalbrace vein in linewith theproxirnal side of thepterostigrna. There are aboutseven and a half cellsunderthe pterostigrna and ten singlecells beyond it, betweenthe costalrnargin and vein RA. Area betweenRA and RPI below the pterostignla,nan'owed, veins RPI and IRI gentlycr-rrved underthe distal side of theptcrostigma with only onerow of cellsbetween them.Three rows of cellsare presentbetween veins IRI and RP2 and betweenRP2 and IR2, bef-orethe pterostigma;area between RP2 and IR2 wideneddistally, with ninerows of cells; endof veinsIR2 and RP3-4strongly curved towards the posteriorwing margin with only one row of cellsbetween thern.Vein RP3-4 reaclting the hind tnargin
6
(r - Vcnation 29 oppositethe proximal sideof the pterostigma.Distal portionof veinsRPl, IR1, RP2 and IR2, slightly zigzagged. Discussion This damselflyis placedin the fossil family Sieblosiidaebecause of the following venationalcharacters: wing relativelybroad and denselyreticulated, pterostigma elongateand fairly wide so that the areabelow the pterostigmabetween RA and RPI is very narrow.Veins RPI and IRl curvedbelow the distalpart of the pterostigma;areabetween RP2 and IR2 wideneddistally; one row of cellsbetween IR2 and RP3-4.The specimenmuch resemblesthe new genuslîalolesles (Nnr- et al., in press)recently discovered in the easternfossiliferous section of the Monte Castellarosite,, especially in theshape and size of thepterostigma and the numerous crossveinsbelow and beyondit. Moreover,it is quite similar to ltaloleslesin the followingfeatures: the veins RPI andIRI arenot stronglycurved under the distal sideof thepterostigma and include only onerow of cellsbetween them; there are threerows of cells betweenveins tRl and RP2 and the distalend of RP3-4is stronglybent towards the posterior wing margin.Unfortunately,the lack of the first half of the wing with importantvenational characters, does not permit a sure attribution. Fossildamselflies belonging to thefamily Sieblosiidaeare especially known from easternand westerndeposits of the Oligoceneand Miocene of E,urope,with six generadescribed up to now: Stenolestes(Scudder, 1895); Oligolesres (Schmidt, 1958);Paraoligolestes (Nel & Escuillié,1993); Parastenolestes (Nel & Paicheler, 1994);Italolestes and Miostenolestes(Nel, Petrulevicius,Gentilini & Martínez- Delclòs,in press). Taxon Caloptera Belyshev& Haritonov,1983 Taxon Calopterygoidea Selys,1850 Family Calopterygidae(?) Selys,l853 Calopterygidse ('/) (SpeciesG - Fig. 7) (Genusand speciesundetermined) Material- Specimenno. 1735and 1735a(part and counterpart);coll. Gentilini; Museodel Tenitorio, Riccione (Rimini, EmiliaRomagna, Northern Italy); Gessoso- "Strato solfifera Formation,Marne bituminoseunit, degli insetti" level; Upper Miocene(Lower Messinian ) ofMonte Castellaro(Pesaro, Marches, Central ltaly). Description The speciesis known from a wing fragmentwith the base and apex missing. Preservedwing length, 19 mm; maximum width, 8 mm; wing membraneentirely brown colouredand densely reticulated; veins brown; antenodalcrossveins in poor 30 condition.There are32 postnodalcrossveins preserved rarely coincidingwith the crossveinsbelow them,between veins RAand RPl.Vein IRI with origin about eightcells distal to nodus;first onerow of cells,then two rows arepresent between RPI andIRl;course of vein RP2fairly straight;vein lR2 sinuous;one row of cells betweenRP2 and IR2 with two cell rows in the median/distalportion; three rows of cells preservedbetween veins IR2 and RP3-4 and betweenRP3-4 and MA, endof vein MP curvedtowards the hind margin;threerows of cellspresent between veins MA and MP.Thecubital and anal areaare poorly preserved. Discussion The specimenis too incompletefor an accurateattribution, therefore the fossil wing hasbeen placed in thetaxon Calopterygoidea and provisionally in thefamily Calopterygidae,because ofthe followingvenational characters: wing broad,densely reticulated(venation fine and close) and brown coloured, longitudinal veins curved towardsthe posteriorwing margin;postnodal crossveins not alwaysaligned with crossveinssituated below them, between veins RA andRPI;origin of IRI relatively closeto thenodus, vein IR2 sinuous. FossilCalopterygoidea are rare in Cenozoicand Neozoicdeposits and only the following generaand speciesare known, until now. Eucalopteryxatavina (Cockerell,1920) from theMiddle Eocene of theGreen River Formation,Wyoming (U.S.A.),may be placedin the taxon Calopterygoidea(family uncertain); Calopteryxatratu (Selys,1853), (sensu Esnrt & AsnHtNe, 1957)from the Upper Plioceneof the Oya-Formation,Kazusa (Japon), has to be consideredas undeterminedspecies of the genusCalopteryx; Sapho armissani (Nel, 1987)is known from the Oligocene-Miocenedeposit of Armissan(France); Umma sp. incertaesedis and Calopterygoidea( family, genusand speciesincertae sedis), 7 Fig. 7 - Venationof Calopterygidac('?) (Species G): left wing. Scale:3 mm. 31 havebeen described by Nrl & PntcnElun( 1992),from the Upper Oligoceneof Aix-en Provenceand frorn the Mio-Plioceneof Sainte-Reine(France); two specimensof Calopterygoidea(family, genus and species incertae sedis), are known from the Eoceneof the GreenRiver Forrnation,Wyoming (U.S.A.); Calopter.yx ondonc:ensishas been described by Nrr-& Bntsnc'(1994a), fiom theUpper Miocene of Saint-Bauzrle.Ardeche(France). Concerning the ltalianCalopterygifonnia, an EoceneZygopfera belonging to the fossil farnily Bolcathoridaeand the genus Bolc'othorz,has beendescribed by Gr-.Nrtr-rNr(2002) from the Monte Bolca site (Verona,Northern ltaly) and a hindwingbelonging to thefarnily Epallagidae and genusEpulloge (Charpentier, 1840), has been figured but not describedby Cnvnl- r.o& Gnrlr,rrr(1987), fiorn the UpperMiocene of Costiglioled'Asti (Piedmont, NorthernItaly). E,xtantCalopterygoidea occllr in ternperate,warm ternperatehabitats and evenin torrid zones.The taxon is representedin WesternEurope by two fàrnilies,two generaand six species:the fan-rily Calopterygidae with thegenus Calopte4ur (Leach, Itll5) and fìve speciesand the farnilyE,pallagidae, with the genusEpulluge (Charpentier,ltì40) and or-rly one species. Suborder Anisoptera Selys,1854 Family Libellulidae Leach,1tì 15 Subfamily Syrnpetrinae Tillyard,1917 Genus Dcieliu Kirby,1889 Deielia sarue n. sp. (Fig. tl) Holotype- Specimenno. 1293,1293a(part and counterpart); coll. Gentilini; Museo delTcrritorio, Riccione (Rimini, Emilia Rotnagna, Northern Italy). Typelocality - MonteCastellaro (Pesaro, Marches, Central ltaly). Typehorizon - UpperMiocene, Lower Messinian, Gessoso - solfìfèra Fonnation, "strato Marnebituminose unit, degliinsetti" level. E,tyrlology Thisspecies is dedicatedto Sara,granddaughter of thesenior author. Diagnosis Thencw species can be distinguished by thefbllowing venational characters: wing relativelyshort, broad. densely reticulated and crossed by a browrrband. Arculus situatedbetween the 1ìrst and second antenodals; sectors of arculusstalked; discoidal triangletwo-celled; vein MP short,curved and reaching the posterior wing rnargin crppositethe level of thenodus. Vein CuA ztgzaggedand strongly bent towards the hind rnargin;pser-rdo-lRl originates beneath the rniddleof the pterostigrna;RP2 sinuous;Rspl subtending two lows ol'cells.Pterostigr-r'ra pale brown to yellowisl-r, rrnbraced.elongate and sttrtttotrntirtg tu o cl'ossreins. -)a Description The speciesis known from a left forewing characterizedby a brown crossband betweenthe nodus and the proximal side of the pterostigma:the colour pattern coversthe areabetween vein RP1 and branchesof Mspl and it does not reach rearwardthe hind margin. Lengthof forewing,23 mm; width at nodus,8.5 mm; distancefrom baseto nodus, 1l mm (thenodus is situatedat about48% of thewing length);distance from nodus to pterostigma,6.7 mm; distancefrom nodusto wing apex,I 1.7mm; distancefrom baseto arculus,2.8mm.There are eight antenodal crossveins between costal margin andScP, all alignedwith the correspondingcrossveins between ScP and RA. Seven antesubnodalcrossveins are present basal of the subnodus,with a moderately "cordulegastrid short gap" (sensuBEcHLy, 1996); there are ieven postnodalcrossveins betweencostal margin and RA andonly threepostsubnodals between RA andRpl, with the first two not in line with the correspondingcrossveins situated " above;a long libellulidgap" present directly distal ofthe subnodus.Pterostigma (length,2.7 mm; width,0.7 mm) elongate,with upperand lower marginsthickened; there are oneand two half cellsbelow the pterostigma and five crossveinsbeyond it, between vein RA and the anteriorwing margin.Arculus nearerto first antenodalcrossvein than to second;sectors of arculusrising from a very short stalk,situated near the baseof MP; hypertrianglefree of crossveinsand with costalside (MA) weakly curved;discoidal triangle transverse, narrow, pointed and divided into two cellsby a crossvein:length of its basalside, 2.1 mm;length of its costalside, I mm; lengthof its distalside, 2.6 mm. Subdiscoidaltriangle three-celled with costalside (pséudo- analvein PsA) slightly concave: four paranal cells are present out ofthe subtriangle. Basalspace free and cubitalcell with CuP-crossingsituated about I mm basalof arculus;anal areawith two-three rows of cells;vein CuA stronglybent towards the wing marginand with five posteriorbranches: three-four rows of cellsare present betweenCuA and the hind margin.Vein MP relatively short and ending on the 8 - Fig. 8 Deielia surae n. sp., holotypeno. 1293:venation of left forewing. Scale:5 mm aa JJ posteriorwing margin,opposite the level of thenodus. Basal postdiscoidal area with three rows of cells and postdiscoidalfield slightly widened distally.Vein Mspl subtendingtwo rows of cells in the medianportion; there are threebranches of Mspl reachingthe wing border.RP3-4 and MA run parallelwith one row of cells presentin the proximal and medianarea and two rows, distally.First branching of RP (midfork),situated3.2 mm basalof the subnodusand origin of IR2 atabout2.5 "O" mm. Vein RP2 alignedwith subnodus,one lestine oblique vein betweenveins RP2 and IR2, situated0.6 mm distalof the subnodus;only one bridgecrossvein basalof the obliquevein; courseof vein IR2 straightexcluding the distal portion moderatelycurved; Rspl long,gently arched, ending on vein IR2 and subtendinga maximumof two rows of cells;about five secondarylongitudinal veins distinctly originatingon Rspland reaching the hind margin.VeinRP2 well undulate:one row of cellsin the basalarea between RP2 and IR2 andtwo rows of cellsin the median anddistal portion. Vein RPl and RP2divergent with threesingle cells basally and "libellulid two-threerows ofcells before the pterostigma; obliquevein" (sensu BecHLv, 1996),strongly bent and moderately sinuous.Vein lRl zigzaggedand forked below thedistal side of thepterostigma; pseudo-lRl originates on RPI beneaththe middle of the pterostigma;three rows of cellsbetween RPl and pseudo-lRland about sevencell rows betweenIRI and RP2distallv. Discussion WithinAnisoptera the new speciesshares synapomorphies with thefollowing taxa introducedby BecHr-v( 1996): Cavilabiata (distal parl of antesubnodalatea, free of crossveins);Brachystigmata ( pterostigmacovering only I -3 completecells; pterostigmalbrace vein absent;RP3-4 and MA parallelup to the hind margin); Paucipostnodalia(BncHlv, 2002),( basalpart of postsubnodalarea free of crossveins);Italoansida (forewing discoidaltriangle divided into two cells;forewing subdiscoidaltriangle three-celled with posteriormargin angulated); Anauriculida (a "libellulidoblique vein" presentin the basalarea between RPI and RP2; Rspl moreor lesscurved and rejoining IR2, distally; vein Mspl well distinctand ending on MA); Libellulida( primary antenodalsindistinct in the forewing;basal area betweenRA and RPI unicellularfor 2-5 cells);Eulibellulida (sectors of arculus stalked;a concavesupplementary sector present between veins RPI and RP2). The specimenshows venational characters of the subfamilySympetrinae (arculus situatedbetween the first and secondantenodals; sectors of arculusjoined for somedistance from origin;costal side of forewingdiscoidal triangle, narrow) and thewing venationmostly resembles that of the genusDeielia. The new speciesis placedin this genusbecause of the following features:wing denselyreticulated andcrossed by a brownband; last antenodal crossvein complete; sectors of arculus rising from a short stalk; discoidaltriangle narrow, pointed and two-celled; postdiscoidalfield with threerows of cells;vein CuA zigzagged;Rspl subtending two rows of cells;pterostigma elongate, covering more than one cell and with upper and lower marginsmoderately thickened. Deieliq sarae n. sp. can be 34 distinguishedfrom the only extantspecies, Deielia phaon (Selys,1883), by the shorterand broader forewing, the highernumber of cells,the subdiscoidaltriangle less elongatedtowards the wing baseand with the posteriormargin angulated. Moreover,it differs in the less numerousdouble cells betweenRP3-4 and MA distally,the vein MP relativelyshort and ending opposite the level of thenodus and the brown marking situatedbetween nodus and pterostigma,that doesnot extend forward to touch the costaand rearwardto touch the posteriorwing margin. It is interestingto note that most of the venationalcharacters listed above are also presentin the genusBrachythemis ( mainly the wing crossedby a colouredband, the sectorsof arculusstalked, the discoidaltriangle and the subdiscoidaltriangle freeor two/three-celledand the vein Rsplsubtending two rowsof cells),but in the extantspecies of this genusthe lastantenodal crossvein is usuallyincomplete. At the presentday, the genusDeielia is representedby a singlespecies limited to specificareas of the Far East:Japan, from Hokkaidoto the Ryukyus,Taiwan, Korea,Manchuria and North and SouthChina.The female specimens of Deielia phaon,oftenshow pale yellowish body and banded wings: in northemJapan species, crossbandsare broad and deeply coloured while in southemareas (Taiwan, Ryukyus and Cina),the wing colour pattern is much degeneratedand often absent.The speciesis usuallyfound in quiet,shallow water environments, where reed maces normallygrow. The discoveryof a fossildragonfly belonging to the genusDeièlia in the Monte Castellarosite, indicates that this genus already existed during the Lower Messinian in westernE,urope and the presentdistribution in EasternAsia is probablyrelated to climaticchanges occurred during the Plioceneand Pleistocenetimes. Subfamily Pantalinae Jacobson& Bianchi.1905 Tribe Trameini Tillyard,1917 Genus Trapezostigma Hagen,1849 Trapezostigmu barbaresii n. sp. (Fig. 9) Holotype- Specimenno. 1295,1295a(partand counterpart); coll. Gentilini; Museo delTerritorio,Riccione (Rirnini, Emilia Romagna, Northern Italy). Type locality- MonteCastellaro (Pesaro, Marches, Central ltaly). Typehorizon - UpperMiocene, Lower Messinian,Gessoso-solfifera Formation, "strato Marnebituminose unit, degliinsetti" level. Etymology- This speciesis dedicatedto a friendof ours,Dr. FabrizioBarbaresi amateurpalaeontologist of Rimini. Diagnosis The new speciesis characterizedby the following venationalcharacters: wing 35 venationfairly dense;pterostigma short and pointed distally; discoidaltriangle narrow,crossed and benttowards the anal area;postdiscoidal field with five rows of cells;vein Rsplsubtending a maximumof four cell rows andvein Mspl a maximum of three rows of cells; vein MP reachesthe hind margin before the level of the nodus. Description The speciesis representedby a left, hyaline forewing with the apex and most of theposterior margin missing. Preserved wing length,43 mm; width at nodus,8.8 mm; distancefrom baseto nodus,23 mm; nodussituated at about50% of the wing length;distance from nodusto pterostigma,l7 mm; distancefrom baseto arculus, 5.4 mm. There are fifteen antenodalcrossveins between costal margin and ScP with the first sevencrossveins in line with the correspondingcrossveins situated beneaththem. Only twelve crossveinspresent between ScP and RA; eleven "cordulegastrid antesubnodalcrossveins with a long gap", preservedbasal of the subnodus.There are ten postnodalcrossveins (the first moderatelyslanted towards thenodus) between costal margin and RA andnine postsubnodal crossveins between RA and RPI more or lessaligned with the crossveinssituated above, except for "libellulid thelast two. A distinct gap" is presentdistal of thesubnodus. Pterostigma ( length2.7 mm;width, 0.8 mm) short,brown, moderately thickened on upperand lowermargins and with proximaland distal sides not parallel: the distal side is fairly concaveand pointed towards the costal margin. Arculus oblique, slightly concave and nearerto the first than to the secondantenodal crossvein; sectors of arculus rising from a relatively short stalk situatednear the lower part of the arculus. Hypertriangle(length, 6.3 mm; width, 0.5 mm) free of crossveinsand strongly elongatedwith costalside gently curved near the arculus. Discoidal triangle narrow, pointed,oblique in positionand four-celled: length of thebasal side, 3.3 mm; length of thecostal side, 1.2 mm; lengthof thedistal side, 4.1 mm. Subdiscoidaltriangle eight-celledand very irregular(strongly zigzagged) on its innerside; pseudo-anal vein PsA,slightly curved only at the distalend. There are ten paranalcells and a 9 Fig. 9 - Tr-apezosÍigmttberhcrre.sii n. sp., holotype no. 1295: venation of left fbrewing. Scalc: 5 mm. 36 maximumof four-five rows of cells in the analarea; basal space free; cubitalcell with CuP-crossingsituated 1.9 mm basalof the arculus.Postdiscoidal area with five rows of cells;Mspl well distinctand subtendingthree rows of cells in the medianpart; veins RP3-4and MA runningparallel in the porlion preserved,with only one rows of cells betweenthem; distal part of MP zigzaggedand ending oppositethe level of thenodus. The firstbranching of RP is situated5.2 mm basal of the subnodusand the secondbranching (origin of IR2),at about3.7 mm. Obli- "O" quevein betweenRP2 and IR2 situatedlmm andone celldistal ofthe subnodus; only one bridgecrossvein strongly bent present below the subnodus;veins RP2 and IR2 parallelsided and with only one row of cellsbetween them; Rspl well archedand subtending a maximumof four rows of cells.Basal area between RPI andRP2 including four single cells: the second crossvein, is developedas "libellulid obliquevein". The apicalplanate (vein IRI) is weaklyconcave and subtends three rows of cellsbefore the pterostigma. Discussion The new speciesshares synapomorphies with the fbllowing taxa introducedby BEcttlv( I 996):Cavilabiata ( distalpart of antesubnodalarea, free of crossveins); Cristotibiata(pterostigma with distalside more oblique than proximal side and with Iengthless than 8 timeswidth); Brachystigmata( pterostigma short and covering only l-3 completecells; RP3-4 and RA parallelup to the hind margin); Paucipostnodalia(BecHlv, 2002), ( basalparl of the postsubnodalarea free of crossveins);Neobrachystigmata (posterior part of the arculusshorter than the anteriorpart); Paneurypalpidomorpha ( anterior margin of hypertriangledistinctly "O" convex, veinMA beingmore or lessarched basally; oblique vein situatedonly I -2 cells distalof the subnodus);Eurypalpida (forewing pseudo-anal vein PsA, hypertrophied);Anauriculida (vein Rspl curved and rejoining the vein IR2 distally; Msplwell distinct);Libellulida (second crossveins between RPI andRP2 developed "libellulid as obliquevein"; basalarea between RA and RPI unicellularfor 2-5 cells);Eulibellulida (sectors of the arculusstalked; primary antenodal crossveins Ax I andAx2, indistinguishable from thesecondaries; a supplementary sector present betweenRPI andRP2; the most distal antenodal crossvein between costal margin and ScPdeveloped as antenodaloblique vein); Pantalinae(Jacobson & Bianchi, 1905),( Rspland Mspl similarto a primarylongitudinal vein, with 2-3 rowsof cells between,in forewingsand/or hindwings). Trapezosfigmabarbaresii n. sp. belongsto the subfamily Pantalinaeand genus Trapezostigma(Hagen, 1849),because of the following venationalcharacters: pterostigmawith endsnot parallelland longer in forewingthan in hindwing;numerous antenodaland postnodalcrossveins not completelyin line with crossveinsbelow them; forewing triangle narrow, elongateand with costal side relatively short; postdiscoidalarea generally with four cell rows; Rspl and Mspl subtendingtwo- threerows of cells (the type genusTramea (Hagen, I 861) hasto be rejectedsince junior objectivesynonym of Trapezosîigma(Hagen, 1849). 37 The genus Trapezostigmahas a relativelypoor fossil recordup to now: only the rp..Èr Trapezostigmamiocenica (Gentilini, 19S9)has been previously described from the UpperMiocene of Monte Castellaroand the baseof a hindwing is known from the Upper Mioceneof Alba (Piedmont,Northern ltaly). On the contrary,the fossilgenus Paleotramea (NEl & Papnztnx,1985), a genusshowing a wing venation - relativelysimilar to Trapezostigma,is known from the Upper Oligoceneof Aix en Provenceand the Oligo - Miocenefossiliferous locality of Bes-Konak(Turkey), where numerousspecimens have been found. The new speciesTrapezostigma barbaresii differs from Tropezostigmamiocenica in the following characters: the first half of the wing is narrowerand more denselyreticulated; the antenodal crossveinsare more numerous ( 15/ 1 2 insteadof l2ll I ), thepterostigma is pointed distally and shorter,the postdiscoidalfield has five cell rows insteadof four, the vein Mspl subtendsthree rows of cellsinstead of two andRspl four rowsof cells, insteadof three. At the presentday, speciesbelonging to the genus Trapezostigmaoccur in the Temperateand Tonid zonesof the whole earth(NEnoHnv & wEsrpR*, 1975). Five otherspecimens belonging to the genusTrapezostigma have been discovered in theUpper Miocene ofthe MonteCastellaro locality (Gessoso-solfifera Formation, "strato degliinsetti" level), but the attributionto a speciesis difficult becausethe materialis too incomplete.A fairly well preservedhindwing is describedand figured below.Available material: specimen no.208 (distal half of a rightforewing); no.l89, l89a (distalhalf of a hindwing);no.202,2\Za(basalhalf of a righthindwing, brown coloured;this specieshas been tentatively placed in the genusTrapezostigma, but importantvenational characters of the distal part of the wing are missingand the attributionis problematical);no.l 296,1296a(righthindwing in quitegood condition, with wing venationfairly similarto thatof the specimenno.1297: the dragonflies mightbe conspecific);no.1297,1291a(a left hindwing). Trapezostigma sp. (SpeciesH - Fig. 10) Material- Specimenno. 1297,1297 a(parl and counterpart); coll. Gentilini; Museo delTerritorio, Riccione (Rimini, Emilia Romagna, Northern ltaly);Gessoso-solfifera "strato Formation,Marne bituminoseunit, degli insetti" level; Upper Miocene (Lower Messinian)of Monte Castellaro(Pesaro, Marches, Central ltaly). Description The specimenis known from a singleleft hindwing characterizedby threebrown spotson thewing base,covering the basal half of thecubital cell, the hypertriangle, the discoidaltriangle, the first threecells of the postdiscoidalarea and a small portionof the membranebetween the fork of Cuspland somecell rows of the anal field. Hindwinglength, 50.4 mm; width at nodus,,11.7mm; distancefrom baseto nodus,19.7 mm; nodussituated at39oh of thewing length;distance from nodusto 38 pterostigma,30.5 mm; distancefrom base to arculus,5 mm. There are seven antenodalcrossveins between costal margin and ScP,all aligned with the correspondingantenodals situated between ScP and RA. Five antesubnodal crossveinsare presentbetween RA and RP/RP1-2,with a relatively long "cordulegastridgap" basal of the subnodus;eleven postnodal crossveins not in line with theten corresponding crossveins between veins RA andRPl; a long"libellulid gap"of the postsubnodalcrossveins is presentdistal of the subnodus.Pterostigma (length,2.8mm; width,0.7mm) short,brown coloured, unbraced and covering one and a half cells.There are five crossveinsbetween the costalmargin and vein RA beyondthe pterostigma.The arculusis slightly sinuousand in line with the basal sideofthe triangle;sectors of arculusstalked. Hypertriangle (length4.I mm;width, 0.6 mm) free of crossveinsand with costalside strongly curved; discoidal triangle free:length of the basalside, 1.5 mm; lengthof the costalside, 3.9 mm; lengthof thedistal side, 3.4 mm; vein MAb straight.Basal space free, cubital cell with only the CuP-crossingsituated about 2.2 mm, basalof the arculus.Anal loop well developed,boot-shaped and including thirfy-five cells; vein Cuspl(midrib) forked, stronglyangulated and with a toe fairly straight.Anal areagîeatly expanded with numerouscell rows between the anal loop and the hind, rounded angle of thewing. VeinCuA sigmoidallycurved and forkedinto CuAa and CuAb; one row of cells betweenveins MP andCuA. Basalpostdiscoidal area with threerows of cellsand discoidalfield stronglywidened distally. Mspl well distinctand subtending only one row of cells;distal part of Mspl zigzagged.Veins RP3-4 and MA runningparallel with onerow of cellsbetween them: distal area between RP3-4 and MA. narrowed: RP3-4and MA reachingthe posterior wing margin,opposite the origin of vein IR I . Firstbranching of RP at 5.8 mm basalof the subnodus;IR2 originateson RPI-2 aboutbelow the most distalantesubnodal crossvein; vein RP2 alignedwith the "O" subnodus;oblique vein situatedbetween RP2 and IR2, 1.7mm distalof the subnodus.No extrabridge crossveinsbetween RP l-2 and IR2. Veins RP2 and IR2 parallelwith only onerow of cellsbetween them; vein Rsplelongate, gently arched,ending on IR2 andsubtending a maximumof threecell rows;seven-eight Fig. l0 - Venationof Trapett.stigmasp. (SpeciesH): left hindw'ing.Scale: 5 mm. 39 rows of cells are presentbetween Rspl and the posterior wing margin in the narrowestpart. Basal area between RP 1 and RP2with four singlecells: the second "libellulid crossvein,developed as obliquevein" is sinuousin shape.VeinIRl elongate, moderatelyconcave in the medianpart and subtendingthree rows of cells before the pterostigma;pseudo-lRl with origin on RPI near the proximal end of the pterostigma. Discussion The fossil wing describedabove belongs to the genusTrapezostigma because of thefollowing venationalcharacters: hindwing long, pointed, very broadand spotted towardsthe base;pterostigma relatively short; anal loop elongate,boot-shaped, includingnumerous cells and with a midrib(vein Cuspl) well angulated;vein CuA sigrnoidallycurved at base;RP3-4 and MA strongly bent towardsthe posterior wing margin,distally. Vein IRI weaklyconcave: first onerow andthen two-three rows of cellsbefore the pterostigmabetween RPI and IRI; vein Rsplsubtending a maximumof threecell rowsand rejoining vein IR2 distally. The baseof a hindwing belongingto the genusTrapezostigma, figured by Cn- vALLo& GnllErrr ( 1987),from theUpper Miocene of Alba (Piedmont,Northern Italy),has been described by NEl & PnrcHElen ( 1993b)in an accountof thetribe Trameini.This wing fragmentmuch resembles the hindwing described above, in the venationand evenin the positionof the threebrown spotspreserved on the wing base,therefore the two specimensmight belong to the samespecies. Moreover,a new genusand a new speciesof Pantalinae(Pisaurum coloretum Gentilini,1989), are known from the UpperMiocene of MonteCastellaro. This speciesis basedon a singleright hindwing and it is quitedistinct from thefossil hindwing of Trapezostigmaexamined above, by the following venational characters:the wing of Pisaurumis relativelyshort and greatly expanded towards thewing base,most of the first half of the wing is brown coloured;there are five antenodalcrossveins, ten postnodalsand six postsubnodalsinstead of seven antenodals,eleven postnodals and ten postsubnodalspresent in Trapezostigma. The pterostigmaof Pisaurum is short,,dilated and has the distal side weakly oblique: in Trapezostigmathe pterostigmais narrow with the distal end well oblique. Moreover,the genusPisaurum differs from Trapezostigmain having the analloop foot-shaped,strongly elongate and entirely developed between the discoidaltriangle and the distalhalf of the cubitalcell, insteadof an anal loop with a long gaff and the heel cells shifted distal of the discoidaltriangle. In Pisaurumthe vein IRI subtendsonly one-twocell rows beforethe pterostigma, insteadof two-three,the vein RP2 is sinuousrather than straightand the vein Rspl is shorterand subtendsonly one-tworows of cells insteadof three. 40 List of the known fossil Odonata from the Upper Miocene of Monte Castellaro Currently225 specimens( 135adults and 90 larvae)in 7 families,l2 genetaandl8 speciesare known from this fossiliferouslocality, as follows: four families of Zygopterawith threegenera and a new speciesand three families of Anisoptera with nine generaand seventeennew species. SuborderZygoptera Family Lestidae GenusLesÍes (species undetermined): a left wing. Family Lestidae(genus and speciesundetermined): body and wing remainsand a basalright wing fragment (this paper). FamilyCoenagrionidae GenusIschnura (species undetermined): a right forewing.Subfamily lschnurinae (genusand species undetermined): a left hindwing(this paper). FamilySieblosiidae Genusltalolestes. Italolestes stroppai (Nel, Petrulevièius,Gentilini & Martínez' Delclòs,in press):a completespecimen. Italolestes (?) (speciesundetermined): a wing apex(this paper). TaxonCalopterygoidea FamilyCalopterygidae (?) (genusand species undetermined): the medianportion of a wing (thispaper). SuborderAnisoptera FamilyLibellulidae GenusSltmpetrum. Sltmpetrum italicum (Gentilini, 1989):twelve forewingsand ten hindwings.Sympetrum elongutttm (Gentilini, 1989):fourteen forewings and nine hindwings.Sympetrum krzeminskii (Gentilini, 1989):a right forewing. Sympetrummarinum (Bagli & Gentilini,2003):the distalhalf of a left forewing. Sympetrumsp.: four specimens.Libellulidae (genus and speciesundetermined): twentyeight specimens. GenusDeielie. Deielia sarae n. sp. (this paper):a left forewing. GenusCelithemis. Celithemis zavattinii (Bagli & Gentilini,2003):a right hindwing. GenusLibellula. Libettula mediterranea(Gentilini, 1989):a right forewing. Zr- bellulaadriatica (Bagli & Gentilini'2003):distal half of a left hindwing. GenusPisaurum. Pisaurum coloratum (Gentilini, 1989): a right hindwing. GenusTrapezostigma. Trapezostigma miocenica (Gentilini, 1989):a left forewing previously describedby the author as Trameamiocenica (Tramea has to be regardedas junior synonym of Trapezostigma).Trapezostigma barbaresii n. sp. (this paper).Trapezostigma sp.: two forewingsand threehindwings (this paper). FamilyCorduliidae 4l GenusEpitheca. Epitheca miocenica (Gentilini,1992): a right hindwing. Epitheca annae(Gentilini,1992): a left hindwing.Subfamily Corduliinae: a left forewing. FamilyAeshnidae GenusAnax. Anax cryptus(Gentilini & Peters,1993): four forewingsand three hindwings.Anax cf. imperator (Leach, 1815): three forewings,two hindwings and two abdomenremains. Anax cf. parthenope(Selys, 1839):a left hindwing. Anax sp.: sevenforewings and threehindwings. GenusAeshna. Aeshna messiniqna (Gentilini & Peters,1993): a right hindwing. Aeshnaghiandonii (Gentilini & Peters,1993): a right hindwing. Aeshna multicellulata(Gentilini & Peters,1993): a left hindwing. Aeshnasp.: a left forewing,, a wing apex,the basal part of a hindwingand a femaleabdomen. FamilyAeshnidae: a wing fragmentand a forewing. Odonatalarvae (family, genus and speciesundetermined): 90 specimens. Conclusions Therecent finding of thefossil family Sieblosiidaeand the discovery of thegenera Ischnura andLestes as well as the occurrenceof severalgenera and speciesof dragonflies,suggest an interestingbiodiversity in the Monte Castellaro paleoenvironment,with the presenceof four familiesof OdonataZygoptera and threefarnilies of OdonataAnisoptera at least.Finds of damselfliesand dragonflies now living in differenthabitats (temperate to subtropicalor tropicalareas) might be relatedto climatic changesof the Late Miocenein the Mediterraneanbasin, as suggestedalso by the lithostratigraphiccharacters of the MonteCastellaro section and by the faunaand flora found in the bituminousmarls of the Gessoso-solfifera Formation.However, there is muchto be doneand further collecting of fossilsas well asstudies on otherinsect orders, which have not yet beeninvestigated, will be necessaryfor increasingour knowledgeof this Messinianenvironment. Ac knowledgemen ts We are gratefulto Dr. GúnterPeters of Berlin and Dr. SyoziroAsahina of Tokyo, for commentson somefossil damselfliesand dragonflies.Our sincerethanks to Dr. GúnterBechly of the StaatlichesMuseum fiirNaturkunde, Stuttgart, for helpful suggestionsand Dr. DanielaGrossi, director of theMuseo del Territorio of Riccione, for accessto palaeoentomologicalcollections. 42 References Bac;rrL. & GENrrLrNrG., 2003 - Nuove libellulefossili del Miocenesuperiore di Monte Castellaro(Pesaro, Marches, Central ltaly) (OdonataLibellulidae). Quaderno di Studie Notiziedi Storia Naturaledella Romagna,18: 37-50. Br.c'Hr-yG., 1996- MorphologischeUntersuchungen am Flùgelgeàderder rezenten Libellen und deren Stammgruppenvertreter(lnsecta; Pterygota;Odonata), unter besonderer Berúcksichtigungder PhylogenetischenSystematik und der Grundplanesder Odonata. Petalura.2:l-402. Bsculy G., 2000 - PhylogeneticSystematics of Odonata.Website on the Internet'.www. bechly.deiphylosys.htm. Br..c'Hr-yG., 2000a - A new fossil damselflyspecies (lnsecta: Odonata: Zygoptera: Coenagrionidae:lschnurinae)from Dominican amber.Stuttgarter Beitrrige zur Norurkunde,Serie B (Geologietrnd Palciontologie).299: l-9. Brc'gryG..2002 - The first fossilrecord and first New World recordfor the dragonflyclade Chlorogomphida( lnsecta: Odonata:Anisoptera:Araripechlorogomphidae n. fam.) from the Crato Limestone(Lower Cretaceous,Brazil). Stuttgarter Beitrcige zttr Naturkunde SerieB (Geologieund Palciontologie),328:l-l l. Bun'rolRNrMnnc'nElrr D. & MRnrorrrLrnrr M., 1989- Le Messiniend'ltalie du pointde vue palynologique.Boltettino della SocietàPaleontologica Italiana,23 (2-3):183-ltl8. CnvnrroO. & Galre r-tr A., I 987- Studidi CarloSturani su Odonati e altri insettifossili del Messinianoalbese (Piemonte) con descrizionedi Oryctodiplaxgypsorum n. gen. n. sp. (OdonataLibellulidae). Bollettino della SocietàPaleontologica Italiana,26 (l- 2\: l5l-116. EsnrrT.& AsnnrNnS., 1957- On two Terliarydragonflies species from theOya-Formation in Kazusa-NagasakiPrefecture . Kontyu.25: 82-86. Gr,N'r'rLrNrG.,1989 - TheUpper Miocene dragonflies of MonteCastellaro (Marches, Central Italy) (OdonataLibellulidae). Memrtrie clella Soc'ietà Entomologica ltalianu,6T (2): 251-21t. Gr:N'r'rr.rNrG., 1992 - Two new speciesof Epithec'afrom the Lower Messinianof Central Italy (OdonataCorduliidae). Bollettino clella Società Entomologica Italirtna,l23 (3): 201-208. Gr,N'r'rr-tNrG.& Pr,'r'r-.nsG.,1993 - TheUpper Miocene aeshnids of MonteCastellaro, Central Italy,and their relationshipsto extantspecies (Anisoptera: Aeshnidae\. Odonatologica. 22(2):t47-tl8. GnNlrr-rNrG..2002 - Fossil damselfliesand dragonflies from the Eoceneof Monte Bolca, Italy (lnsecta:Odonata). Studi e Ricerchesui Giacimenti Terziari di Bolca, Miscella- nea Paleontologica,9: 7-22. Hr,NNrc;W., 1966- Phylogeneticsystematics. University o.f lllinois Press,Urbana: 263 pp. Hr,NNr<;W., 1969- Die Stammesgeschichteder Insekten. Kramer, Frankfurt: 436 pp. Nr,r,prrnuJ. G. & Wr,s-lrellM. J., 1975- Amanual of the Dragonfliesof NorthAmerica (Anisoptera)including the GreaterAntilles and the Provincesof the MexicanBorder. lJniversitvo/'California Press. Berkeley and Los Angeles:615 pp. Nur-A. & PnpnzrnNM.,1985 - Unenouvelle espece d'Odonate fossile du stampiend'Aix-en- Provence(B.d. R. ) (OdonataLibellulida e). L' Entomo lo giste, 4 | (2) .5 5-59. Nr,l A. & PnpnzrnNM., 1990- LesCoenagrionoidea fossiles de l'Oligocènedu sud-estde la France( Zygoptera:Platycnernididae, Coenagrionidae). OcJonatologica,19 (3) : 251-262. 43 Nul A. & PatcHElrnJ. C., 1992- Les Odonatafossiles: état actueldes connaissances. Huitiernepartie : Les Calopterygoideafossiles (Odonata, Zygoptera). Bulletin de la Sociefeentomologique de France, 97 (4): 381-396. Ner- A., MnnrÍNEz-DelclosX., ParcHnr-enJ.C. & HENnornvM., 1993- LesAnisozygoptera fossiles.Phylogénie et classification.(Odonata) . Martinia, 3: I -31 I . NEl A. & P,qrc'uEr-enJ. C.. 1993a- Les Odonatafossiles: etat actuel des connaissances. 7. Les Coenagrionoidea(Odon., Zygoptera). Entomologica gallica,4 (2-3): 59-63. NEr-A. & Pnrc'HEr-nnJ.C., 1993b - Les Libellulidaefossiles. Un inventairecritique (Odon., Anisoptera,Libelluloidea). Entomologica gallica, 4 (4): I 66-190. Nr,r-A. & P,qrc'uEr-r-.r Authors' addresses: GiuseppeGentilini viaAdriatica, 78 | - 47843Misano Adriatico (RN) e- m o i I : g_gentiIi ni. ads 12003 (!l ibero.it LorisBagli Museodel Territorio viaLazio,l0 l-41036 Riccione(RN) e-mail: bagliloris@l ibero. it 44