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Giuseppe Gentilini & Loris Bagli FOSSIL ZYGOPTERA AND

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Giuseppe Gentilini & Loris Bagli FOSSIL ZYGOPTERA AND 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 andstraight. 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
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