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Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1B - Pag.: VOLUME.p0006.pdf 10:56:23 10-11-18

10-11-18 10:56:23 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1B - Pag.: VOLUME.p0003.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1B - Pag.: VOLUME.p0011.pdf 02.p65 10:56:23 mediterranea esub-mediterranea)abbiamocollezionatoun’interessanteassociazionefaunistica,notevoleperquantitàdiesempla aBifronszone aSerpentinum/LevisoniedellabaseZona (secondolavigentezonazionestandard dellepaleoprovincie della esemplari diammonitidelToarciano, spessoottimamenteconservatieanchedigrandidimensioni.NegliorizzontidelToarciano raccolta stata fossiliferi delfrontedicava,cosìcomedallanotevolequantitàdetritirimossadall’attivitàestrattiva,è nodulare emarne),litologicamenteequivalenteaicoevicalcarinodularirossidelRossoAmmoniticoUmbro-Marchigiano.Dai live Hettangiano) alCalcare MaiolicadelCretaceo. NellesezionidellaMarconessa diCingoliilToarciano sipresenta infaciesgri SIELPA, hadenudatounvastofronte dicavaincuiaffioralasuccessionegiurassicaridotta,chevadalCalcare Massiccio(Sin oggetto diinteressedapartedeipaleontologiitalianifindallafinedell’Ottocento.Negliannipiùrecentil’attivitàestrat nella CavadellaMarconessadiCingoli(Macerata).Inumerosifossiliammonitiealtrimolluschicollezionatiinquestaloca Macerata, Italia)] rare show acertainamountofprovincialism thataffectedsomeHildoceratinaelineagesofthelowerToarcian. Apennines, whereitoccurswithotherendemicHildoceratinaefauna.TheseammoniteassemblagesfoundinourApenninessections (1932) tothegenus Leukadiella studio, einoltre new species( ammonite genera,suchas of ammonoidssometimeslarge sized.From thelowerandmiddleToarcian layersoftheMarconessa quarrysectionswecollectedse Here, thenodularlimestoneandgrey marloftheBugarone Formationhaverecorded agreat amountofwellpreserved inn is historicallymeaningful,becauseinthepasttwocenturiesItalianpalaeontologistshavefiguredammonoidspecimensfrom“M where alowerJurassicoutcrop yieldedaremakable Toarcian ammoniteassemblage,veryrichinearlyHildoceratinae() Italy). We foundtheearliest lower Toarcian oftheApennines(MarcheanApennines, Cingolites ISSN 0375-7633 F. Venturi, DipartimentodiScienzedellaTerra, UniversitàdegliStudidiPerugia,PiazzaUniversità,I-06100Italy; S. Sassaroli,Via S.Michele33,I-60030Rosora(AN),Italy;[email protected] collezionato formeappartenentiaigeneripiùtipicidelToarciano inferiore: ottimo statodiconservazioneeilnumerotaxarappresentati.Oltrealnuovogenere the topof 1976) orthe Mouterde, 1967;Pelosio,1968; Elmietal.,1974;Gabilly, Nejdia Frechiella già nota( ammonite zonation fortheMediterranean Province(or biostratigraphic dataneed arevisedearlyToarcian nevertheless, veryquestionable andournew of bothindexes and Elmietal.,1994).Inthe Apenninestheoccurrence Cingolites Umbro-Marchigiano, altre formesembranodistribuiteancheinaltribacinidellaTetide mediterranea.Allostatoattualedellaricerca ilgenere sembrerebbe quindicomprovare l’esistenzadiuncertogradoprovincialismo nellafaunaammoniticadelToarciano inferiore. sembrano essere confinatenellaTetide mediterranea.Ilritrovamento diquesteforme,ilcuiresoconto dettagliatorichiedeult This workconcernstheammonitesofnewgenus RIASSUNTO - ABSTRACT -Newbiostratigraphicdataare provided from theMarchean Apennines(Marconessa quarry, nearCingoli,Macerata,Italy KEY WORDS-Apennines,Tethyan palaeobiogeography, Toarcian, Ammonitina,,. 10-11-18 Dactylioceras e Cingolites Dactylioceras C. stefaninii , sembraesseredistribuitoprincipalmentenell’AppenninoedassociatoadaltreformediHildoceratinaeancorainedite,c Rarenodia , (Hildoceratinae),whichwerecollectedinthe C. clavatus Frechiella Hildaites levisoni -Questolavoro trattaunnuovogenere diammonitiHildoceratinaedelToarciano inferiore raccolticoncriteriostratigrafico and [ Cingolites Hildaites serpentinus Mercaticeras from theMarcheanApennines(Cingoli,Macerata,Italy) ), mainclusadaMerla(1932)nelgenere , H. serpentinus (moltorari).Alnuovogenere Rakusites Nejdia , , 97 Bollettino dellaSocietàPaleontologicaItaliana,49(2),2010,97-118. Modena,31luglio2010 C. picenus FOREWORD Phymatoceras n.gen.,anewlowerToarcian Hildoceratinae(Ammonitina) Hildaites , , , nuovogenerediHildoceratinae(Ammonitina)delToarciano inferioredell’AppenninoMarchigiano(Cingoli, Taffertia Lytoceras , . To thepresent-day thegenus Orthodactylites Cingolites and Zone ( Zone , Orthildaites , , C. spiralis , Neotaffertia and Rarenodia Calliphylloceras Stefano S sensu H.levisoni specimensnear Zone( , , ) andanalreadyestablishedspecies( Mesodactylites , Guex,1976 , Cingolites Praepolyplectus Martanites ASSAROLI , Phylloceras sensu , is, Mercaticeras attribuiamo3nuovespecie( Harpoceras , Cingolites ,

Praepolyplectus Nodicoeloceras &FedericoV , . We establishherein anewgenus Polyplectus

serpentinum other hand,accordingtoDonovan(1958) with SubmediterraneanandSubborealProvinces.Onthe (Page, 2003,2004)unsuitabletocorrelatethisProvince West Tethyan), beinginouropinion theStandardZonation F°124 INO-F°117 IISO;Carta Geologica d’ItaliaF°124 1; CartaIGM Carcatora Mt.andS.Angelo Mt.(seeFig. cut byRudiellecreekintheJurassic sequences,between outcrop islocatedontheorographic leftsideofthevalley Marconessa quarrynearCingoli (Macerata,Italy).This of ammonitespecimensfrom theLiassicoutcropof Europe, arenotsuitablefortheApennines. typical indexspeciesfortheearlyToarcian ofNW , . Fraquesteforme,alcunesembranoendemichedelbacinoTosco- seemstohaveaMediterraneanTethys dispersal,mainlyinthe Taffertia Hildaites We carriedoutabed-by-bedbiostratigraphic sampling , , , Lytoceras , Polyplectus Orthodactylites Rakusites , C. stefaninii ENTURI Orthildaites and 06/09/10, 13.49 Cingolites C. clavatus 4 3 2 1 Device Resolution2400DPI , Harpoceras falciferum Calliphylloceras , , Martanites Phymatoceras ), whichwaspreviouslyassignedbyMerla e , Mesodactylites oggetto delpresentelavoro,abbiamo Hildoceras , C. picenus Cingolites 4x4 3x3 2x2 1x1 , Praemercaticeras 50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 , , Phylloceras , connuovitaxaancorain Mercaticeras [email protected] , towhichweascribethree e tiva, condottadalladitta C. spiralis , unagrandequantitàdi Nodicoeloceras eriori pubblicazioni, , whicharethe , einfineanche , Harpoceras , inferiore delle ) eunaspecie lità sonostati Mercaticeras Leukadiella gia (calcare arconessa”. emuriano- er moulds ri, illoro . Thesite seem to Tetide

, very veral

10-11-18 97

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, , 10:56:23 Data: 10/08/201008:32:49-Lavoro:MD17220-Bollettino_Paleontologia_Vol_49_N_2Segnatura:1BBlack Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1B - Pag.: VOLUME.p0014.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1B - Pag.: VOLUME.p0010.pdf 10:56:23 10-11-18

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10-11-18 10:56:23 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1A - Pag.: VOLUME.p0001.pdf 1 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1A - Pag.: VOLUME.p0009.pdf 10:56:23 10-11-18

YellowMagentaCyan 10-11-18 10:56:23 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1A - Pag.: VOLUME.p0016.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1A - Pag.: VOLUME.p0012.pdf 02.p65 4 3 2 1 Device Resolution2400DPI 10:56:23 Italy. Fig. 1-LocationmapoftheMarconessaquarry, Cingoli(Macerata), 98 Formation, yielded overtheyearsaveryrich fossilfauna, known RossoAmmonitico Umbro-Marchigiano lithologically andchronologically comparabletothewell- stage, whichareinagreymarly-nodular limestone Umbrian-Marchean Apennines. ThestrataoftheToarcian sequences, typicalofthetectonic structuralhighsofthe has uncoveredalargeportion oftheJurassicrock current quarrying,carriedoutbytheSIELPA Company, stone, whichislocallycalled“Marconessastone”.The originated fromtheancientquarryingofornamental Romitorio ofS.AngeloMt.Thisdenominationpossibly 1990). occurs (Ceccaetal., Bugarone FormationoftheUmbrian-MarcheanApennines Maiolica acondensedrocksuccessioncomparabletothe Formation. BetweentheCalcareMassiccioand (Pozza, 1989,1992)andfinisheswiththeCalcareMaiolica This outcropstartswiththeCalcareMassiccioFormation MC andGeologicalFieldTripn°9,inPasseri,1994,Ed.). The siteisknownasMarconessaquarryofthe 10-11-18 4x4 3x3 2x2 1x1 /0 /929 /749 /51/02/03/04/050 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 98 Bollettino dellaSocietàPaleontologicaItaliana,49(2),2010 facies

we subdivideour Undicosta Zone ina section fromthesuddenappearance of being theearlyToarcian post-OAEcharacterizedin our Polymorphum orMirabile zones) oftheseProvinces, the earliestToarcian (excludingtheTenuicostatum and correlated. Hildoceratinaemay bealsousefultocorrelate in theEuropeanProvince, which maybeeasilythus broadly dispersalbothintheWest Tethyan Province and Nejdia Department oftheEarthSciences–PerugiaUniversity. studied andfiguredaretemporarilyhousedinthe (2005). Dezi &Ridolfi(1975,1978)andSassaroliVenturi ammonites oftheMarconessaquarrywerestudiedby sized ascalcareousinnermoulds.Morerecentlysome which areoftenwellpreservedandsometimesverylarge Marconessa quarryaremarkableamountofammonites, number ofamateurscollectedinthedetritus from the“Marconessa”ofCingoli.Overyearsa (1939), haveoftenquotedandfiguredspecimenscoming Bonarelli (1899,1899),Merla(1932)andRamaccioni ammonite specialists,suchasMeneghini(1867-1881), mainly ofammonitesandothermolluscs.Severalearlier Guex, 1973and shapes arealreadywell-knownsuchas Hildaites propose hereaprovisionalzonationbasedontwotypical Marconessa quarrysectionsforthelowerToarcian, we Standard Zonationisverydifficulttoapplyour of theBifronsZone.BecauseMediterraneanTethys first occurrenceofthegenus described speciesandsomenewones.We observedthe which aregenerathathererecordedwithalready fertia yielding thegenera nodular limestoneandgreymarloccur, withseveralbeds occurrence ofthegenus thickness occurs:herewehaveobservedthefirst anoxic event(OAE)acalcareouslayerofvariable In sections2and3,abovetheblackshalesofoceanic the fossiliferousbedswherespecimenswerefound. sections 2and3(Figs.2-4)anumberedarrowindicates two sectionsofMarconessaquarry. Inthelogsof angustisiphonatum sub-chronozones ( occurrence tothedisappearanceof chronozone, commonlycharacterizedfromthefirst 1994). We provisonallysubdivide ourBifrons Tethys MediterraneanStandardZonation(Elmi etal., be provisionallycomparedtotheLevisoniZoneof early Toarcian StriatusandUndicostachronozonescould Cingolites Calliphylloceras Mercaticeras Harpoceras

All specimensoftheSassaroli-Venturi collectionhere The ammonitesdescribedherewerecollectedfrom OFTHEMARCONESSA , , Phymatoceras Mesodactylites taxacollectedintheApenninessections,both , aswellseveral , , 06/09/10, 13.52 Praepolyplectus Leukadiella Hildaites undicosta QUARRY SECTIONS and , H. sublevisoni Lytoceras Taffertia , veryrare H. semipolitum , Hildaites Nodicoeloceras , Hildaites Frechiella Hildoceras , , Rakusites andthenewgenus Polyplectus . Abovethislayerthe , (Merla,1932).Both Dactylioceras H. lusitanicum Hildoceras and Hildaites crassus ) beingthisgenus Hildaites striatus Hildaites , atthebottom Phylloceras , Orthildaites , Martanites Rarenodia , , infour Neotaf- . Thus, , and 10-11-18 H. , , , , 10:56:23 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1A - Pag.: VOLUME.p0013.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1A - Pag.: VOLUME.p0005.pdf 10:56:23 10-11-18

10-11-18 10:56:23 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1A - Pag.: VOLUME.p0004.pdf Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:1APag.:VOLUME.p0008.pdf Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile 210x290mmStampatore:Moderna Data: 10/08/201008:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2-Segnatura: 1ABlack Job creato:Data:10-11-18 10:56:23 10-11-18 10:56:23

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YellowMagentaCyan 10-11-18 10:56:23 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1A - Pag.: VOLUME.p0016.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:1APag.:VOLUME.p0012.pdf 4 3 2 1 Device Resolution2400DPI 10:56:23 99 10-11-18 4x4 3x3 2x2 1x1 /0 /929 /749 /51/02/03/04/050 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 Neolioceratoides schopeni Zone by the “Tethyan Standard” (Elmi et Standard” Zone by the “Tethyan

06/09/10, 13.53 sp. indet. and and other ammonites, chiefly Hildoceratinae, in the early Polymorphum layers that the dark marly In fact, it seems al., 1994). of the on a calcareous bed the OAE lay which record where a ) Domerian (upper top of the About logs, Figs. 2, 4). surface occurs (see hardground a calcareous bed into the 20 cm under this hardground, has recorded the last occurrence of Corniola Formation Jaraphyllites which are forms commonly (Gemmellaro, 1886), in the uppermost Pliensbachian believed occurring Ferretti, 1975), although the latter (Cantaluppi, 1970;

Cingolites n.gen. from the Toarcian of the Apennines of the Toarcian from n.gen.

sub- Cingolites Cingolites is very rare O. douvillei Orthildaites douvillei Orthildaites S. Sassaroli, F. Venturi - Venturi F. S. Sassaroli, 99 10-11-18 In our Marconessa outcrop we observe a gap in the In our Marconessa Toarcian. Fig. 2 - Log of the Marconessa quarry section 2 (part 1) and ranges of chronozone, although the real although the real chronozone, sub-chronozone and an sub-chronozone or probably absent in the Appennines where, nevertheless, absent in the Appennines or probably we genus occur (which samples of this the oldest and still unnamed but unfortunately collected, Zone our Striatus cannot here subdivide unpublished). We still unpublished. being both index species to the underlying early Toarcian fossil record with regard Zone by the to the Tenuicostatum beds pre-OAE, ascribed Zonation (Page, 2003, 2004) or NW European Standard

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10-11-18 10:56:23 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1B - Pag.: VOLUME.p0003.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:1BPag.:VOLUME.p0011.pdf 10:56:23 10-11-18 06/09/10, 13.59

4 3 2 1 Device Resolution2400DPI Bollettino della Società Paleontologica Italiana, 49 (2), 2010 Italiana, Società Paleontologica della Bollettino 4x4 3x3 2x2 1x1 50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 and other ammonites in the Marconessa quarry section 2, part 1: from Levisoni Zone (Striatus and Undicosta and other ammonites in the Marconessa quarry section 2, part 1: from Levisoni 100 Cingolites

10-11-18

Tab. 1 - Ranges of Tab. zones) to lowermost Bifrons Zone. 100

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10-11-18 10:56:23 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1B - Pag.: VOLUME.p0015.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:1BPag.:VOLUME.p0007.pdf Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile 210x290mmStampatore:Moderna 10:56:23 10-11-18

10:56:23 YellowMagentaCyan 10-11-18 10:56:23 Job creato: Data: 10-11-18 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1B - Pag.: VOLUME.p0002.pdf

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4 3 2 1 Device Resolution2400DPI 4x4 3x3 2x2 1x1 50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 10-11-18 10:56:23 Data: 10/08/201008:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2-Segnatura: 1BBlack Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1B - Pag.: VOLUME.p0014.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:1BPag.:VOLUME.p0010.pdf 10:56:23 101 10-11-18 (Tabs. 1, 2 and 3), but 1, 2 and (Tabs.

06/09/10, 14.02 The stratigraphic occurrence of the ammonoid fauna occurrence The stratigraphic across both sections of the Marconessa quarry studied is quarry studied sections of the Marconessa across both tables displayed in the here broadly our and provisional being are incomplete these data detailed progress (a more and study still in sampling description of new collected taxa report with a formal

n.gen. from the Toarcian of the Apennines of the Toarcian from n.gen.

Cingolites Cingolites and other ammonites in the Marconessa quarry section 2, part 2: Bifrons Zone. S. Sassaroli, F. Venturi - Venturi F. S. Sassaroli, 101 Cingolites 2010). 10-11-18 Tab. 2 - Ranges of Tab. form is believed by few authors occurring also in the occurring also by few authors form is believed (see or Mirabile zones) (Polymorphum earliest Toarcian data review and new 2002). (For an updated Macchioni, the to OAE and across in relation on the ammonites the Apennines, see boundary in Pliensbachian-Toarcian Bilotta et al.,

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10:56:23 YellowMagentaCyan 10-11-18 10:56:23 Job creato: Data: 10-11-18 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1B - Pag.: VOLUME.p0002.pdf

Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:1APag.:VOLUME.p0008.pdf Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile 210x290mmStampatore:Moderna Data: 10/08/201008:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2-Segnatura: 1ABlack Job creato:Data:10-11-18 10:56:23 10-11-18 10:56:23

10-11-18 10:56:23 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1A - Pag.: VOLUME.p0001.pdf 1 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:1APag.:VOLUME.p0009.pdf 10:56:23 ». 10-11-18 Evolute Hyatt, 1867 Hyatt, 1867 Hyatt, Hyatt, 1867 ILDOCERATACEAE ILDOCERATINAE ILDOCERATIDAE 03/08/10, 13.42 Family H Subfamily H and other ammonites, chiefly Hildoceratinae, in the early Superfamily H According to Arkell et al. (1957, p. L259) the

Gèczy (1967, p. 127) has reviewed Schindewolf’s (1963) Gèczy (1967, p. 127) has reviewed Schindewolf’s Hildoceratinae are characterized by shells « planulates with quadrate whorl section, tricarinate- and falcate ribs which may be bisulcate venter, on whorl side interrupted by longitudinal groove Cingolites

Hyatt, 1889 Zittel, 1884 Bollettino della Società Paleontologica Italiana, 49 (2), 2010 Italiana, Società Paleontologica della Bollettino MMONITINA 102 MMONOIDEA YellowMagentaCyan Order A Suborder A SYSTEMATIC PALAEONTOLOGY SYSTEMATIC 10-11-18

will be provided by us in further papers). Furthermore, some meaningful ammonite taxa for biostratigraphy, chiefly the ranges of Hildoceratinae, Mercaticeratinae, are plotted against the logs (Figs. 2, 3 and 4). Fig. 3 - Log of the Marconessa quarry section 2 (part 2) and ranges of Toarcian. 102

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10-11-18 10:56:23 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 1A - Pag.: VOLUME.p0004.pdf Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:2APag.:VOLUME.p0024.pdf Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile 210x290mmStampatore:Moderna Data: 10/08/201008:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2-Segnatura: 2ABlack Job creato:Data:10-11-18 10:56:24 103 10-11-18 and (Blaison, Hildaitoides , characterized by Parahildaites Mercaticeras . We agree with these authors. We . . According to Guex (1974) the of the South America Palaeoprovince 06/09/10, 14.03 must be included in the subfamily 10:56:24 We are unable to carry out a critical evaluation about are We with tricarinate-bisulcate venter and strong, straight and spiny ribs, should be excluded from the Hildoceratinae (2000) According to Macchioni & Venturi subfamily. Renziceras Leukadiellinae Mediterranean genus small sized shapes, should be included in the subfamily Mercaticeratinae the taxonomic range of the genus Atacamiceras in our opinion, described by Hillebrandt (1987) show, only a seeming affinity with European Hildoceratinae. These American taxa, in fact, have rectiradiate, or gently 1967), found in the Arabian Palaeoprovince, because we have not enough data. The genera

and other ammonites, chiefly Hildoceratinae, in the early Toarcian. n.gen. from the Toarcian of the Apennines of the Toarcian from n.gen.

, . Cingolites Cingolites Cingolites Mercaticeras , Atacamiceras , , which is a shape Hildoceras , Renziceras Hildaitoides , S. Sassaroli, F. Venturi - Venturi F. S. Sassaroli, 103 Orthildaites , Parahildaites , diagnosis of the subfamily, which includes the diagnosis of the subfamily, Hildaites 10-11-18 We think that the genus We typical of the West Mediterranean Tethys characterized Tethys Mediterranean typical of the West by an evolute coiling and wider than high whorl section Renziceras suture line in diagnosis, and he wrote that this group is characterized by a «nombre réduit de lobes ombilicaux et par une selle externe sans incision médiane». Howarth (1992, p. 165) provided a broader diagnosis than above quoted authors, and for the first time he emphasizes the variability of the whorl section, which is «quadrate or elliptical» with «keeled or tricarinate-bisulcate venter», and of the ornament, which shows «Ribs vary from fine to strong, and from straight to falcate or strongly angled». Howarth includes into this subfamily eight genera: Fig. 4 - Log of the Marconessa quarry section 3 and ranges of (1963)

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4 3 2 1 Device Resolution2400DPI 4x4 3x3 2x2 1x1 50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 10-11-18 10:56:25 Data: 10/08/201008:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2-Segnatura: 2BBlack Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0030.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:2BPag.:VOLUME.p0026.pdf 10:56:25 10-11-18

10-11-18 10:56:25 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0031.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:2BPag.:VOLUME.p0023.pdf Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile 210x290mmStampatore:Moderna 10:56:25 10-11-18 - From very 06/09/10, 14.07 diagnosis of Hildoceratinae

Amended

evolute to moderately involute shells. Variable whorl evolute to moderately involute shells. Variable section, which may be sub-quadrate, low or high sub- rectangular and sub-trapezoidal, with wide or narrow which may be sub-rounded (mainly in the body venter, chamber) sub-tabulate, bisulcate and tricarinate-bisulcate. The keel is more or less raised, strong or fine. The ornament varies showing ribs from fine to coarse; they are usually falcate-falcoid or sigmoidal (sinuous- §flexuous) in the inner whorls, more or less retroverted near the umbilical edge and more or less projected on the ventro-lateral edge, and sometimes moderately falcate,

, and 10:56:25 Orthildaites , YellowMagentaCyan . Parahildaites Bollettino della Società Paleontologica Italiana, 49 (2), 2010 Italiana, Società Paleontologica della Bollettino Hildoceratinae. Therefore, Hildaites

Hildoceras and other ammonites in the Marconessa quarry section 3: from Levisoni Zone (Striatus and Undicosta zone) and other ammonites in the Marconessa quarry section 3: from Levisoni Zone (Striatus and 104 Cingolites do not have the typical traits described by (new genus) and

10-11-18

we suggest an amended diagnosis of this subfamily, which we suggest an amended diagnosis of this subfamily, includes only the genera that show a geographic dispersal in Subboreal, Submediterranean and Mediterranean Palaeoprovinces, such as Cingolites arched, and very thin ribs with rounded section. These have an evolute ovoidal-rounded whorl shells, moreover, section, and they have also a rounded or sub-acute venter, never bisulcate. Since the genera to lowermost Bifrons Zone. Tab. 3 - The ranges of Tab. Hildaitoides Arkell, Gèczy and Howarth, we think that they should be excluded from the subfamily 104 10:56:25

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Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:2BPag.:VOLUME.p0022.pdf 10:56:25 , . 105 10-11-18 has have ) occur . These , whilst stefaninii Cingolites ; in fact, it

Cingolites Cingolites C. Hildaitoides spiralis crassus Cingolites

Cingolites

shells. C. H. and n. sp. larger sized, and in Mercaticeras and - At the present state Mercaticeras is distinguished from , is very wide, swollen and , is very wide, swollen has typically straight and has typically Hildoceras 87, 259, 431, 432, 433, 802 , picenus

Atacamiceras Cingolites C. - Because of its typical strongly shows coarser ribs with a hildaitic , (Guex, 1973) in having a wide Cingolites . In fact, the former genus shows a . In fact, the former Orthildaites Orthildaites Cingolites clavatus has some degree of similarity with also differs from the taxa figured by also differs from the taxa figured 06/09/10, 14.13 in having a coarser ornamentation with in having a coarser ornamentation crassus

has a suture resembling both the latest clavatus - Eight well preserved specimens of medium (Figs. 5, 7, 11; Pl. 1, figs. 1-4) (Figs. 5, 7, 11;

H. C. is the most similar to and the earliest Orthildates Material Derivatio nominis Cingolites Nevertheless, Cingolites Palaeogeographic dispersal already described by Merla (1932), occurs in Southern Alta Brianza) and Greece Alps (Entratico, Alpe Turati, (Lefkas Island). and small size (MSA: 86 and 804), all from Marconessa quarry. clavate and coarser ribs. in the Apennines only. The fourth species in the Apennines only. Mercaticeras ribs and a more clavate, irregularly raised and not falcoid Besides, they evolute, swollen and wide whorl section. differ in size, being These American ammonoids, in fact, have a rounded or These American ammonoids, in fact, sulci, and straight sub-elliptical whorl section, without ribs, which are neither flexuous nor clavate. Hildaites crassus sulci and a strong tricarinate-bisulcate venter with deep E and L lobes. keel and a simple suture line, with short However, rursiradiate-falcate appearance and they are also not clavate. In any case, the earlier shapes of examination reveals remarkable differences, which differences, reveals remarkable examination this new Hildoceratinae establishment of justify the fact, genus. In faded in the proximal ribs, regularly raised, rectiradiate near the besides, they start distal segments; and in the they disappear completely near the umbilical edge and On the contrary, ventro-lateral edge. Hillebrandt (1987) as typically sinuous and projected ribs even in the outer typically sinuous and enlarged and swollen on the whorls, characteristically thus giving rise to the clavi and a ventro-lateral edges, area; besides, the ribs start from coronate ventral they have not a regularly raised umbilical wall and sub-quadrate whorls section, appearance. The differently from strongly tricarinate-bisulcate venter it always shows a suture distinguishes also with deep sulci. The taxon stratigraphic occurrence. The later stefaninii from very simple suture, similar to Orthildaites Hildaites was assigned by Merla (1932), Pinna (1963) and was assigned by Merla (1932), Pinna and typical Zanzucchi (1963) to this well-known Mediterranean genus. the same stratigraphic occurrence of likeness and affinity make plausible a common evolutionary descent. of knowledge, three of the four species of (namely,

n.gen. from the Toarcian of the Apennines of the Toarcian from n.gen.

, 3 ) is 3 n. sp., Haug, the ribs Cingolites Cingolites Hildoceras Orthildaites Bifrons Zone.

picenus the ribs usually

(Merla, 1932). douvillei

C. , n. sp. ). Usually, the ribs ). Usually, Cingolites O. lobe, a dentiform U 2 n. gen. ), one of which (U Hildaites 2 n. sp., stefaninii new genus includes the

Cingolites showing an almost dentiform From the location of the saddle. The sutures show also 1 C.

1 and U S. Sassaroli, F. Venturi - Venturi F. S. Sassaroli, - 3 Buckman, 1923, but a closer and Cingolites clavatus as well the ribs disappear before as well the ribs disappear

105 shells, mainly C. Cingolites clavatus Cingolites - orthus Genus n. sp. and

- Very evolute shell with a sub-quadrate - Very - - At a glance this taxon may be confused with O. Hildoceras Orthildaites spiralis Orthildaites

10-11-18 Among the genera of this subfamily only Among the genera ammonitic, i.e. usually The sutures are typically Remarks Stratus typicus - Nodular limestone and grey marl of The ornament shows spaced, coarse, swollen and The suture is simple, poorly jagged, with distanced Derivatio nominis Diagnosis Type species Type C. Material are single, but sometimes they are also bifurcating or are also bifurcating but sometimes they are single, edge fine tubercles near the umbilical bundled showing on the ending of the ribs and dimples. The or bullae also: in ventro-lateral edge varies edges; among come until the ventro-lateral and a few whereas, in the ventro-lateral edges; come until the ventro-lateral edge enlarged and swollen, come until the ventro-lateral venter. forming a typical coronate stage and in the grown-up stage, shows, both in the juvenile cuts the ribs, forming a typical more a spiral groove that smooth band. or less wide dorsal (jagged) and showing spaced simple, poorly indented but normally the lobes. The ratio of the E/L lobes varies, ES saddle is wider than LS the upper Undicosta Zone and the lowermost which is placed on the umbilical wall. The ES saddle is about twice as wide than LS some A lobe. 1884 and almost straight and rectiradiate in the grown-up stage in the grown-up and rectiradiate almost straight (mainly in usually small sometimes dentiform. The hidden internal usually small sometimes dentiform. The lobe (I) is simple, long and narrow. Marconessa quarry near Cingoli (Macerata, Italy), where Marconessa quarry near Cingoli (Macerata, this taxon was found. having whorl section and wide tricarinate-bisulcate venter, more or less deep sulci. The umbilical edge is rounded; the umbilical wall is more or less high, sloped or vertical. unevenly risen ribs, which are as wide as the interspace between ribs. These more or less coarse and flexuous ribs start from the umbilical wall, they are gently retroverted in the umbilical-proximal segment, stretched in the median segment and more or less projected on the ventro-lateral edges, where they reach the maximum growth forming a very swollen, enlarged and like a clavus area. Therefore, from the ventral view a typical coronate area can be observed. lobes; it shows a narrow E lobe, as long as the L lobe or only just longer than L; a short U two umbilical lobes (U following species:

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4 3 2 1 Device Resolution2400DPI 4x4 3x3 2x2 1x1 50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 10-11-18 10:56:25 Data: 10/08/201008:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2-Segnatura: 2BBlack Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0030.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:2BPag.:VOLUME.p0026.pdf 10:56:25 10-11-18

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10:56:25 YellowMagentaCyan 10-11-18 10:56:25 Job creato: Data: 10-11-18 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0018.pdf

Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:2APag.:VOLUME.p0024.pdf Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile 210x290mmStampatore:Moderna Data: 10/08/201008:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2-Segnatura: 2ABlack Job creato:Data:10-11-18 10:56:24 10-11-18 10:56:24

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YellowMagentaCyan 10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0032.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:2APag.:VOLUME.p0028.pdf 4 3 2 1 Device Resolution2400DPI 10:56:24 10-11-18 4x4 3x3 2x2 1x1 /0 /929 /749 /51/02/03/04/050 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100

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C. n. sp. Hildaites crassus from the celebrated S. Anna n. sp.: a) section (x 1) and b) suture and from - From the location of recording, - Nodular limestone and grey marl Undicosta Zone.

ex-situ and furthermore both from the various Cingolites picenus - 06/09/10, 14.16 - A well preserved specimen MSA FV1 of - Nine specimens of medium-large size Orthildaites Cingolites clavatus Cingolites Occurrence Material Derivatio nominis Holotype Stratus typicus (Figs. 6 a-f, i-l, n; Pl. 1, figs. 5-8; Pl. 2, figs. 1-5, 9)

from Marconessa quarry, some of which well preserved from Marconessa quarry, (MSA: FV1, 88, 91, 105, 207, 359, 434, 855 and 870). specimens of medium size (SAF 9 and SAF 18) Two were collected quarry of Furlo Pass (Pesaro). shapes of the Marchean Apennines, which long ago was inhabited by the ancient people named “Piceni”. ornament, which clearly distinguish other Fig. 5 - (x2) of holotype MSA 87; c) section (x1) and e) suture (x3) of (x2) of holotype MSA 87; c) section (x1) and (x4) of paratype paratype MSA 259; d) section (x1) and f) suture MSA 86. medium-large size with the body chamber (Figs. 6 a-b; Pl. 1, figs. 7a-b), collected from Marconessa quarry, section 2, bed 8b. Zone. of the upper Undicosta

lobe, 3 Cingolites Bollettino della Società Paleontologica Italiana, 49 (2), 2010 Italiana, Società Paleontologica della Bollettino - Very evolute shell - Very with an almost dentiform A 1 lobe and a dentiform U 2 106 - Nodular limestone and grey marl - Platycone very evolute shell, sub- are unevenly raised, more or less swollen; - The variously sized paratypes are very - A well preserved specimen MSA 87 of - for the strongly clavate appearance of the for the strongly clavate appearance . (Legend: D = whorl diameter; U = umbilical width; Wh = . (Legend: D = whorl diameter; 10-11-18 Paratypes Diagnosis Holotype description The flexuous, coarse and spaced ribs have a moderately The suture is simple, showing distanced lobes; an E Holotype Stratus typicus C. clavatus

which is placed on the umbilical wall. The ES saddle is about twice as wide than LS similar to the holotype, so that see the holotype diagnosis. They all show the typical characters of the lobe. clavatus breadth; ratio U/D; ratio Wb/Wh = Wb/ whorl height; Wb = whorl mm; they are usually taken at the maximum Wh. The measures are in shell size). Tab.4 - Measurements of the holotype and paratypes of - Measurements of the Tab.4 106 quadrate whorl section, wide tricarinate-bisulcate venter quadrate whorl section, wide tricarinate-bisulcate sparse ribs are with keel. The sinuous, coarse and The venter. characterized by clavi forming a coronate from other ammonitic suture is simple. It differs Cingolites ornament, for the coarser ribs and for the sub-quadrate ornament, for the coarser ribs and for whorl section wider than high. retroverted proximal segment, a long stretched medial segment and a short and gently projected distal segment. The ribs end enlarged and swollen on the ventro-lateral edges, here reaching the maximum thickness; thus, they take on a typical claviform appearance and forming a Besides, the ribs coronate area viewed from the venter. of lobe as long as L lobe, which is short and wide; a moderately developed U of the Undicosta Zone. with a sub- characterized by a slow growth of coiling, than high. The quadrate whorl section noticeably wider deep and wide wide tricarinate-bisulcate venter shows sulci, with a strong and moderately raised keel. The swollen sides show an almost vertical umbilical wall with rounded edges. they are 24 in number in the last whorl of the phragmocone. medium size with a portion of body chamber (Figs. 5 medium size with a portion of body Marconessa a-b; Pl. 1, figs. 1a-b), collected from 2, bed 6. section quarry,

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YellowMagentaCyan 10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0032.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0028.pdf 4 3 2 1 Device Resolution2400DPI 10:56:24 10-11-18 4x4 3x3 2x2 1x1 /0 /929 /749 /51/02/03/04/050 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100

10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0029.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0021.pdf 02.p65 10:56:24 claviform, but theclavidisappearonbody chamber, phragmocone theribtermination appearsgently segment ontheventro-lateral edges.Inthe segment andmoderatelyprojected withshortdistal segment, stretchedorgently archedinthemedial when theyappearslightlyretroverted intheproximal the innerwhorls,moreflexuous intheouterwhorls, almost verticalandhigherintheouterwhorls. wall ismoderatelyhighandroundedintheinnerwhorls, with deepsulciandmoderatelystrongkeel.Theumbilical quadrate whorlsection.Theventeristricarinate-bisulcate sinuous intheouterones.Theammoniticsutureissimple. rectiradiate intheinnerwhorlsandtheybecomemore strong, gentlyclavateandsparseribsarealmost whorl section,tricarinate-bisulcateventerwithkeel.The Fig. 6- 9; l)section(x1)andn)suture(x4)ofparatypeMSA359. 88; d)section(x1)andi)suture(x4)ofparatypeMSA6;e)j)105;k) The ribsarestrong,spaced,almostrectiradiatein Holotype description Diagnosis 10-11-18 Cingolites picenus -Platyconeevoluteshell,sub-quadrate 107 n.sp.:a)section(x1)andb)suture(x3)ofholotypeMSAFV1;c)f)paratype S. Sassaroli, F. Venturi - - Evoluteshellwithasub- Cingolites Cingolites

n.gen. from theToarcian oftheApennines gr. picenus: g)drawing(x1);h)section(x1)andm)suture(x3).

Fig. 7- Undicosta Zone; b) specimenMSA804(x1),section 2,bed.8b. Cingolites clavatus 06/09/10, 14.17 n.sp.:a)specimenMSA433(x1), from paratypeSAF

10-11-18 107

10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0020.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0022.pdf 02.p65 10:56:25 All photographsare naturalsizeprovided. Figs. 5-8 Figs. 1-4 holotype. variance intheLlobe,usuallyshorterthanthatof mainly intheouterwhorls.Thesuturealsoshowsa sometimes appearmorefalcoidthanintheholotype, the ornament,withmoreorlessstrongribs,which and ashortAlobe. longer thanElobe;moderatelydevelopedumbilicallobes 29 ribsinthelastwhorlofphragmocone. when theribsprogressivelybecomefaded.We count picenus andofspecimens Tab. 5-Measurementsofholotypeandparatypes 108 only. Itdiffers from and theappearanceofgentleclaviinphragmocone narrower venter;moreregularlyrisenandlesscoarseribs having anarroverandlessswollenwhorlsection; It finallydiffersfrom coarse ribs,alesswideventerandmoreflattenedsides. the lowermostBifronsZone. flexuous ribsintheouterwhorlstoo. tricarinate-bisulcate venterandlessswollenmore Occurrence Remarks Paratypes The sutureissimplewithaLlengthened,which 10-11-18 8 -paratypeMSA91,section2,bed 8c. 7 -holotypeMSAFV1,section2, bed8b. 6 -paratypeMSA105,collectedfrom detritusoftheuppermost 5 -paratypeMSA207,collectedform detritusoftheuppermost - Cingolitespicenus 4 -paratypeMSA431,section2,bed8c. 3 -paratypeMSA259,section2,bed8a. 2 -paratypeMSA86,section2,bed8b. 1 -holotypeMSA87,section2,bed6. - Cingolitesclavatus -Thisspeciesdiffersfrom -Theparatypesshowsomevariancesin -Fromuppermost C. 108

spiralis C. Cingolites

stefaninii n.sp. n.sp. inhavinglessarchedand Bollettino dellaSocietàPaleontologicaItaliana,49(2),2010 gr.

picenus inhavingawider Undicosta Zoneto C. .

clavatus EXPLANATION OFPLATE 1 Cingolites in

with serpenticoneappearance. respectively. Lecceti section(BossoValley, Pesaro):BU1andLE1 Maranghi quarrysection(BuranoValley, Pesaro)andfrom Marconessa quarry;twofragmentscollectedfrom clavatus and foritsundoubtedtransitionaltraitsbetween C. whorls, whichareunevenlyrisenandstronglyclavatelike specimen showsalmostrectiradiateribsintheinner stratigraphic occurrence,whichislowerthan raised andthinkeel. tricarinate-bisulcate, butlesswidethan wide inthebodychamber. The venteristypically whorl sectioninthephragmocone,onlyjusthigherthan Marconessa quarry. preserved specimenofthesamesizeMSA844,bothfrom 157 ofmediumsizewiththebodychamberandabadly chamber, like become progressivelyfalcoidandfadedinthebody regularly risenandmorecoarseflexuousribs,which including thebodychamber, theornamentshowsamore U is wideandshort;adentiformAlobelongnarrow ornament. collected fromdetritus

2

Undicosta Undicosta lobe. Occurrence The specimenMSA157isveryinterestingforits Remarks Material The sutureshowstheElobeaslongL,which Derivatio nominis Material clavatus and (Figs. 6g-h,m;Pl.2,figs.6-7) zone(section3). zone(section2). -OnewellpreservedspecimenMSA21from -Very evoluteshellwithasub-quadrate -AverywellpreservedspecimenMSA ; whereas,intheouterwhorl,whichis (Fig. 8;Pl.2,figs.8,10-11) C. 06/09/10, 14.20 C. -Upper Cingolites spiralis

Cingolites picenus

picenus

- Foritstypicalevolutecoiling specimen MSA884hasthesame

Undicosta Zone. . Infact,theornamentofthis and gr.

they losetheclavi.The picenus n.sp. C.

clavatus C.

picenus , with 10-11-18 C. , 10:56:25 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0019.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0027.pdf 10:56:25 10-11-18

4 3 2 1 Device Resolution2400DPI 4x4 3x3 2x2 1x1 50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 10-11-18 10:56:25 Data: 10/08/201008:32:49-Lavoro:MD17220-Bollettino_Paleontologia_Vol_49_N_2Segnatura:2BBlack Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0030.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0026.pdf 10:56:25 10-11-18

10-11-18 10:56:25 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0031.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0023.pdf Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile210x290mmStampatore:Moderna 10:56:25 10-11-18

10:56:25 YellowMagentaCyan 10-11-18 10:56:25 Job creato: Data: 10-11-18 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0018.pdf

Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0022.pdf 10:56:25 10-11-18

10-11-18 10:56:25 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0019.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0027.pdf 10:56:25 10-11-18

4 3 2 1 Device Resolution2400DPI 4x4 3x3 2x2 1x1 50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 10-11-18 10:56:25 Data: 10/08/201008:32:49-Lavoro:MD17220-Bollettino_Paleontologia_Vol_49_N_2Segnatura:2BBlack Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0030.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0026.pdf 10:56:25 10-11-18

10-11-18 10:56:25 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0031.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0023.pdf 02.p65 Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile210x290mmStampatore:Moderna 10:56:25 10-11-18 109 S. Sassaroli, F. Venturi - Cingolites

n.gen. from theToarcian oftheApennines

10:56:25 03/08/10, 13.45 YellowMagentaCyan Pl. 1 10-11-18 109

10:56:25

Job creato: Data: 10-11-18 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0018.pdf

Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0024.pdf 02.p65 Formato foglio:640x880 STAMPA:B+VPiegatura:16moconvertibile 210x290mmStampatore:Moderna Data: 10/08/2010 08:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2 -Segnatura:2A Black Job creato:Data: 10-11-18 10:56:24 compressed shell, sub-quadratewhorl section, of theupperUndicostaZone. Pl. 2,figs.8a-b),coming from Marconessaquarry. the detritusofupperUndicostaZone(Figs.8a-c; 110 (x4) ofparatypeBU1. (x1) ofparatypeLE1;e)drawing(x1),g)sectionandh)suture and c)suture(x3)ofholotypeMSA21;d)f)section Fig. 8- Diagnosis Stratus typicus Holotype 10-11-18 Cingolites spiralis -SpecimenMSA21collected -Very evolutealmostserpenticone -Nodularlimestoneandgrey marl 110 n.sp.:a)drawing(x1),b)section(x1) Bollettino dellaSocietàPaleontologicaItaliana,49(2),2010 ex situ 10:56:24 from

and smallsize (MSA: 264,281,388,435,843, 845,847, similar to some traitsoftheribsappearanceandsutureare about twiceaswidethanLS 1930 ? the umbilicallobeswithadentiformU lobe; anElobeaslongL,alittledevelopedAlobe,and count 23ribsinthelastwhorlofphragmocone. lateral edgesandformingacoronateventralarea.We a gentlyclaviformappearance,endingontheventro moderately projecteddistalsegment,wheretheyhave appearance; theyareswollenandenlargednearthe medial segments,characterizedbyanalmostprorsiradiate from theumbilicalwall,withstretchedproximaland are broaderthantheinterspacebetweenribs.Theribsstart low umbilicalwallisroundedtoo. and moderatelydeepsulci.Thesidesareroundedthe coiling. Thewideventeristricarinate-bisulcatewith which iswiderthanhigh,characterizedbyaslowgrowth serpenticone appearanceandsub-quadratewhorlsection, is verysimple. concave-prorsiradiate appearance.Theammoniticsuture clavate, verysparceribsarecharacterizedbyanalmost tricarinate-bisulcate venterwithkeel.Thecoarse,gently spiralis Tab. 6-Measurementsoftheholotypeandparatypes close affinitywith and thestronglytricarinate-bisulcateventerrevealits evolute appearance.Besides,theclaviformcoarserribs Hildaites lobe islongerthanLlobe. and venter;theydiffer inthesuturelineonly, wheretheE similar totheholotypeinornament,whorlsection

1932 1963 1963 1995 Occurrence The sutureisverysimple,showingashortandwideL The ornamentshowscoarseandspacedribs,which Holotype description Material Remarks Paratypes . - M Hildoceras (Lillia)Mercati Mercaticeras stefaninii Mercaticeras stefaninii Mercaticeras stefaninii Praemercaticeras sp.indet.P shellsforitstypicalstronglyserpenticone Cingolites stefaninii ITZOPOULOS Hildaites crassus -Ninewellpreservedspecimens ofmedium -Thisspeciesdiffersfrom (Fig. 9;Pl.3,figs.1-4,6,8-9) -Thetwofragmentedparatypesarevery 06/09/10, 14.22 -UpperUndicostaZone. , Pl.6,figs.6a-b. Cingolites clavatus -Very evoluteshellwith Merla- Z Merla-P M 1 . saddle. ERLA (Merla,1932) (Hauer) var. ETTINELLI , Tav. 6,figs. 8-9. ANZUCCHI INNA 3. ThesaddleESis , Tav. 10,fig. 14. Orthildaites etal.,Pl.2,fig.5. . Furthermore, ellenica , Tav. 19,fig.1. Cingolites (Renz) and 10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0017.pdf 2 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0025.pdf 10:56:24 10-11-18

YellowMagentaCyan 10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0032.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0028.pdf 4 3 2 1 Device Resolution2400DPI 10:56:24 10-11-18 4x4 3x3 2x2 1x1 /0 /929 /749 /51/02/03/04/050 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100

10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0029.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0021.pdf 10:56:24 10-11-18

10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0020.pdf Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0024.pdf Formato foglio:640x880 STAMPA:B+VPiegatura:16moconvertibile 210x290mmStampatore:Moderna Data: 10/08/2010 08:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2 -Segnatura:2A Black Job creato:Data: 10-11-18 10:56:24 10-11-18 10:56:24

10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0017.pdf 2 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0025.pdf 02.p65 10:56:24 quarry specimensmustbeascribed. which boththeMerla’s specimenandtheMarconessa section andtheornamentaretypicalof Mercaticeras species 485), allfromMarconessaquarry. 846, 886andtwofragmentedspecimens(MSA:411 and (x4) ofthespecimenMSA388. e) section(x1)andh)suture(x3)ofspecimenMSA886;g) MSA 485;d)section(x1)andf)suture(x3)ofspecimen281; suture (x3)ofspecimenMSA264;c)section(x1) Fig. 9- the mediumsized specimens. disappear onthebodychamber andintheouterwhorlof which isvisibleamongsmall specimensonly. The clavi edges, hereformingacoronate areaintheinnerwhorl, umbilical wallandtheyendclavate ontheventro-lateral of mediumsizedspecimen. Theribsstartfromthe rectiradiate; theyaregently flexuous intheouterwhorl is lowwithroundededge. sides areflattenedorbarelyrounded.Theumbilicalwall The sulciarenotmuchwideandmoderatelydeep. quadrate whorlsectionfromtricarinate-bisulcateventer. Amended diagnosis Remarks The strongandspacedribsarealmoststraight 10-11-18 Cingolites stefaninii stefaninii -ThesespecimensareveryclosetoMerla’s . Nevertheless,thecharactersofwhorl , whichwasascribedtothegenus 111 -Very evolute shellwithasub- S. Sassaroli, F. Venturi - (Merla, 1932):a)section(x1)andb) Cingolites Cingolites , to

n.gen. from theToarcian oftheApennines

Tab. 7-Measurementsofthespecimens of 280. sp. indet.3:f)suture(x6)andg)section(x1)ofthespecimenMSA section (x1)ande)suture(x3)ofspecimenMSA155. and c)suture(x3)ofspecimenMSA150. Fig. 10- Cingolites 03/08/10, 13.46 sp. indet.1:a)section(x1),b)drawing(x1) Cingolites Cingolites stefaninii YellowMagentaCyan sp. indet.2:d)

Cingolites

10-11-18 111

. 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0032.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0028.pdf 4 3 2 1 Device Resolution2400DPI 10:56:24 10-11-18 4x4 3x3 2x2 1x1 /0 /929 /749 /51/02/03/04/050 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100

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10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0020.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0022.pdf 10:56:25 10-11-18

10-11-18 10:56:25 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0019.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0027.pdf 10:56:25 10-11-18

4 3 2 1 Device Resolution2400DPI 4x4 3x3 2x2 1x1 50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 10-11-18 10:56:25 Data: 10/08/201008:32:49-Lavoro:MD17220-Bollettino_Paleontologia_Vol_49_N_2Segnatura:2BBlack Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0030.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0026.pdf 02.p65 10:56:25 moderately developed. is lowandwide;theumbilicallobesAlobe simple sutureshowstheElobeaslongLlobe,which the ventro-lateraledges,mainlyinouterwhorls.The strong andalmostrectiradiateribsshowgentleclavion section andtricarinate-bisulcateventer. Themoderately the bodychamber, comingfromMarconessaquarry. Subzone). collected prorsiradiate. Unfortunately, thisspecimenwasnot outer whorltheribsbecomefiner, archedandalmost characters ofthisspeciesintheinnerwhorls,but larger specimenMSA886showsthemaindiagnostic evolute whorlsectionandmoreorlessstrongribs.The column, showsomevariancesinhavingamoreorless specimens of Hildaites pseudolevisoni inner whorlsaremoresimilartothelaterspecimensof the whorlsectionappearanceandribscourseof affinity with of theornamentandsuturelinerevealacertain never formingarealcoronateventralarea.Sometraits evolution, fineranddenserribs,whicharegentlyclavate lobes withamoreorlessdevelopedlanciformU lobe, theAlobemoderatelydevelopedandumbilical 112 dentiform U Figs. 1-5,9 transitional shape. All photographsare naturalsizeprovided. Figs. 6-7 Figs. 8,10-11 It differsfrom Remarks Material Occurrence Our specimens,variouslylocatedinthestratigraphic The simplesutureshowstheElobeaslongL Occurrence 10-11-18 in situ -Evoluteshellwithasub-quadratewhorl -OnespecimenMSA150withaportionof 3 -paratypeMSA870,collectedfromdetritusoftheuppermost 2 -paratypeMSA88,section2,bed8b. 1 Cingolitespicenus - -paratypeMSA359,collectedfromdetritusoftheuppermost 3 Cingolites - -paratypeSAF18,collected 9 -paratypeSAF9,collected 5 -paratypeMSA855,collectedfromdetritusoftheuppermost 4 1-paratypeLE1,collected 11 0-paratype BU 1,collected 10 -holotypeMSA21,collected fromdetritusoftheuppermost 8 Cingolites spiralis - -specimenMSA844,collectedfromdetritusof theuppermost 7 -specimen MSA157,section3,bed6e. 6 , whichisplacedontheumbilicalwall. Cingolites Hildaites crassus (Figs. 10a-c;Pl.3,fig.5) -LowerUndicostaZone. - FromlowerBifronsZone(Sublevisoni . Cingolites C. 112

clavatus . Ontheotherhand,coiling, gr. (Venturi, 1981)andtheearlier picenus sp. indet.1 byhavingalessershell Bollettino dellaSocietàPaleontologicaItaliana,49(2),2010 . Therefore,itisaclearly n.sp. n.sp. ex situ ex situ ex situ ex situ fromdetritusoftheuppermost fromdetritusoftheuppermost . . EXPLANATION OFPLATE 2 2 anda

and alonglobeL. is simple,poorlyjagged,withAlobenotmuchdeveloped more straightandstrongintheouterwhorls.Thesuture bisulcate venter. Theribsareflexuousintheinnerwhorls; section, onlyjusthigherthanwide,andabroadtricarinate- chamber, MSA155,comingfromMarconessaquarry. Tab. 8-Measurementsofthespecimensundetermined short Llobe. not reallyclavate.Thesutureissimplewithawideand wall, areweaklyprojectedontheventro-lateraledges, section. Therectiradiateribs,startingfromtheumbilical quarry. the bodychamber, MSA280,comingfromMarconessa is anothertransitionalshape. Thus, itisforsometraitssimilarto more rectiradiatenotclavateribsintheouterwhorls.

It differsfrom Remarks Remarks Occurrence Material Material Undicosta Undicosta Undicosta Undicosta Undicosta Undicosta Undicosta -Evoluteshellwithasub-quadratewhorl -Evoluteshellwithasub-quadratewhorl -Amedium-large specimenwiththebody -Afragmentofthephragmoconeand zone(section3). 06/09/10, 14.24 (Figs. 10d-e;Pl.3,fig.7) zone. zone(section2). zone(section3). -LowerUndicostaZone. zone(section2). Cingolites Cingolites zone. zone. C.

(Figs. 10f-g) clavatus sp. indet.3 sp. indet.2 and C. Hildaites crassus

spiralis inhaving Cingolites 10-11-18 . It . 10:56:25 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0031.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0023.pdf Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile210x290mmStampatore:Moderna 10:56:25 10-11-18

10:56:25 YellowMagentaCyan 10-11-18 10:56:25 Job creato: Data: 10-11-18 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0018.pdf

Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0022.pdf 10:56:25 10-11-18

10-11-18 10:56:25 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0019.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0027.pdf 02.p65 10:56:25 10-11-18 113 S. Sassaroli, F. Venturi - Cingolites

n.gen. from theToarcian oftheApennines

03/08/10, 13.48 4 3 2 1 Device Resolution2400DPI 4x4 3x3 2x2 1x1 50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100

Pl. 2

10-11-18 113

10:56:25 Data: 10/08/201008:32:49-Lavoro:MD17220-Bollettino_Paleontologia_Vol_49_N_2Segnatura:2BBlack Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0030.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0026.pdf 10:56:25 10-11-18

10-11-18 10:56:25 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0031.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0023.pdf Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile210x290mmStampatore:Moderna 10:56:25 10-11-18

10:56:25 YellowMagentaCyan 10-11-18 10:56:25 Job creato: Data: 10-11-18 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0018.pdf

Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0024.pdf Formato foglio:640x880 STAMPA:B+VPiegatura:16moconvertibile 210x290mmStampatore:Moderna Data: 10/08/2010 08:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2 -Segnatura:2A Black Job creato:Data: 10-11-18 10:56:24 10-11-18 10:56:24

10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0017.pdf 2 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0025.pdf 10:56:24 10-11-18

YellowMagentaCyan 10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0032.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0028.pdf 02.p65 4 3 2 1 Device Resolution2400DPI 10:56:24 All photographsare naturalsizeprovided. Fig. 7 Fig. 5 above treatedfossilrecordevidences. questions intotheevolutionaryframeworkshownby study. However, weproposeheresomeprovisional the lowerToarcian Hildoceratinaewouldneedaseparate Subzone). indet. 1and2foritsstratigraphicoccurrence. 114 Figs. 1-4, 6, 8-9 - - Figs. 1-4,6,8-9 is missing. the OAErecordinginfigs.2and4)whenfossilrecord surface infigs.2and4)theriseofsealevel(see occurred inatimespanbetweenthefall(seehardground were maybelocatedintheApenninesorelsewhere,clearly evolutionary changesfromthecommonancestors,which well differentiatedinmanyspeciesandvarieties.These in oursectionsabovetheOAEblackshalesarealready of thefossilrecord.InfactHildoceratinaeoccurring Eldredge &Gould(1972),butclearlytotheincompleteness of Hildoceratinaeisnotduetoa“punctuation” seems tobesudden.However, thissuddenappearance theappearanceofHildoceratinae Zone (ourStriatusZone) preserved anyammonites,thusinthelowermostLevisoni black shalesofMarconessaquarrysectionshavenot Pliensbachian)-Toarcian transition.Forinstance,theOAE usually poorormissingacrosstheDomerian(late collections oftheEuropeanandMediterraneansections by thefossilrecord,beingunfortunatelyammonoid likely, althoughtherearenodirectevidencesprovided from thelateDomerianProtogrammoceratinaeisvery (see Bilottaetal.,2010).ThedescentofHildoceratinae previous anoxicbiologicalcrisisoftheearlyToarcian appear toreplacethetaxathathadbecomeextinctin fauna whennewgeneraandspeciesofHildoceratinae observe aremarkableadaptiveradiationoftheammonoid after theoccurrenceofblackshalesOAE,we Hildaites unfortunately stillunpublishedtaxaofthegenus striatus In thefossilrecordofstudied A moredetailedaccountoftheevolutionarytrends Occurrence It noticeablydiffersfromtheother In fact,wehavefoundinthelowerToarcian ( 10-11-18 zone)oftheMarconessaquarryseveralnewand 4x4 3x3 2x2 1x1 : someshapesaremoderatelyinvolute,withflat THE DESCENTOF /0 /929 /749 /51/02/03/04/050 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 - Cingolites - Cingolites 9 -specimenMSA435,collected from 8 -specimenMSA843,section2, bed10g. 6 -specimenMSA485,section2,bed10a. 4 -specimenMSA886,collectedfromdetritusofthelowermost BifronsZone(section2). 3 -specimenMSA845,collected 2 -specimenMSA388,collected 1 -specimenMSA264,section2,bed10a. Cingolites stefaninii -LowerBifronsZone(Sublevisoni 114 sp. indet.2,specimenMSA155,section 3,bed6b. sp. indet.1,specimenMSA150,section 3,bed6c. Bollettino dellaSocietàPaleontologicaItaliana,49(2),2010 CINGOLITES (Merla,1932)

Apennines sections, Cingolites ex situ ex situ EXPLANATION OFPLATE 3 Hildaites detritus sensu fromdetritusoflowermostBifronsZone(section2). fromdetritusoftheuppermostUndicostaZone(section3). sp.

ofthelowermostBifronsZone (section 2). Marchean basinhasbeenobserved.To thepresent-day nodular limestoneoccurringintheTuscan-Umbrian- Cingolites transitional andundeterminedshellsreferableto Merla (1932).We havefurthermorecollectedsome the samepelagicdepositional (Lefkas Island)(Pettinellietal.,1995,Pl.2,fig.5),where 19, fig.1;Pinna,1963,Tav. 10,fig.14)andGreece Apennines, theSouthernAlps(Zanzucchi,1963,Tav. was previouslyassignedtothegenus forthcoming paper. framework forthissubfamilywillbeattemptedbyusina our collection.Anewandexhaustivephylogenetic descent, asshownbyseveraltransitionalspecimensof that allHildoceratinaetaxaarelinkedbyacommon Apennines outcrops,wehopewillbeabletoprove Considering thegrowingbiostratigraphicdatafrom from thesmallsizedshapesof genus appears. Finally, intheearliest Bifronschronthegenus those ofthegroup bisulcate venterandcoarserribsintheornament,like from agroupof Cingolites late portionoftheUndicostachron,whenfirstly remarkable evolutionaryeventoccurredinthemiddle- when thegenera former gentleammonitesinthe in thestratigraphicsequence,andtheyreplacetotally strong ribs.Theselastrobustonesbecomemoreabundant instead evolute,withbisulcateortricarinateventerand and narrowventerfineribswhereasotherformsare recognized speciesas spiralis of Hildoceratinae. We havealsoestablishedthreenewspecies Cingolites Marconessa quarryallowedustoestablishthenewgenus PALAEOGEOGRAPHIC DISPERSALOF

Cingolites The richnessofourcollectioncomingfrom IN THEMEDITERRANEANTETHYSREALM Mercaticeras REMARKS ANDDISCUSSIONONTHE

) andweascribedtothisgenusanalready It ispossiblethatatthebottomofthiszone andlater . , whichweincludeinthesubfamily Cingolites (namely, 06/09/10, 14.31 Orthildaites Hildaites originatedfrom Orthildaites C. Hildaites crassus C.

stefaninii stefaninii

characterizedbytricarinate- clavatus and facies Hildaites undicosta Cingolites originated,probably (Merla,1932),which C. stefaninii ofredorgreymarly isreportedinthe , Cingolites C. Mercaticeras

picenus (Guex,1973). CINGOLITES appear. Hildoceras , probably and group. zone,

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10-11-18 10:56:24 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2A - Pag.: VOLUME.p0020.pdf Data: 10/08/2010 08:32:49 - Lavoro: MD17220 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:2APag.:VOLUME.p0024.pdf Formato foglio:640x880STAMPA:B+VPiegatura:16moconvertibile 210x290mmStampatore:Moderna Data: 10/08/201008:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2-Segnatura: 2ABlack Job creato:Data:10-11-18 10:56:24 10-11-18 10:56:24

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n.gen. from the Toarcian of the Apennines of the Toarcian from n.gen.

Cingolites Cingolites S. Sassaroli, F. Venturi - Venturi F. S. Sassaroli, 115 10-11-18

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10-11-18 10:56:25 Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0019.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:2BPag.:VOLUME.p0027.pdf 10:56:25 , . C. . 10-11-18 and Hildaites Cingolites Mercaticeras (collected from picenus Mercaticeras

and C. , stefaninii We collected also several We clavatus

Hildoceras seems to have its descent from , C. of the Undicosta Zone characterized CONCLUSIONS Cingolites. ACKNOWLEDGMENTS Cingolites 06/09/10, 14.34 Orthildaites , plus three new species which we ascribed Hildaites , being this new genus confined chiefly to the (all collected from the beds of the Undicosta (all collected from the beds of the The lower Toarcian Apennines outcrops preserved The lower Toarcian The genus Many of these taxa show a clear provincialism like We thank Prof. Maria Paola Maceratini, Prof. Johannes S. thank Prof. Maria Paola We Rosen’s (1992) warning on some constraints of the (1992) warning on some constraints Rosen’s

a remarkable early Hildoceratinae fossil record in the a remarkable early Hildoceratinae fossil (Macerata, layers of the Marconessa quarry near Cingoli of Italy), where we collected several species spiralis the first time in Zone), which have been described for this paper and a later species C. was previously the beds of the Bifrons Zone), which assigned by Merla (1932) to the genus Cingolites Apennines or anyway to the Mediterranean Tethys (Southern Alps and Lefkas Island, Greece). Generally speaking, several ancient Hildoceratinae seem to have an as shown early occurrence in the Mediterranean Tethys, by the recording of our Apennines sections. However a fauna more detailed account for this early Toarcian sampled by us in the Apennines outcrop of the Marconessa quarry will be provided in further papers. the ecology, and hence endemism is limited to the species endemism is limited and hence the ecology, barriers, not only physical level. Nevertheless or genus fauna factors obstruct the and climatic but also ecological on and eustasy affect and both plate tectonics dispersal a long distinction between Thus, a sharp these factors. scale (i.e. between the vicariance and a short temporal in palaeobiogeography)and the historical issues seems possible. The role of the centres of to be not practically therefore is not yet ruled out. origin and spreading data, such as those related to the palaeobiogeographical fossil record or the stratigraphical incompleteness of the mistake in taxonomic judgements, is correlation and the that the short biostratigraphic and useful. It is possible palaeobiogeographic account provided in this paper could incompleteness of the fossil record. be conditioned by the that there are some evidences Nevertheless, we believe between for the claim of provincialism and endemism Hildoceratinae the European and Mediterranean ammonoid faunas of the early Toarcian. Orthildaites to the new genus attributable to undetermined and transitional specimens the present-day knowledge these genera. To includes four species: a group of Pignatti, Dr. Massimiliano Bilotta and Prof. Enrico Serpagli for kindly Pignatti, Dr. revising the manuscript and for their useful suggestions, which are also grateful to Prof. Mikhail We helped us to improve this paper. Rogov and Prof. Louis Rulleau, who gently reviewed this paper, for very useful critical comments. by evolute shells with tricarinate-bisulcate venter and coarser ribs; the same group has probably originated the later genera

C. seems of these status 4 3 2 1 stefaninii Device Resolution2400DPI

C. , seem to be endemic , seem to herein designed, i.e. herein designed, Bollettino della Società Paleontologica Italiana, 49 (2), 2010 Italiana, Società Paleontologica della Bollettino 4x4 3x3 2x2 1x1 spiralis

50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 C. zone. n. sp.: a-b-c) specimen MSA 802 (x1), Cingolites 116 and . Undicosta picenus

C. , a priori

Cingolites clavatus

10-11-18

Page (1996, 2003, 2004, 2008) has recently provided Page (1996, 2003, 2004, Westermann (2000a, 2000b; see also Cecca, 2002) (2000a, 2000b; Westermann At species and genus level, our data related to the At species and genus

an useful synthesis of the current palaeogeographical adopting simple knowledge on ammonoids, Subrealm and hierarchical biogeographic units (Realm, Realm, the Province) and distinguishing two European During the late Pliensbachian- Boreal and the Tethyan. includes a Subboreal the Boreal Realm early Toarcian and in the Province (northern Britain, Russian Platform) Realm includes a West same substages the Tethyan Subrealm, which was divided in several Tethyan Province Tethyan) Provinces: a Mediterranean (or West Bulgaria, North Austria, Hungary, (southern Spain, Italy, (northern Spain, Africa); a Submediterranean Province Germany, southern and central France, southern Arabo-Malgach) Switzerland) and an Ethiopian (or Madagascar). Province (Ethiopia, Arabia, Turkey, and the same the boundaries Obviously, of the Apennines. On the other hand, On the other of the Apennines. the other species of the other units provided an updated framework of biogeographical defined by in the light of the new concept of biochore, a geographic the overall endemism of its biota within temporal limit area (chorotype) and within a any definition has its problems. (chronotype). However, For instance, the choice of the quality and quantity of taxa to define a biogeographic unit is somehow subjective. to solve the question leads him to (2002) effort Cecca’s establish a difference between a long temporal scale, when provincialism occurred in relation to vicariance processes, and a short temporal scale (i.e. ammonite biozones), when different taxa occurred in relation to clavatus being affected Provinces are changing in space and time, ecology and climate change. by plate tectonics, eustasy, 116 Fig. 11 - Fig. 11 microconch from stratigraphic occurrence and geographical dispersal of stratigraphic occurrence therefore seem to confirm the the early Hildoceratinae ammonoid faunal realms and existence of different and the Mediterranean areas provinces in the European Furthermore, our data about during the early Toarcian. of these taxa allow to conjecture the endemic occurrence for the ammonite faunas a certain degree of provincialism For these reasons the concepts of of the Apennines. spreading therefore cannot be centre of origin and rejected to have only a Western Mediterranean Tethys dispersal Tethys Mediterranean a Western to have only Greece). Southern Alps, (Apennines,

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4 3 2 1 Device Resolution2400DPI 4x4 3x3 2x2 1x1 50 40/60 30/70 20/80 10/90 5/95 4/96 3/97 2/98 1/99 0/100 10-11-18 10:56:25 Data: 10/08/201008:32:49-Lavoro: MD17220-Bollettino_Paleontologia_Vol_49_N_2-Segnatura: 2BBlack Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2 - Seg.: 2B - Pag.: VOLUME.p0030.pdf Info: MD17220_Bollettino_Paleontologia_Vol_49_N_2-Seg.:2BPag.:VOLUME.p0026.pdf 10:56:25 , . In 117 10-11-18 28 (1): 17: 145-

, 119: 35- , 119: , 92: 171- , Bollettino Bollettino Broidieia e (Maubeuge, Quaderni del Phymatoceras , Bollettino della , 6: 123-124. Palaeontographia Neolioceratoides 33: 4-60. Bollettino del Servizio Rivitsa Italiana di Renz (1919). in Space and Time. in Space and Time. Pseudolillia , 44 (3): 231-236. , 39 (3): 319-339. Hildoceras , 13 (2): 67-98. , nuovo genere di Brachiopodi Pragmateiai tes Akademias 31 (2): 207-222. Pseodomercaticeras , , 110 (1): 109-114. , 110 Leukadiella Geological Survey of Denmark and Monograph of the Paleontographical Monograph del Lias inferiore di Cingoli (Marche, Memorie della Società Italiana Scienze Comunicaçoes dos Seviços Gèologicos Comunicaçoes dos Seviços , 6: 287-432. . Ineson J. & Surlyk F. (eds.). The Jurassic Ineson J. & Surlyk F. , 1: 23-59. , 114: 97-158. , 114: In Part I: 1-106 (1992) - Part II: 107-200 (1993). (1992) - Part II: 107-200 Part I: 1-106

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, 163: 49-68. Palaeogeography, , 158: 1-13. Bollettino della Società Paleontologica Italiana, 49 (2), 2010 Italiana, Società Paleontologica della Bollettino 118 Farinacci A. & Elmi S. (eds.). Rosso Ammonitico & Elmi S. (eds.). Rosso Farinacci A. In YellowMagentaCyan 10-11-18 Consequences sur la stratigraphie et la taxonomie des et la taxonomie sur la stratigraphie Consequences Ammonitina. Roma, pp. 581- Proceedings, Edizioni Tecnoscienza, Symposium 602. for the new guidelines review and revision under Mesozoic: and nomenclature. biogeographic classification Palaeoclimatology, Palaeoecology Palaeoclimatology, nomenclature in Paleobiogegraphy: an attempt at order. nomenclature in Paleobiogegraphy: Palaeoecology Palaeoclimatology, Palaeogeography,

Westermann G.E.G. (2000a). The marine faunal Realms of the (2000a). The marine G.E.G. Westermann 118 Westermann G.E.G. (2000b). Biochore classification and G.E.G. (2000b). Westermann

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