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Abteilung A: Palaozoologie - Stratigraphie

Volume 290

Theammonite genus Vascoceras CHOFFAT, 1898 (family Vascoceratidae DouviLLE, 1912) in the Iberian Trough, Spain

by

FERNANDO BARROSO-BARCENILLA and ANTONIO GoY, Madrid

With 5 plates and 6 text-figures

E. Schweizerbart'sche Verlagsbuchhandlung

(Nagele u. Obermiller) • Stuttgart 2010 Palaeontographica Abt. A: Palaeozoology- Stratigraphy Vol. 290, Issue 4-6: 199-235, Marz 2010 ©E.Schweizerbart'sche Verlagsbuchhandlung 2010

The ammonite genus Vascoceras CHOFFAT, 1898 (family Vascoceratidae DouviLLE, 1912) in the Iberian Trough, Spain

by

FERNANDO BARROSO-BARCENILLA and ANTONIO GOY

with 5 plates and 6 text-figures

Zusammenfa ssung Die zu der Gattung Jlascoceraszugeschriebenen Ammoniten der Sammlungen von WrEDMANN (Universitat Ttibingen, Deutsch­ land) sowie GoY, CARRETERO und MELENDEZ (Universidad Complutense de Madrid, Spanien), die aus dem Ober Cenoman und dem Unter Turon des Iberischen Trogs stammen, sind revidiert worden. AnschlieBend neue Vertreter der Arten Jlascocerasgamai, V.charoni sp.nov., V.barcoicense, V. durandi, V. cauvini, V. amieirense, V. harttii und V. kossmati sind studiert und prasentiert worden. Untersuchungen iiber die Morphologic sowie die geographische und stratigraphische Verteilung dieser Ammoniten haben gefiihrt schlieBlich zu der Feststellung verschiedener phylogenetischer Beziehungen zwischen ihnen und zu der Unterscheidung von zwei Hauptphasen in der Entwicklung der Familie Vascoceratidae die durch die aufeinanderfolgende Dominanz von "primitiven" und von "entwickelten" Jlascoceras gekennzeichnet sind. Schliisselworter: ober Cenoman-unter Turon- Ammonoidea-Vascoceratidae-Vascoceras- neue Art- Iberischer Trog-Spanien

Summary The ammonites assigned to the genus Vtzscoceras of the WIEDMANN (Universirat Tiibingen , Germany) and GoY, CARRETERO and MELENDEZ (Universidad Complutense de Madrid, Spain) collections obtained from the upper Cenomanian and lower Turonian of the Iberian Trough have been revised. Subsequently, new representatives of the species Vtzscoceras gamai, V. charoni sp. nov., V.barcoicense, V.durandi, V. cauvini, V. amieirense, V. harttii and V. kossmati have been studied and presented. Finally, studies of the morphologies and the geographical and stratigraphical distributions of these ammonites have led to the identification of several phylogenetic relationships between them, and to distinguishing two main phases in the evolution of the family Vascoceratidae, characterised by the successive dominance of the "primitive" Vtzscocerasand of the "evolved" Vtzscoceras. Key words: upper Cenomanian- lower Turonian- Arnmonoidea- Vascoceratidae-Va scoceras- new species -Iberian Trough-Spain

Contents

Zusammenfassung ...... V. charoni sp. nov...... Summary ...... V.barcoicense CHOFFAT, 1898 ...... Introduction ...... V.durandi (THOMAS & PERON 1889) ...... Historical background ...... V.cauvini CHUDEAU, 1909 ...... Revision of Vtzscocerasfrom the Iberian Trough in the V. amieirense CHOFFAT, 1898 ...... Universitat Ttibingen and the Universidad Complutense V.hart tii (HYATT 1870) ...... de Madrid ...... V.kossmati CHOFFAT, 1898 ...... New data on Jlascocerasin the Iberian Trough ...... Conclusions ...... Systematic Palaeontology ...... Acknowledgements ...... Family Vascoceratidae DouvrLLE, 1912 ...... References ...... Genus JlascocerasCHOFF AT, 1898 ...... Explanation of the Plates ...... V.gamaiCHOF FAT, 1898 ......

Addresses of the authors: FERNANDO BARROSO-BARCENILLA, Departamento de Paleontologia, Instituto de Geologia Econ6mica (CSIC), Facultad de Ciencias Geol6gicas, Universidad Complutense de Madrid. 28040 Madrid, Espafta,Spain. I Departamento de Geologia.Edificio de Ciencias, Universidad de Alcala de Henares, 28871 Alcala de Henares, Espafta,Spain. E-mail: [email protected] ANTONIO GoY, Departamento de Paleontologfa, lnstituto de Geologfa Econ6mica (CSIC), Facultad de Ciencias Geol6gicas, Universidad Complutense de Madrid, 28040 Madrid, Espafta, Spain.

0375-0442/10/0290/0004 $ 16.65 © 2010 E. Schweizerbart'sche Verlagsbuchhandlung, D-70176 Stuttgart 200 Barroso-Barcenilla & Goy

Introduction raining the most complete and typical marls of the In this paper, firstly, a revision of the ammonites studied interval in the Iberian Trough, as asserted by assigned to the genus Vascoceras CHOFFAT, 1898, ob­ FLo�ET et al. (1982), FLo�ET (1991), SANTAMA­ tained fr om the upper Cenomanian and lower Turo­ RiA-ZABALA (1991, 1992, 1995) and SEGURA et al. nian of the Iberian Trough that are held in the Univer­ ( 1993 ). In order to establish a more precise taxonomic classification,the original types attributed to this fam­ sitat Ttibingen (UT), Germany, and the Universidad Complutense de Madrid (UCM), Spain, is presented. ily that are held in the Museu do Instituto Geol6gico e Mineiro de Lisboa, Portugal, and the Musee National These centres hold the WIEDMANN (JW), GoY (AG), CARRETERO (CM) and MELENDEZ (MH) collec­ d 'Histoire Naturelle de Paris, France, were also stud­ tions, which now contain the largest number of repre­ ied. Thirdly, the obtained data are interpreted, and several conclusions concerning the systematic, distri­ sentatives of the fa mily Vascoceratidae DouviLLE, 1912, collected from the Iberian Tr ough. Secondly, a bution and evolution of the Vascoceratidae are pre­ taxonomic analysis of the members of this genus ob­ sented. In the present paper, the palaeogeographical tained during this research is developed. Geographi­ division (Text-fig. 2) and the ammonite zonation cally, the field work was carried out in the upper Ceno­ (Text-fig.3) for the upper Cenomanian and lower Tu­ manian and lower Tu ronian outcrops situated in the ronian of the Iberian Trough proposed by BARROSO­ localities ofPuentedey (PU), in the north of the prov­ BARCENILLA et al. (2009) have been fo llowed. ince of Burgos, of Fuentetoba (FT), in the centre of Soria, and ofCantalojas (CC), Galve de Sorbe (CG), Historical background Condemios (CA; CB), Somolinos (CS) and Tamaj6n From the beginning of the 20'h century many ce­ (TA), in the north ofGuadalajara, Spain (Text-fig. 1). phalopods have been collected from the upper Ceno­ Stratigraphically, the ammonites presented here manian and lower Tu ronian, not only in the Iberian were mainly collected from the Margas de Puentedey Trough but in the whole of Spain, and assigned to the (FLO�ET et al. 1982) and Margas de Picofrentes family Vascoceratidae DOUVILLE, 1912. KAR REN­ (FLO�ET et al. 1982) fo rmations, deposited in the BERG (1935) identified a significantnumbe r of repre­ inner and the marginal environments of the platform, sentatives of the genus Vascoceras CHOFFAT, 1898, in respectively. Thes e formations are remarkable fo r con- the Outer Navarro-Cantabrian Platform and the

ca1 Cl8lllll�caJrbir�caJ • Puentedey Fuentetoba #/b / &J!!' alve de Sorbe"-; 71©1/!fJ� Cantalojas j Somolino�/!f;�&J� �\®�T Condemios � S>� 0 Tamajon ���el MADRID

00.

200km

Text-fig.1. Geographic provenance of the specimens presented in this paper. The ammonite genus Vtzscoceras CHOFFAT, 1898 201

JL" Bas<(Ue Bnl�de�· I I Fucnteh)l)a T T Protoatlantic �======��Ocean

Text-fig.2. Divisions followed in the Iberian Trough, the approximate locations of the outcrops studied and the geographic boundaries between their different palaeogeographic areas.

North-Castilian Sector. WIEDMANN ( 1960, 1964) trasted with the ones obtained in other regions of the obtained numerous members of the group in the Iberian Trough by BARROSO-BARCENILLA (2006) Outer Navarro-Cantabrian Platform and the North­ and have been presented here. Castilian and Central sectors, many of them in Puent­ Other relevant contributions to the knowledge of ed ey, Fuentetoba and Somolinos. WIEDMANN (1975) the upper Cenomanian and lower Tu ronian cephalo­ presented several ammonite sequences containing var­ pods in the Iberian Trough and in other adjoining ious representatives of Vtzscoceras from the Upper Cre­ and closely related palaeogeographical regions, such as taceous of the Central Sector. MOJICA & WIEDMANN the North-Cantabrian and the Pyrenean basins, (1977) analysed some ammonite sequences from the both located in the north of Spain, were reported Cenomanian of the Central Sector, which included by BARROSO-BARCENILLA (2007), BARROSO­ several members of the genus. WIEDMANN & KA.UFF­ BARCENILLA & GoY (2007, 2009) and BARROSO­ MAN (1978) and WIEDMANN (1979) identified nu­ BARCENILLA et al. (2009). merous ammonites attributed to Vascoceras in the Outer Navarro-Cantabrian Platform, the North Cas­ Revision of Titscoceras from the Iberian Trough tilian Sector and the La Demanda Area. CARRETERO­ in the Universitat Tri.bingen and the MoRENO (1982) collected some representatives of Universidad Complutense de Madrid the genus in several outcrops of the Levantine Plat­ Regarding the revision of the ammonites assigned fo rm, and in Cantalojas, Galve de Sorbe, Condemios to the genus Vtzscoceras CHOFFAT, 1898, obtained and Somolinos. MELENDEZ-HEVIA ( 1984) identified from the Iberian Trough that are held in the numerous members of Vtzscoceras in Condemios, So­ WIEDMANN (Universitat Tubingen, Germany) and molinos and Tamaj 6n. FLOQ.!:!:ET (1991) cited some GoY, CARRETERO and MELENDEZ (Universidad representatives of the genus in the northern halfof the Complutense de Madrid, Spain) collections, it is im­ Iberian Trough and in the Basque Basin, in the north portant to emphasize certain remarks. It has not been of Spain. SEGURA et al. ( 1993) analysed several strati­ possible to find specimens of all the members of graphic sequences deposited in the Cenomanian-Tu­ Vtzscoceras cited by WIEDMANN (1960, 1964, 1975, ronian transition of some parts of the Central Sector, 1979), MOJICA & WIEDMANN (1977) and WIED­ one of them in Somolinos, and mentioned various MANN & KAuFFMAN (1978) in the UT. Therefore, representatives of the genus. BARROSO-BARCENILLA the results of the investigation at this research centre (2004) identified several members of Vtzscoceras in the are based exclusively on the specimens that are now upper Cenomanian and lower Turonian of the north­ hosted in theJW Collection. Likewise, the method by ern margin of the North-Castilian Sector, specifically which the fossils of the CM Collection have been in Puentedey. His conclusions on the genus were con- numbered and identified has hindered an adequate 202 Barroso-Barcenilla & Goy

IberianTrough. Spain. �= =���0�i BARROSO-BARCE-,ILLA Substage TRI)(;FR&Kl'Nl'EDY (l996), 131-.l'GTSON( 1996)and Cl al. (2009) GRADSTEIN cf a l . (2�) / .• m l

Clr�ffaticeras (('lto.lfath·eras) qmwsi

Spathllt'.l Neocardio<:era.v (Jermrogenceras) juddii .\/Jbconciliatus

Vascocera.r gamm -\fetoicocerat MelOicocercJ.\ gesbmammt ge.dinianum \lt'ttNcocerru ge5flfuanwn

Metoicoceras Neulohlles mosbyemt' \.'ibrayewws \Co/ohue� Calycocerar 1 nbra)WUIUS (Proeucall·coccrav) Calyl-ocera\ guerang(•rt (Ca/ycv.K'enu) Euca��'COCt!f'fl.f na�imfare f'OU(.'j

Te xt-fig. 3. Biostratigraphic zonation followed in this work and its correlati on with the standard scale. and individualized tracking of its ammonites. There­ d 'Histoire Naturelle, Paris, France; SGP, Servi<;os fore, the revision of the taxa cited by CARRETERO­ Geol6gicos de Portugal, Lisboa, Portugal; UCM, MORENO (1982) has only been partially carried out, Universidad Complurens e de Madrid, Spain. All the and the referen ces to her work in the synonymy ofthe specimens presented here are held in the Departamen­ taxonomic section are necessarily imprecise. In spite to de Paleontologia of the UCM. of these difficulties, some conclusions on the present situation of the Jlascoceras in the JW, AG, MH and Systematic Palaeontology CM collections have been reached and presented here Family Vascoceratidae DOUVILLE, 1912 (Text-fig. 4). Neoptychitinae COLLIGNON, 1965a (p. 70)

New data on Vascoceras in the Iberian Trough Diagnosis: Genera with spires showing variable In this section, a systematic description of the new sections and degrees ofinvolution that rapidly become records of the genus Jlascoceras CHOFFAT, 1898, ob­ smooth. Maturity with smooth surfaces, or maintain­ tained during fieldwork carried out by the firstauthor ing only umbilical or ventrolateral tubercles, or some as well as of the unpublished memb ers of the group spaced ribs. Sutural el ements usually shallow, although identified in the AG and MH collections is given. The in some genera these may be deep and differen tiated. terminology used fo r the descriptions is based on the Discussion: CHOFFAT (1898) defined the genus glossary of morphological terms proposed by BAR­ Jlascoceras, and suggested the possible relationship ROSO-BARCENILLA (2008). Measurements have been between his new group and Ammonites superstes made with an adjustable caliper, and are given in tenths KosSMAT, 1897. HYATT (1903) included the genera of millimetre and in percentages of the diameter of the Jlas coceras, Tolypeceras HYATT, 1903, and Barrois iceras shelL For comments or location of specimens, the fo l­ DE GROSSOUVRE, 1894, within the fam ily Cosmocer­ lowing abbreviations are used here: UT, Universirat atidae HYATT, 1900. DOUVILLE ( 1912) differentiated Ttibingen, Germany; ICZN, International Code of the "vascoceratines" between the fo rms with normal Zoological Nomenclature; MNHN, Musee National first lateral lobe derived from the "pulchelliides"_ Vascoceras CHOFFAT, 1898, and related groups ---- Original classification Collection Stratigraphical origin Geographical origin Remarks - V. gamai CHOFF A r, I 898, and JW,AG,CM Mainly Guadarrama Area, but also North- Almost all V.gamai Subzone - V. cf. gamai and MH Castilian Sector and La Dcmanda Area - - Specimen ofMELE �DtZ-HFVIA (1984. p. 88, pi. 3, fig. 3 a-b) wilh narrow umbilici and persistent ribs: mundae CHOFFAT, 898 AG gamai Subzone Central Sector, mainly Guadarrama Area V. I V. V. barcoicense - V. cf. mwulae JW V. gamai Subzone Central Sector

o S. (/.) malladae Z ne, M. nodosoides Subzone Dimensions close to V. rog s.wuvrei, but by ossible ribbing and high stratigraphical origin is advisable to reject the V.grossouvrei CHOFFAT, I 898 JW Inner Castilian Platform � and, with doubts, W. munieri Subzone speci tC assignation of these ammonites 1- V. cf. barcoiceme CHOFFAT, S. (/.) mal/adae Zone and M. nodosoides Subogival and compressed section, tectifonn venter, narrow umbilicus, com lex suture (with four saddles) and JW North-Casti lian and Central sectors e 1898 Subzonc stratigraphical range close to evolule and compressed Clrof!aticeras( lrof!aticeras) HYA.II, 1903 '"� - V.d"ro"di{T,OM"& From C. (C.) quaasi Zone to M. nodosoides North-Castilian and Central sectors I 889) and V.cf. durandi Subzonc - - V. douvil/ei CHOFFAT, I 898, From C. (C.) quaasi Zone to M.nodosoides JW North-Castilian and Central sectors and V. cf. douvillei Subzone

V. cf. doul'illei CM - Guadarrama Area -

V. cauvini CHUDEAU. 1909 CM - Guadarrama Area

cf. Zone V. amieirense CHOFFAT, S. (J.) subconci/iarus and lower Most of these specimens are anributable t o other taxa, such as the number 957 that may be assigned to V. durandi , JW North-Ebro Area and Central Sector 1898 Turonian but some other ammonites could be transitional Vascocer as - f- Upper part of C. (C.) quaasi Zon and S. (!.) V. harttii (HYA"IT, 1870) AG andMH e Guadarrama Area malladae Subzone ·-

V. cf. harttii JW Lower Turonian La Dcmanda Area :?(1> I'> V. kossmati CHOFFAT, 1898, S. (1.) malladae Zone and M.nodosoides s JW South-Ebro and La Oemanda areas - and V. cf. kossmati Subzonc s 0 V. harttiforme CHOFFAT, Mainly S. (/.) ma/ladae and M. nodosoides Mainly Central Sector, but also South-Ebro JW - :::1 I 898, and V. cf. harttiforme zones Area ;::;· (1> V. cf. carteri (BARRER, I 957) AGand MH S. (!.) malladae Subzone Guadarrama Area Specimens that should be assigned to V. kossmati 1-- �:::1 V. globosum (REYMENT, S. (/.) malladae Zone and M.nodosoides � JW South-Ebro Area "' 1954b) Subzone -- 1940, Zone V. triangu/are FARAUO, S. (/.) malladae and M. nodosoides V. triangulare must be considered as a synonym of Choffaticeras(Chof!aticeras) douvillei (PbRON, 896),1 as JW South-Ebro Area � and V. cf. triangulare Subzonc indicated by BARROSQ-BARCENILLA & GOY (2007) 8 CllOFFAT, 1898, V. silmnense V. siii'Gnense seems restricted to the upper Cenomanian 1898; KENNEDY et al., 1989) and, thus, the � JW S. (/.) ma/ladae and M. nodosoides zones - (CHOFFAT, and V. cf. silvanense stratigraphical distribution attributed by WIEDMANN to these specimens may be incorrect __ �"' (COLLIGNON, V. rumeaui Two morpbologics: monites with tbe proportions, narrow umbilici aodjuvemle ornamentationof V. barcoicense; CM . Guadarrama Area am (") 1957) specimens with the dimensions and lack of ornamentationof V.durandi � ::c: V. (Pachyvascoceras) 0 JW . La Dcmanda Area Globose and involute ammonites close to V. kossmati .,., crassum (FURON, 1935) .,., - From S. (J.)subconciliatus Zone to S. (!.) Specimens from .subconci/iatusand C. (C.) quaasi zones with unded and depressed section: V. durandi; Paravascoceras spp. and MH Guadarrama Area S. (J.) r:o AG ammonites Zone and Subzone t an u a and omp d section: V. mal/adae Subzone from C. (C.) quaasi S. (/.) malladae w1th n g l r c rc))� amiei� -�...... � 00 PlesiOI'Oscoceras sp. AG Upper part of V. gamai Subzone G uadarrama Area Specimens with the diagnostic features of V. clraroni sp. nov. '-0 00 - Subzone Upper part of V.gamai and low r Not previously classified ammonites with narrow umbilici, early lack of umbilical tubercles and persistent ribs: - AG e Guadarrama Area part of S. (J.) subconciliarus Zone V. harcoicense N I 0 I \.;) Te xt-fig.4. Ammonites attributed to the genus Va scocera s CHOFFAT, 1898, and related groups in the WrEDMANN (JW), GOY (AG), CARRETERO (CM) and MELENDEZ (MH) collections. 204 Barroso-Barcenilla & Goy

Within the "vascoceratines" he included the genera and ?Microdiphasoceras COBBAN et al., 1989, within Vas coceras and Fagesia PERVIN�IERE, 1907. SPATH the family ofDOUVILLE. ( 1925) suggested Vascoceratidae as the name fo r this Regarding the phylogeny of the Vascoceratidae, group, which was finally accepted, and included the REYMENT (1955) considered that the origin of the genera Vascoceras, Tho masites PERVIN�IERE, 1907, family could be located in Nigericeras. However, in and Plesiovascoceras SPATH, 1925, within the family. 1979 the same author stated that several of the Acan­ REYMENT (1954a) integrated the genera Vascoceras, thoceratidae taxa could be involved in the origins of Pa ra ma mmites FuRON, 1935, Nigericeras the Vascoceratidae, and therefore he indicated that the ScHNEEGANS, 1943, Neoptychites KossMAT, 1895, latter group could be considered as a polyphyletic fa m­ and Ezilloella REYMENT, 1954a, within the Vascocer­ ily. CooPER ( 1979) indicated that Vascoceras di­ atidae. KUMMEL & DECKER ( 1954) described the ge­ artianum (D 'ORBIGNY 1850) gave rise to Pa ravas­ nus Sp ath ites , assigning it to this same family. WRIGHT co ceras and Vascoceras. WRIGHT & KENNEDY (1981) in MooRE (1957) stated that the genera Nigericeras, considered that Vascoceratidae derived from Acan­ Sp ath ites, Go mbeo ceras , Ezilloella , Pa ravascoceras thoceratidae by ornamental reduction and sutural FuRON, 1935, Pa chyvas co ceras, Vasco ceras, Para ma m­ simplification. ZABORSKI (1996) also defended the mites, Plesiovascoceras, Fagesia and Neoptychites belong common origin of Pa ravasco ceras and Vascoceras, indi­ to the Vascoceratidae. cating that both groups seemed to have originated BARBER ( 1957) attributed to this family the same from Nigericeras, and maintained that his new genus genera as WRIGHT in Moo RE ( 1957), although he Pseudovascoceras could derive from Cunningtoniceras considered Para cantho ceras and Pa chyvascoceras to be COLLIGNON, 1937. synonyms of Paravasco ceras. WIEDMANN ( 1960, Distribution: From the upper Cenomanian to 1964) asserted that the subfamily Pseudotissotiinae the upper Turonian of many countries, mainly belong­ HYATT, 1903, in which he included Coilopoceratidae ing to the palaeogeographical environment of the HYATT, 1903, should be integrated into the Vascocer­ Tethys. The earliest records of the family correspond atidae. Furthermore, he stated that Discovascoceras to North American specimens of V. diartianum from CoLLIGNON, 1957, shows a strong resemblance to the Sciponoceras gra cile Zone, almost equivalent to the Ch ojfaticeras HYATT, 1903. KENNEDY et al. (1980) Meto ico ceras geslin ianum standard Zone. In the Iberi­ transferred Spathites, and its synonym Fallotites WIED­ an Trough, the Vascoceratidae have been identified MANN, 1960, to Mamm itinae HYATT, 1900. WRIGHT from the V.ga ma i Subzone to the middle Tu ronian. & KENNEDY (1981) divided the family into the Vas­ coceratinae, with normally rounded ventral area with­ Genus VascocerasCH OFFAT, 1898 out keels, and the Pseudotissotiinae, with relatively Pachyvascoceras FuRON, 1935, p. 58, type species Pa chvascoceras involute whorls and flatfla nks. CHANCELLOR ( 1982) crassum FURON, 1935, by subsequent designation ofREYMENT, presented a summary of the modifications es tablished 1954b, p. 257. in the taxonomic organisation of the Vascoceratidae, Paracanthoceras FURON, 1935, p. 59, type species by original since the work of WRIGHT in MoO RE ( 1957) until designation Va scoceras (Paracanthoceras) chevalieri FURON, the year in which his study was written. RENZ (1982) 1935. Pa ra vascoceras FuRON, 1935, p. 60, type species by original proposed the inclusion ofPara ma mmites within Acan­ designation f!ascoceras cauvini CHUDEAU, 1909. 1894. thoceratidae DE GROSSOUVRE, KENNEDY et Broggiiceras BENAVIDEs-CAcERES, 1956, p. 469, type species by al. (1989) and COBBAN et al. (1989) suggested that original designation Broggiiceras olssoni BENAVIDES-CACERES, Nigericeras should be in cluded within the Acanthocer­ 1956. atidae. CHANCELLOR et al. ( 1994) stated that the Greenhornoceras COBBAN & ScoTT, 1972, p. 84, type species by pseudotissotiids seemed to be derived directly from original designation f!ascoceras ( Greenhornoceras) birchbyi the acanthoceratids, and therefore recognised the fa m­ COBBAN & SCOTT, 1972. ily status of the fo rmer group, as KENNEDY (1994). Provascoceras COOPER, 1979, p. 123, type species by original designation Ammonites diartianus D · 0RBIGNY, 1850. ZABORSKI (1996) assigned Pa ravasco ceras and his Na nnovascoceras RENZ & ALVAREZ, 1979, p. 978, type species new genus Pseudovascoceras to the Acanthoceratidae. by original designation Na nnovascoceras intermedium RENZ & WRIGHT in KAESLER (1996) included Vascoceras, ALVAREZ, 1979. Ezilloella , Neoptych ites, Fagesia, In fobricaticeras Type species: vascoceras gamai CHOFFAT, 1898. Although CoBBAN et al., 1989,Rubroceras CoBBAN et al., 1989, some authors stated that the designation of this species as The ammonite genus TlascocerasCHO FFAT, 1898 205

genotype corr esponds to CHOFFAT (1898) or ROMAN (1938), SCHNEEGANS ( 1943) considered the subgenera Para­ the firstauthor who explicitly mentioned V gamai as the type of canthoceras and Pachyvas co ceras as synonyms of Par­ Vascoceras was DIENER (1925, p. 182), as indicated by avascoce ras . He also gave generic status to Paravascocer­ CHANCELLOR et al. (1994). as, and suggested that this group should include the Diagnosis: Variable morphology with oval, sub­ species with the typical suture lines of Tlascoceras, with triangular or rounded embracing whorl section, suba­ oval whorl section and with ribs. REYMENT (1954b) cute, arched or plane ventral region, steep umbilici, considered that the name Paracanthoceras should and generally globose and involute adult cadicone. maintain priority over the term Paravascoceras, but Only the first whorls may show umbilical tubercles later he ( 195 5) again preferred the designation Par­ and also, in some species, small and rounded ribs cross­ av asco ceras fo r this group. BENAVIDES-CACERES ing the ventral area, constrictions and fe eble ventrola­ (1956) described the species B. olssoni and B. hum­ teral tubercles. The ornamentation disappears progres­ boldti, which show smooth inner whorls and adult or­ sively during ontogeny, although the umbilical tuber­ namentation composed of strong ribs, and on the basis cles can remain until maturity. �ite simple and ir­ of these fe atures he proposed the genus Broggiiceras. regular suture lines, with three bifid orindented lobes WIEDMANN (1960, 1964), taking Ammonites and wide saddles on each flank.Th e firstlobes are wide reveliereanus CouRTILLER, 1860, as type, described and low, with palm-shaped terminations. Fallotites, the group in which he included V.subc on cil­ Discussion: CHOFFAT (1898) established the ge­ iatum. COLLIGNON (1965b) maintained Paravas­ nus Tlas coceras and noted that the species he described co ceras as a differentgenus. FREUND & RAAB (1969) and included in this group could be divided into fo ur did not consider Discovas co ceras CoLLIGNON, 1957, conjuncts. The firstis composed of taxa with only one as a differentiated genus, and included it in Paravas­ type of tubercles and wide umbilici, such as V.gamai, co ceras, together with Paracanthoceras and Pachyvas­ including its variety subtriangularis, and V. mundae. co ceras . COBBAN & ScoTT (1972) described Tlas co cer­ TI1e second comprises subglobose species with round­ as ( Greenhornoceras) as the most recent subgenus of ed umbilical margins, such as V. silvanense, V. douvil­ Tlascoceras. These authors indicated that this subgenus lei and V. adonense. The third is composed of globose differsfr om V. ( Tlascoceras) in developing a larger invo­ taxa with sharp umbilical margins, such as V. amieir­ lution grade and maintaining a subrectangular whorl ense and V. kossm ati . Finally, the fo urth comprises the section. ScHOBEL ( 1975) concluded that several spe­ multi tuberculate species V. subconcili atus. This author cies should be considered as synonyms of V. cauvini, remarked, however, that some specimens with inter­ whereas Pachyvasco ceras should be included in Par­ mediate fe atures cannot be clearly assigned to any of av as co ceras, which fo r this author maintained the ge­ these fo ur categories. neric status. BERTHOU & LAUVERJAT ( 1975) studied PERVIN�IERE ( 1907) considered that V. dou­ the Va scoceratidae in its type region, and BERTHOU et villei is a synonym of V. du randi, and suggested that al. ( 1975) included in the genus Tlas co ceras the groups V. amieirense might be included within the morpho­ of V.gam ai-mundae, of V. barcoicense, of V. douvillei logical variability of V. du randi. FUR ON ( 1935) dif­ and V. amieirense, of V. kossm ati and of V.sub con cili a­ fe rentiated fo ur subgenera in Tlas co ce ras. The firstone, tus. The first is composed of specimens with evolute Pachyvascoceras, includes globose and involute species and quite compressed shell, umbilical tubercles that with umbilical tubercles, deep umbilici and lack of or­ disappear near the adult body chamber, rounded or namentation at maturity, such as V.silvanense, V. dou­ subtriangular whorls and feeble ribs during early on­ villei, V. adonense, V. amieirense, V. harttiforme and togeny. The group of V. barcoicense comprises more V.kos sm ati. The subgenus Param ammites groups non­ involute specimens with fe eble plications on the adult globose and tuberculate taxa, such as V. subconcili atus body chamber. The group of V. douvillei and V. amiei­ and V.polymo rphum PERVIN�IERE, 1907. The sub­ rense is characterised by specimens that differ in the genus Paracanthoceras includes involute and non-glo­ whorl section. The group of V. kossm ati, a taxon con­ bose species, less ornamented and with complex suture sidered indistinguishable from V. harttiforme, com­ lines, such as his new species V. chevalieri. Finally, the prises involute and globose specimens with narrow subgenus Paravascoceras groups compressed and invo­ umbilici and flattened whorls of subrounded or sub­ lute taxa with ribs and simple suture lines and without triangular section. The fifth and last group includes tubercles, such as V. cauvini CHUDEAU, 1909. V. subcon cili atus. Finally, BERTHOU et al. ( 1975) con- 206 Barroso-Barcenilla & Goy

eluded that, with the exception of V subcon cili atus, all giiceras, Greenhornoceras, Provascoceras and Nanno­ the original material ofCHOFFAT (1898) could be as­ vascoceras. Regarding Paravas co ceras, there is no doubt signed to Vas co ceras, and that this genus should only be that its type, V cauvini, is an unusual species among divided into Vs. str. and V (Pachyvascoceras), which those of the genus of CHOFFAT, as it fo llows a slightly includes the species V douvillei, V amieirense and different ontogenetic development. Nevertheless, this V kossm ati. species lacks the juvenile ornamentation of the Acan­ CooPER (1978) considered Greenhornoceras as a thoceratidae, and shows simple suture lines. Therefore, synonym of Vas coceras. REYMENT & CHANCELLOR it seems to be more appropriate to assign V cauvini, ( 1978) studied the common occurrence of members and consequently Paravascoceras, to the family Vas­ of Vas coceras in Spain and West Africa. RENZ & ALvA­ coceratidae, specifically to thegenus Vas co ceras. REZ ( 1979) described Nannovascoceras on the basis of As regards the phylogeny of the group, although several tiny ammonites with certain constrictions dur­ fo r many years it was believed that the origin of Vas­ ing their young stages. CooPER (1979) proposed co ceras was in Nigericeras SCHNEEGANS, 1943, Provascoceras as a new genus fo r small Cenomanian WRIGHT & KENNEDY (1980) and KENNEDY & specimens with marked ribs on the ventral area and WRIGHT (1994) indicated that the genus of CHOF­ numerous tubercles on the adult body chamber. FAT could be derived directly from Protacanthoceras WRIGHT & KENNEDY (1981) considered that the di­ SPATH, 1923. CHANCELLOR et al. (1994) concluded agnostic fe atures of Pro vasco ceras did not justify its that the origin of Vas co ceras, and therefore of the Vas­ generic status. RENZ ( 1982) included Greenho rno ceras coceratidae, was in Protacanthoceras proteus WRIGHT in the synonymy ofPs eudoneoptychites LEANZA, 1967. & KENNEDY, 1980, exactly in P.proteus vascoceratoides BERTHOU et al. ( 1986) included in the genus of WRIGHT & KENNEDY, 1987, by loss of the ventrola­ CHOFFAT the species V di arti anum (D 'ORBIGNY teral and siphonal tubercles. ZABORSKI ( 1996), ob­ 1850), V harttii (HYATT 1870), Vgam ai , V barcoi­ serving the ornamentation and the bifidfo rm of the cense, V kossm ati, V cauvini CHUDEAU, 1909, Vpioti first lateral saddles of V diartianum, indicated that (PERON in FoURTAU 1904), V birchbyi COBBAN & this species is transitional between the Acanthocerati­ SCOTT, 1972, and, as a possible synonym ofHYATT 's dae and the Vascoceratidae. taxon, V durandi (THOMAS & PERON 1889). They Distribution: Upper Cenomanian and lower Tu­ considered V amieirense and V douvillei to be con­ ronian of Portugal, France, Spain, the UK, Germany, specific fo rms of V durandi; V mundae, V adonense Croatia, north, west and east of Africa, Madagascar, and Ammonites (?V) grossouvrei of V gamai; and the Middle East, Brazil, Peru, Colombia, Mexico, V harttiforme of V kossm ati. These authors also indi­ Venezuela, the USA and Japan. Its oldest records oc­ cated that V silvanense CHOFFAT, 1898, V tavense cur in the S. gracile Zone, almost equivalent to the M. FARAUD, 1940, and V triangulare FARAUD, 1940, are geslinianum standard Zone. In the Iberian Trough, the nomin a dubia. Finally, BERTHOU et al. ( 1986) high­ genus ranges from the base of the Vgam ai Subzone to lighted the complexity of the classificationof the Ni­ the upper part of the S. (I.) malladae Subzone of the gerian species assigned to Vas coceras. KENNEDY ( 1994) Inner Castilian Platform. Nevertheless, Vasco ce ras is excluded V (Pachyvascoceras) tri angulare from this ge­ better represented in the Central Sector than in the nus, attributing it to Choffaticeras HYATT, 1903, as he North-Castilian Sector. considered this species as a synonym of Choffaticeras ( Choffaticeras) douvillei (PERON 1896). ZABORSKI ( 1996) proposed the genus Pseudovascoceras, designat­ Vt:zscocerasgamai CHOFFAT, 1898 ing V. nigeriense Wo oDS, 1911, as its type species. (Plate 1, figs.A-L; Text-fig. 5 A) Within his new group he included specimens without tubercles during mid and advanced ontogeny and of­ 1898 Vttscoceras gamai CHOFFAT, p. 54, pi. 7, figs. 1-4; ten with strong ventral ornamentation, and integrated pi. 8, fig. 1; pi. 10, fig. 2; pi. 21, figs. 1-4. var. Pseudovascoceras and Paravascoceras in the Acan­ 1898 Vttscoceras gamai subtriangularis CHOFFAT, p. 55, pi. 7, fig. 5; pi. 21, fig.5. thoceratidae DE GROSSOUVRE, 1894. 1898 Vttscoceras mundae CHOFFAT, p. 56, pi. 10, fig. 1; On the basis of the above remarks, there would pi. 8, figs. 2-4; pi. 21, figs. 6-8, 10. seem to be insufficient grounds fo r maintaining the 1898 Vttscoceras adonensis CHOFFAT, p. 59, pi. 9, fig.3; separation of Pachyvascoceras, Paracanthoceras, Brog- pi. 21, fig.12 a-b. Theammonit e genus Vas co ceras CHOFFAT, 1898 207

1898 Ammonites (?vascoceras) grossouvrei CHOFFAT, p. cf. 1979 Vtlscoceras cf. gamai CH OFFAT - WIEDMANN, p. 68, pl. 9, figs. 1-2; pl.22, figs.37-38. 203. ? 1898 Ammonites (? vascoceras) c£ barcoicensis CHOFFAT, 1981 Vascoceras gamai CHOFFAT - WRIGHT & p. 67, pl. 16, fig.11; pl. 22, fig. 36. KENNEDY, text-fig.29 g-i. aff1920 Vascoceras affgamaiCHOFFAT - BosE, p. 216, pl. cf. 1982 Vtlscocerascf.gamai CHOFFAT- CHANCELLOR, p. 15, figs. 3-5, text-fig.3. 97, figs.26-28. 1928 vascoceras gamai CHOFFAT - DouvrLLE, p. 13, 1982 vascoceras gammai [sic] CHOFFAT- CARRETERO­ fig.3; pl. 1, fig.4. MORENO, P· 255. ? 1957 vascoceras gamai CHOFFAT - COLLIGNON, P· 1984 Vtlscocerasgamai CH OFFAT- MELENDEZ-HEVIA, 122, pl. 2, fig.2. p. 85, pl. 2, fig. 1 a-b; pi. 3, figs. 1-2. c£ 1960 Vascocerascf. gamai CHOFFAT- WIEDMANN, pp. 1984 Va scoceras mundae CHOFFAT - MELENDEZ­ 712,720. HEVIA, p. 88 (only). ? non 1960 Vascoceras (Pachyvascoceras) grossouvrei CHOFFAT 1986 Vtlscocerasgamai CHOFFAT - BERTHOU et al., p. - WIEDMANN, PP· 712, 720. 66, pl. 2, figs. 1-12; pi. ?3, figs.1-3, 5-7, I 0, 13- 1960 Vtlscoceras (vascoceras) gamai CHOFFAT 14. WIEDMANN, p. 723. 1989 Vtlscoceras gamai CHOFFAT - LUGER & c£ 1960 Vtlscoceras (vascoceras) cf. mundae CHOFFAT - GROSCHKE, p. 378, pl. 40, figs.5, 7, text-fig.6 c. WIEDMANN, p. 723. ? 1989 Vtlscoceras cf.gamai CHOFFAT- COBBAN et al., p. cf. 1964 vascoceras (Vascoceras) cf. gamai CHOFFAT - 45, figs. 44, 87 w-aa, ee-rr. WIEDMANN, PP· Ill, 115. 1992 Vtlscocerasgamai CHOFFAT- THOMEL, p. 228, pi. ? non 1964 vascoceras (Pachyvascoceras) grossouvrei CHOFFAT 121, figs.1-2; pl. 121, fig.4. - WIEDMANN, pp. Ill, 115. 1996 Va scoceras gamai CHOFFAT - AMEDRO et al., p. 1964 Vascoceras (vascoceras) gamai CHOFFAT 213, fig. 17 a-d. WrEDMANN, p. 116. 2001 Va scoceras gamai CHOFFAT - CALLAPEZ & cf. 1964 vascoceras (vascoceras) c£ mundae CHOFFAT - FERREIRA, p. 51, pl. 5, figs. 3-6; pl. 6, figs. 1-2, WIEDMANN, p. 116. text-figs. 16.4-11, 20.9. 1965b vascocerasgamai CHOFFAT- COLLIGNON, p.l83 2001 Vascoceras adonense CHOFFAT - CALLAPEZ & (21), figs. 5-7. FERREIRA, p. 64, pi. 8, figs. 4-5; pi. 9, figs. 1-2, 1967 vascoceras gamai CHOFFAT - POLSAK, p. 137, pi. text-figs.19.2-3, 19.5-6, 19.8-10. 83, fig.3. ? 2005 Vascoceras gr. gamai CHOFFAT - MEISTER & 1967 vascoceras (Pachyvascoceras) grossouvrei CHOFFAT ABDALLAH, p. 134, pi. 13, figs. 1-2, 4, text-fig. - POLSAK, p. 137, pi. 83, fig.1. 25. cf. 1975 Vascoceras cf. gamai CHOFFAT - WIEDMANN, p. 2006 Vascoceras gamai CHOFFAT BARROSO­ 141. BARCENILLA, p. 220, pi. 28, figs. a-1; pi. 29, figs. 1977 vascoceras ex. gr. V.gamai CHOFFAT- MO]ICA & a-c, text-fig.66. WIEDMANN, p. 749, pl. 1, fig.1. Type: The holotype designated by CHOFFAT (1898, p. 54, 1978 vascoceras gamai CHOFFAT - WIEDMANN & pl. 7, fig.l; pi. 21, fig. 1) was destroyed. Among the paratypes l

Dimensions: D H (%) E (%) 0 (%) Range 1245-408 548-169 (49-32) 587-185 (56-23) 445-124 (41-29) Mean 963 376 (39) 364 (39) 327 (34)

Description: Specimens with slightly compressed row and rounded margins. In early ontogeny they or depressed rounded whorl section, evolute coiling, show 5-7 large and rounded umbilical tubercles per convex ventral area, convergent flanks, and relatively whorl. From each of these can grow 1-2 sinuous, di­ wide umbilici with subvertical and low walls, and nar- vergent and slightly prorsiradiate ribs, which become 208 Barroso-Barcenilla & Goy

stronger aftercrossing the ventral area. When present, The coincidence of the geographical and strati­ the accessory ribs grow from the flanks,intercalated at graphical distributions of V.ga mai and V.mundae in regular intervals. During ontogeny the whorl section the Iberian Trough, as well as the great similarity be­ becomes slightly compressed, the flanks flatten and tween both fo rms, supports the conclusions of the ornamentation disappears, leaving the adult body BERTHOU et al. ( 1986) in this respect. Likewise, after chamber completely smooth and with a certain ten­ having observed in SGP that the types of V.adonense dency to uncoiling. and A. (?V.)gros souvrei present the typical features of Discussion: CHOFFAT (1898) included V.gamai V.ga mai, in this paper both taxa have also been in­ within the fo rms of the genus he called monotubercu­ cluded in the synonymy of Vga mai. On the contrary, late with wide umbilici, and described other close taxa, V.ga mai var. mahafolensis COLLIGNON, 1965a, ex­ such as V.mundae, V adonensis and Ammonites (? v.) cluded from the synonymy of V.ga mai by BERTHOU grossouvrei. The possibility that V mundae, V.adon ­ et al. ( 1986), shows persistent and strong umbilical tu­ ensis and A. (?V.) grossouvrei were synonyms of bercles and complex suture lines, and seems to be clos­ V.gamaiwas firstsuggested by BERTHOU et al. (1975). er to the groups Sp athites or Pseudaspidoceras HYATT, These authors stated that it is impossible to establish a 1903, than to the genus Vascoceras. limit between V.ga mai and V. mundae, and consid­ The evolute Ammonites (?v.) cf. barcoicensis of ered that V.ga mai var. subtriangularis, V.ga mai var. CHOFFAT (1898, pl. 16, fig. 11; pl. 22, fig.36) lacks lissa CHOFFAT, 1898, V.adonense and A. (?v.)gros ­ ribs and shows an umbilical width closer to V.ga mai. souvrei could be included in the morphological varia­ The fragmented V.ga mai ofCOLLIGNON (1957) has bility of the first taxon. BERTHOU et al. (1986) re­ some fe atures of the same species, bur they are not sig­ garded V.ga mai var. subtriangularis CHOFFAT, 1898, nificantenough to make a precise taxonomic determi­ V. mundae, V. adonense and A. (?V.) grossouvrei as nation. Some of the V. cf. gamai of COBBAN et al. synonyms of Vga mai. They integrated nearly all of the (1989) show quite narrow umbilici, weak umbilical most evolute, compressed and monotuberculate forms tubercles and persistent ribs. These features are close of the genus in this species, and indicated that the to those of V. barcoicense, suggesting that these am­ slight differences in the ornamentation, the degree of monites may be transitional fo rms between this spe­ involution and the outline of the suture lines of cies and V.ga mai. V. mundae, V. adonense and A. (?V.) grossouvrei do Distribution: This species has been identified in not justify their specific separation. Finally, the same the upper Cenomanian, where it reaches its maximum authors also stated that the outline of the suture lines abundance, and in the base of the lower Turonian of cannot be accepted as a feature fo r classifying the Portugal, Egypt, Algeria, Spain, Croatia, France and, members of Vascoceras, and they considered that the possibly, Mexico and the USA. In the Iberian Trough, slightly tabulate fo rm of the ventral area of some juve­ it has been collected from the V. gamai Subzone of the nile specimens of this genus reveals the existence of a Inner Castilian Platform. close relationship with Spathites KuMMEL & DECKER, 1954.

A 8 Te xt-fig.5. Suture lines. A, Vascocerasgamai CHOFFAT, 1898, TA-S-1 05, from the Vascoceras gamai Subzone ofTamaj6n. B, Vascoceras charoni sp. nov., MS-3, holotype, from the Vascoceras gamai Subzone ofTamaj6n. Both are x 3/2. The ammonite genus Vascoceras CHOFFAT, 1898 209

Jlascoceras charoni sp. nov. Ty pes: Holotype is specimen MS-3, and paratypes are specimens TA-R-655, TA-S-95, TA -S-122 and TA -S-556. (Plate 2, figs.A-F; Te xt-fig. 5 B) Derivation of the name: The name comes from the 1984 Plesiovascoceras sp. MELENDEZ-HEVIA, p. 95, pi. 3, mythological fe rryman who carries souls across the river Acheron figs. 4 a-b, 5 a-b. to the realm of Hades. 2006 Vascoceras nov. sp. BARROSO-BARCENILLA, p. 226, pi. 29, Material: 5 specimens; MS-3, TA- R-655, TA -S-95, TA -S- figs.d-i, text-fig. 67. 122 and TA-S-556.

Dimensions: D H (%) E (%) 0 (%) Range 838-405 342- 199 (49-32) 548-295 (76-58) 275-134 (44-30) Mean 589 237 (41) 391 (68) 199 (34)

Diagnosis: Small or medium globose specimens, (HYATT 1870), V.kos sm ati CHOFFAT, 1898, V.gfo ­ strongly ornamented fo r the genus Vas co ceras CHOF­ bosum (REYMENT 1954b), V.pro prium (REYMENT FAT, 1898, and with semicircular or subrenif orm whorl 1954b), V. robustum BARBER, 1957, V.poly gonum section and relatively wide umbilici. They present 6-8 BARBER, 1957, and V.effi pticum BARBER, 1957, do strong umbilical tubercles and 20-25 marked ribs of not normally have strong ornamentations, and fo llow triangular section and the same number of semicircu­ the tendency to adopt almost smooth surfaces. Among lar interspaces per whorl. From each umbilical tuber­ the fe w fo rms of this genus with relatively wide and cle grow one or two main ribs. Each radial band with­ ornamented whorls, V. costatum (REYMENT 1954b) out umbilical tubercles has two or three accessory ribs, shows strong ribs, but it has a highly involute coiling which grow from the outer limit of the umbilical mar­ and lacks the characteristic tubercles of V. charoni. gins. The ribbing crosses the flankswithout changing, V.nigeri ense WooDS, 1911, presents strongly marked but as it approaches the ventral area it is usually less ventrolateral tubercles, which can modify the whorl pronounced, and it may even disappear on the sipho­ section to the extent of developing a subpolygonal nal line. During early ontogeny the most characteristic contour, and notably different ornamentation. V.sil­ fe atures are the rounded ventral ridges produced by a vanense CH oFFAT, 189 8, also has relatively wide whorl siphon located at a very outer position, and the feeble section and strong umbilical tubercles, but during ear­ and elongated ventrolateral tubercles located over the ly ontogeny it lacks ribs. V. rumeaui ( COLLIGNON main and the accessory ribs, which disappear quickly 1957) is one ofthe most ornamented taxa of Vascocer­ with growth. During ontogeny the ornamentation be­ as . Nevertheless, it presents subtriangular section, has comes more distant, the umbilical tubercles get strong­ narrower umbilici, lacks umbilical tubercles and shows er and more rounded, and the ribbing and the sipho­ ribs only during advanced ontogeny. Finally, it has nal ridge tend to decrease. The whorl section during been observed that during early ontogeny, the oldest ontogeny maintains the normal tendency of Vascoceras specimens of V. durandi can show fe eble vestiges of of becoming more compressed, although its height is umbilical tubercles, and therefore exhibit a relatively always by far surpassed by its width. These specimens similar appearance to the representatives of V.charoni. show the typical suture lines of this genus, with broad This new species, however, presents much more robust and high firstlateral saddles, and wide and low second and persistent ornamentation, broader whorl section, lateral saddles. slightly more complex suture lines and lower strati­ Discussion: Few species of Vascoceras present a graphical distribution than V.durandi. relatively wide whorl section and a marked ornamen­ Among the Cenomanian members of Vascoceras, tation. Most of the more ornamented fo rms, such as a tendency towards the progressive loss of umbilical V.di arti anum (o 'ORBIGNY 1850), V.gam ai CH OF­ tubercles and ribs has been observed. V. charoni con­ FAT, 1898, V.ba rcoicense CHOFFAT, 1898, V. cauvini stitutes an atypical fo rm within this process, because CHUDEAU, 1909, V. bufbosum (REYMENT 1954a) its morphology becomes similar to that of the genus and V.obscurum BARBER, 1957, show elevated and Fagesia PERVIN�IERE, 1907. This new species may compressed whorl sections. On the other hand, the therefore be one of the taxa from which the earliest wider and more compressed taxa, such as V. harttii representatives of this group originated. An example 210 Barroso-Barcenilla & Goy

of these first members of Fagesia is F. catinus (MAN­ 1967 vascoceras barcoicense CHOFFAT - POLSAK, P· 136, TELL 1822), whose lowest records were reported by pi. 83, fig.2. CoBBAN et al. (1989) from the N juddii Zone of the 1981 Nigericeras barcoicense (CHOFFAT) - AMARD et al., p. 54, pl. 4, fig. 16 a-b. USA. This biostratigraphic unit is stratigraphically 1984 Va scoceras mundae CHOFFAT - MELENDEZ-HEVIA, close to the V.gam ai Subzone and the S. (])subc on­ p. 88, pl. 3, fig. 3a-b [only]. cili atus Zone of Spain, from whose levels V. charoni 1986 vascoceras barcoicense CHOFFAT - BERTHOU et al., has been collected. p. 70, pl. 4, figs. 1-3. Distribution: This new species occurs in the lev­ 1989 vascoceras barcoicense CHOFFAT exile COBBAN et al., els 10, 11 and, possibly, 12 ofBARROSO-BARCENILLA p. 47, pl. 87, figs. q-s; pi. 89, figs. m-g. (2006) of the upper part of the V.gam ai Subzone and ? 1994 Vascoceras durandi (THOMAS & PERON) CHANCELLOR et al., p. 48, pl. 10, figs. 1-2 [only]. the lower part of the (]) subconcili atus Zone of S. 2001 Vascoceras barcoicense CHOFFAT - CALLAPEZ & Tamajon, in the Guadarrama Area of the Iberian FERREIRA, p. 67, pl. 6, figs. 3-5; pl. 7, figs. 1-2, text­ Trough, Spain. figs. 20.1-3, 20.5. 2006 Vascoceras barcoicense CHOFFAT - BARROSO­ Vascoceras barcoicense CHOFFAT, 1898 BARCENILLA, p. 228, pl. 30, figs. a-i. (Plate 2, figs.G-K) Type: The holotype designated by CHOFFAT (1898, pl. 17, fig. 1 a-c) was destroyed. Among the paratypes conserved in the 1898 Ammonites (?vascoceras) barcoicens is CHOFFAT, p. 67, SGP, BERTHOU et al. ( 1986) selected as neotype the non­ pl. 17, fig. 1;pl. 22, fig. 35 [only]. illustrated specimen 831 of CHOFFAT, from Almoxarife, ? 1907 vascoceras cf. barcoicensis CHOFFAT - PERVIN�IERE, Portugal. p. 335. Material: 6 specimens; TA- R-106, TA-R-634, TA-R-652, ? 1914 vascoceras barcoicensis CHOFFAT - EcK, P· 203, pl. 14, TA-R-99, TA-S-567 and TA -S-572. fig. !.

Dimensions: D H (%) E (%) 0 (%) Range 993-534 415-234 (48-42) 571-304 (58-46) 228-129 (26-19) Mean 737 325 (44) 384 (52) 173 (24)

Description: Involute specimens with subellipti­ The V.cf. barcoicensis of PERVIN�IERE (1907, cal or subrectangular whorl section whose greatest p. 335) was considered a strongly ribbed Vttscoceras breadth is close to the umbilical margins. They have an durandi (THOMAS & PERON 1889) by BERTHOU et arched ventral region, flator slightly convex flanksand al. (1986) and CHANCELLOR et al. (1994). After quite small umbilici with rounded margins. Their or­ studying in the MNHN this specimen, figured by namentation appears on the ventral region and the CHANCELLOR et al. (1994, pl. 10, figs. 1-2), it has outer half of the flanks, and is composed of several been observed that its features, such as the whorl sec­ slightly prorsiradiate ribs, although certain specimens tion, which is only slightly wider than higher, the um­ can also have some small umbilical tubercles on the in­ bilical width, which only reaches 20 % of the diameter ner whorls, which disappear quite quickly. The adult of the shell, and the ornamentation are closer to those body chamber presents a scaphitoid aspect and a broad of V. barcoicense than to those of V. durandi. The aperture. Theyshow variable suture lines with proxi­ V.ba rcoicense of EcK (1914) was considered a doubt­ mate septa. ful V.durandi by BERTHOU et al. (1986). This speci­ Discussion: CHOFFAT ( 1898) described this spe­ men has a relatively depressed whorl section, as indi­ cies and included it in his new genus Vttscoceras. In cated by BERTHOU et al. (1986), and in our opinion contrast, SCHNEEGANS (1943) integrated V. barcoi­ their fe atures may correspond to a wide V.ba rcoicense. cense in Paravascoceras FuRON, 1935, and AMARD et The Paravascoceras c£ barcoicensis of SCHNEEGANS al. (1981) included it in Nigericeras ScHNEEGANS, ( 1943) was considered an evolute V.ca uvini CHUDEAU, 1943. However, as shown by CHOFFAT (1898, pl. 12, 1909, by ScHOBEL (1975). This opinion was not re­ fig. 35), the suture lines of this species show the typical jected by BERTHOU et al. (1986, p. 72), who, never­ pattern of Vttscoceras. theless, maintained it in the synonymy of V. barcoi- The ammonite genus Vascoceras CHOFFAT, 1898 211

cense. The same ammonite was later attributed to and with strong ribbing in the later stages belong to V. (P aravascoceras) cauvini by MEISTER et al. ( 1992), CHUDEAU 's species. Similarly, the presence of orna­ and with some reservations to Pseudovascoceras ni­ mentation in some of the smallest specimens presented geriense (Wo ODS 1911) by ZABORSKI (1996). This in this paper clearly distinguishes V.ba rcoicense from doubtful specimen, recently studied at the MNHN, V. cauvini, but places the fo rmer species close to presents 25-30 weak ribs per whorl but reaches a rela­ V.ga m ai. However, the narrowness ofthe umbilici, the tively high umbilical width, which ranges 26-29 % of robustness of the ribbing and the early lack of orna­ the diameter of the shell, depending on the measured mentation near the umbilical area of these immature flank.For this reason, it is difficultto consider this am­ specimens of V.ba rcoicense distinguish them from the monite a member of V.ba rcoicense. Finally, the N bar­ more evolute members of the genus. V. barcoicense coicense of AMARD et al. ( 1981) was considered a pos­ loses its small umbilical tubercles earlier than its strong sible Paravascoceras cauvini by ZABORSKI ( 1996). ribbing, whereas V.gam ai loses its weak ribbing quick­ In our opinion, this specimen of the MNHN seems to ly and maintains its strong umbilical tubercles until present the typical fe atures of V.ba rcoicense. advanced ontogeny. On the basis of these features, it is In view of the difficulties encountered in distin­ advisable to maintain the specific status of V. barcoi­ guishing V. barcoicense from V. gamai and from cense. Phylogenetically, V. barcoicense is also close to V.cauvini, BERTHOU et al. ( 1975, 1986) emphasised V.gam ai, from which it seems to have evolved by weak­ that the main problem in identifying and establishing ening and early disappearance of the umbilical tuber­ the taxonomic status of the the fo rmer species lies in cles, and increasing involution of the whorls. the ignorance of the morphology and ornamentation Distribution: V. barcoicense occurs in the upper of the inner whorls of its holotype. BERTHOU et al. Cenomanian and lower Turonian ofPortugal, Croatia, ( 1986) also remarked that if this missing holotype in Algeria, Spain, the USA, and, possibly, Tunisia and its early ontogeny had presented some umbilical tu­ Egypt. In the Iberian Trough, it has been identifiedin bercles, or a nearly smooth external surface, it would the upper part of the V.gam ai Subzone and the lower be difficultto maintain the specific separation between part of the 5. (] ) subconcili atus Zone of the Guadar­ V.ba rcoicense and V.gam ai, in the firstcase, or between rama Area. V.ba rcoicense and V.cauvini, in the second. Regarding the separation between V. barcoicense Vilscoceras durandi (THOMAS & PERON, 1889) and V.cauvini, BERTHOU et al. (1986) stated that the (Plate 3, figs.A-G; Plate 4, figs.A-B; Text-figure 6 A) fo rmer species has compressed and involute morphol­ ? aff. 1889 Pa chydiscus aff.peramplus (MANTELL) - THOMAS ogy and, apparently, lacks umbilical tubercles during & PERON in PERON, p. 25, pl. 18, figs. 1-2. early ontogeny. However, they maintained this specific 1889 Pa chydiscus durandi THOMAS & PERON in distinction because V.cauvini shows a subtriangular PERON, p. 27, pl. 18, figs. 5-8. and more involute whorl section and a strong ribbing 1896 Pa chydiscusdurandiTHOMAS & PERON- PERON, at maturity. ZABORSKI ( 1996) separated both species p. 44, pl. 4, fig.1; pl. 5, fig. 1; pl. 17, fig.5. on the basis of the reduced ornamentation of V.ba r­ 1898 rascoceras douvillei CHOFFAT, P· 59, pl. 10, fig. 6; coicense during maturity. Apart from these features, pl. ll, figs. 2-5; pl. 21,figs. 13-15. the different ontogenetic tendencies followed by aff. 1898 Va scoceras sp. aff.douvil lei CHOFFAT, p. 60, pl. 10, fig.3; pl. 21, fig.16. V.ba rcoicense and V. cauvini also make it possible to 1903 Va scoceras durandi (THOMAS & PERON) - differentiate between these two species. Whereas the PERVINQ:!:!IERE,PP · 98-99. fo rmer loses ornamentation during ontogeny, as can 1903 Vascoceras douvilfei CHOFFAT - PERVINQ:!:!IERE, be observed in the specimens presented in this paper, p. 99. the latter develops stronger ribbing during growth. 1907 Va scoceras durandi (THOMAS & PERON) - Therefore, the notable morphologic differences and PERVINQ:!:!IERE, p. 332, pl. 21, fig. 1 a-b, text­ the dissimilar ontogenetic development of V. barcoi­ fig. 125. cense and V.cauvini support the taxonomic separation 1914 Va scoceras durandi (THOMAS & PERON) - EcK, p. 202. of these taxa. Specimens with small umbilici whose or­ ? 1915 Vascoceras durandi (THOMAS & PERON) - namentation disappears during ontogeny correspond GRECO, p. 210, pl. 18, figs. 2-3. to CHOFFAT 's species, whereas specimens with an al­ ? aff. 1920 f/ascoceras sp. nov. ex. aff.adonense CHOFFAT - most smooth surface in the earlier stages of growth BasE, p. 214, pl. 14, fig. 4;pl. 17, fig. 2, text-fig. 2. 212 Barroso-Barcenilla & Goy

1928 Vascoceras sp. DouvrLLE, p. 15, pl. 1, fig.6. 1984 Paravascoceras sp. 4 MELENDEZ-HEVIA, p. 102, 1934 Vascoceras durandi (THOMAS & PERON) - pl. 8, figs. 1 a-c, 2 [only]. FARAUD, pi. 20, fig.8. 1986 Vtzscoceras durandi (PERON) - BERTHOU et al., p. 1940 Vascoceras (Pachyvascoceras) durandi (THOMAS & 72, pl. 4, figs.4-9; pl. 6, figs. 1-6. PERON) - FARAUD, p. 43, pi. 2, figs. 1-2; pi. 8, 1989 Vtzscocerasdurandi (THOMAS & PERON) - LUG ER fig. !. & GROSCHKE, p. 376, pl. 43, figs. 1-2, text-fig.8 cf. 1960 Discovascoceras cf. douvillei (CHOFFAT) - a. WI EDMANN, pp. 712, 721, 723. 1991 Vtzscoceras durandi (PERON) - KENNEDY & 1960 Vtzscoceras (Pachyvascoceras) durandi (THOMAS & SrMMONS, p. 138, pl. 5 a-b. PERON) - WI EDMANN, pp. 714, 720. 1994 Vascoceras durandi (THOMAS & PERON) - cf.1960 Vtzscoceras (Pachyvascoceras) cf.durandi (THOMAS CHANCELLOR et al., p. 48, pl. ?2, fig.1; pl. 10, figs. & PERON) - WrEDMANN, p. 714. 3-4; pl. 11, figs. 1-2; pl. 12, figs. 1-3; pl. 13, figs. cf. 1964 Discovascoceras cf. douvillei (CHOFFAT) - 3-4; pl. ? 14, figs. 2, 5 [only]. WI EDMANN, pp. 111, 115-116. ? aff. 1996 Va scoceras (Paravascoceras) aff. durandi (THOMAS 1964 Vtzscoceras (Pachyvascoceras) durandi (THOMAS & & PERON) - MEISTER & ABDALLAH, P· 10, pl. 4 PERON) - WrEDMANN, pp. 111, 115. fig. 1; pl. 5, fig. 2, text-fig. 5c. cf.1964 Vtzscoceras (Pachyvascoceras) cf.durandi (THOMAS 2001 Vtzscoceras douvillei CHOFFAT - CALLAPEZ & & PERON) - WI EDMANN, P· 111. fERREIRA, p. 69, pl. 9, figs.3-8; pi. 10, fig.1, text­ ? aff1965a Vascoceras (Pachyvascoceras) aff.douvi llei CHOFFAT figs. 20.8-11, ?21.1, 21.2. - durandi (THOMAS & PERON) - COLLIGNON, 2001 Va scoceras durandi (THOMAS & PERON) - p. 43, pi. 394, fig. 1676. CALLAPEZ & FERREIRA, p. 78, pl. 11, figs. 3-4; 1969 Vtzscoceras durandi (THOMAS & PERON) - pl. 12, figs. 1-2,text-figs. 21.3-6. FREUND & RAAB, p. 30, text-fig.6 h-i. 2005 Va scoceras durandi (THOMAS & PERON) - ? 1969 Vtzscoceras harttifOrme CHOFFAT - FREUND & MEISTER & ABDALLAH, P· 135, pl. 14, fig. 1 RAAB, p. 31, text-fig. 6j. [only]. cf. 1969 Vtzscoceras cf.V. adonense CHOFFAT - FREUND & cf.2006 Vtzscoceras cf. durandi (THOMAS & PERON) -EL RAAB, p. 32, pi. 5, fig. 1, text-fig. 7a-b. QoT, p. 118, pi. 25, fig.4; pl. 26, fig. 1 a-b. ? aff. 1973 Vtzscoceras sp. aff durandi (THOMAS & PERON) ­ 2006 Va scoceras durandi (THOMAS & PERON) - MATSUMOTO, p. 29, pJ. 8, fig.1, text-fig. 1. BARROSO-BARCENILLA, p. 232, pl. 30, fig. j; pl. 1975 Pachyvascoceras durandi (THOMAS & PERON) - 31, figs. a-d; pl. 32, figs. a-e, text-fig.68. WIEDMANN, P· 142. Type: The lectotype designated by CHANCELLOR et al. ? aff. 1978 Vtzscoceras sp. aff V.durandi (THOMAS & PERON) ( 1994) is the specimen 1901-11 from Aln Settara, Tunisia, - MATSUMOTO & MuRAMOTO, p. 281, text­ described by PERON (1889, p. 25, pl. 18, fig. 5), and currently fig. 1 a-b. held in the MNHN with the number R7937. 1979 Va scoceras cf. douvillei CHOFFAT - WrEDMANN, Material: 17 specimens; CA-R-28, CB-R-659, CC-R-194, p. 199. CC-R-199, CC-R-203, CC-R-206, CC-R-225, MS-4, TA -R- 1984 Pa ra vascoceras sp. 2 MELENDEZ-HEVIA, p. 102, 101, TA -R-418, TA-R-621, TA-R-624, TA-S-432, TA-S-568, pi. 7, fig.2 a-c [only]. TA-S-573, TA-S-575 and TA-S-85.

Dimensions: D H (%) E(%) 0 (%) Range 1310-676 500-282 (45-34) 770-392 (63-44) 485-210 (38-28) Mean 1000 396 (40) 545 (55) 337 (34)

Description: Slightly involute specimens, almost more acute in the approach to maturity. Their firstlat­ without ornamentation, and with quite variable, de­ eral saddles are large, whereas the remaining saddles pressed and rounded, subtriangular or subogival whorl are relatively small. section, which is always wider than higher. They have Discussion: CHOFFAT (1898) emphasised the wide umbilici with steep and subvertical walls and strong resemblance between Vascoceras durandi blunt umbilical angles. Juveniles may present 6-8 (THOMAS & PERON 1889) and his new species Vas­ small umbilical tubercles per whorl, disappearing coceras douvillei, and based their taxonomic separation quickly during ontogeny. They show a tendency to on the smaller size and the strong ventral ribbing adopt a subtriangular whorl section that becomes of the latter. PERVIN�IERE (1907) included V.dou - The ammonite genus Vascoceras CHOFFAT, 1898 213

villei in the synonymy of V.dur andi. BERTHOU et al. ofFREUND & RAAB (1969) also present a morphol­ ( 1986) remarked that many specimens with interme­ ogy quite similar to that of V. durandi, as noted by diate fe atures can be observed, and concluded that BERTHOU et al. ( 1986) and CHANCELLOR et al. V. douvillei should be considered as a synonym of ( 1994), who attributed these specimens to the species V. durandi. The interpretations made by PERVIN­ ofTHOMAS & PERON. The V.cf. V.adonense ofFRE­ �J:ERE(1 907) and BERTHOU et al. (1986) concern­ UND & RAAB ( 1969) are closer to V.durandi than the ing V.durandi and V.douvi llei have been confirmedby V.harttiforme of the same authors. The V.durandi of the Spanish records of both taxa, and the latter has GRECO (1915), the V.(Pa chyvascoceras) aff.dou villei­ also been considered here as a synonym of the fo rmer. durandi of COLLIGNON ( 1965a), and the V.sp. aff. BERTHOU et al. ( 1986) regarded the separation durandi of MATSUMOTO (1973) and MATSUMOTO between V.durandi and Vascoceras amieirense CHOF­ & MuRAMOTO (1978) have a whorl section and a re­ FAT, 1898, as being based only on the involution and duced umbilical width that seem to separate these the contraction ofthe body chamber, and they consid­ specimens from the species ofTHOMAS & PERON, as ered that the second taxon is conspecific with the first stated by BERTHOU et al. (1986) and CHANCELLOR one. Likewise, these authors highlighted the morpho­ et al. (1994). The V.sp. nov. ex. aff. adonense ofBosE logic proximity between V. durandi and Vascoceras ( 1920) has a notable whorl breadth and lacks umbili­ harttii (HYATT 1870). They stated that the Paravas­ cal tubercles, which seems to indicate that it is slightly coceras aff. hartii of CHANCELLOR (1982, p. 101, closer to V.durandi than to Vascoceras ga mai CH OF­ fig. 34)shows a high umbilical width, with dimensions FAT, 1898. Finally, the V.(P.) aff. durandi ofMEISTER close to those of V.dou villei, and that this specimen, & ABDALLAH ( 1996) exhibits a very persistent rib­ which could represent an intermediate fo rm between bing that is notably different from that of the species the species of HYATT and of THOMAS & PERON, ofTHOMAS & PERON. seems to provide evidence of the close phylogenetic Studying the phylogeny of V.durandi, BERTHOU relationship between these two taxa. BERTHOU et al. et al. (1986) and CHANCELLOR et al. (1994) indicat­ ( 1986) also suggested that V.durandi could be a pos­ ed that the juvenile specimen of V.harttii previously sible synonym of V.harttii. Nevertheless, they main­ cited by CHANCELLOR (1982, p. 118) presents a cri­ tained the specific division of both taxa, on the basis carinate ventral region, similar to that of the members of the narrower umbilical width of V.harttii . The new of the species assigned by CoLLIGNON ( 1957, 1965b) data fo r V.am ieirense and V.harttii presented in this to his new genus Discovascoceras. These authors also paper support their specific separation, as explained indicated that this morphologic similarity could re­ below. veal a certain phylogenetic relationship between the Some compressed juvenile specimens of V.dur an­ species V. durandi and the bicarinate taxon 1homa­ di, such as that presented by PERVINQ£IERE ( 1907), sites madagascariensis CoLLIGNON, 1965a. In this can be hard to differentiate from those of V.gamai, es­ regard, it should be noted that within the morpho­ pecially when their umbilical tubercles are maintained. logical variability of Vascoceras CHOFFAT, 1898, as However, the members of V.ga mai develop a less wide can be observed in V.ga mai, particularly in its variety whorl section and a more evolute coiling, and present subtriangularis, and in V.durandi, forms exist with a striking and strong umbilical tubercles. On the other relatively sharp triangular whorl section. Therefore, hand, Vascoceras cauvini CHUDEAU, 1909, shows a the small ammonite cited by CHANCELLOR ( 1982) more compressed whorl section, a less wide umbilical may be a member of Vascoceras with a more or less width and a different adult ornamentation, whereas triangular whorl section. Likewise, it has been noted Vascoceras angermanni BOSE, 1920, and Vascoceras that in the Iberian Trough V.durandi follows an on­ crassum FURON, 1935, have narrower umbilici than togenetic development similar to that of V.gamai. V.dur andi. This coincidence and the existence of specimens with The Pa ch ydiscus aff.pera mplus (MANTELL 1822) intermediate fe atures, such as ammonite TA-S-568 of THOMAS & PERON (1889) is a poorly preserved (Pl. 3, figs.A-B ), could also indicate that the origin of ammonite that, as indicated by CHANCELLOR et al. V.durandi might be in V.ga mai. ( 1994 ), could be a compressed and highly ornamented Distribution: V. durandi has been collected from V. durandi. The Vascoceras harttiforme CHOFFAT, the upper Cenomanian and lower Turonian of Tuni­ 1898, and V.cf. Vascoceras adonense CHOFFAT, 1898, sia, Algeria, Portugal, Egypt, France, Spain, Israel, 214 Barroso-Barcenilla & Goy

Oman and, probably, Mexico, Madagascar and Japan. Vascoceras cauvini CHUDEAU, 1909 In the Iberian Trough, it is common in the Inner Cas­ (Plate 4, fig.C) tilian Platform, within the subcon cili atus and S. (]) 1909 Vascoceras cauvini CHUDEAU, p. 68, pis. 1-2; pi. 3, C. (C.)quaa si zones. Within the latter biostratigraph­ figs. 1-2,4. ic unit, the morphological variability of V. durandi 1921 Thomasites cauvini (CHUDEAU) - CHUDEAU, p. 463, increases progressively. The stratigraphical distribu­ fig. 1. tion of this species observed in Spain is equivalent to 1933 Vtzscoceras cauvini CHUDEAU - FuRON, p. 268, pi. 9, that determined by CHOFFAT (1898) and BERTHOU fig.17. FURON, et al. ( 1986) in Portugal. In fact, the lowest Iberian ? 1935 Va scoceras (Paracanthoceras) chevalieri p. 59, pi. 4, fig.1 a-b. records are among the oldest ones in the world as­ ? 1935 Va scoceras (Paracanthoceras) chudeaui FURON, p. 60, signed to V.du randi. pi. 4, fig.2. BERTHOU et al. (1986) noticed a progressive in­ 1935 Vtzscoceras (Paravascoceras) cauvini CHuoEAU - crease in the whorl breadth from the earlier to the later FURON, p. 60, pi. 5, fig. 1. representatives of V.du randi. They also indicated that 1935 Va scoceras (Paravascoceras) cauvini CHUDEAU var. the amplitude of the umbilical tubercles seems to de­ semiglabra FuRON, p. 61, pi. 4, fig.3. - SCHNEEGANS, pend on the palaeogeographical origin of the speci­ 1943 Pa ra vascoceras cauvini (CHUDEAU) p. 128, pi. 5, fig.2, text-fig 9. mens, because members of V. durandi collected in . 1943 Pa ra vascoceras cauvini (CHUDEAU) var. evofuta Portugal show thicker tubercles than those obtained SCHNEEGANS, p. 130, pi. 8, fig. 2, text-figs. 10-11. in North Africa. From the earlier to the later Spanish 1943 Pa ra vascoceras cauvini (CHUDEAU) var. injlata specimens of V.durandi presented in this paper it can SCHNEEGANS, P· 131. be observed that the whorl breadth increases gradually aff. 1957 Pa ravascoceras aff. cauvini (CHUDEAU) - BARBER, as the size and persistence of the vestigial tubercles de­ p. 37, pi. 14, figs.2-3; pi. 32, figs.8-9. creases. In fact, the umbilical tubercles seem to be 1969 Pa ravascoceras cauvini (CHUDEAU) - FREUND & RAAB, p. 20, pi. 3. figs.1-3. primitive structures in V.durandi, probably evolved ? 1969 Pa ravascoceras tavense (FARAUD) - FREUND & RAAB, from V.gam ai, and their thickness and persistence are p. 23, pi. 2, fig. 9, text-fig. 5 e-g. possibly related to the stage of evolution of the speci­ 1975 Pa ravascoceras cauvini (CHUDEAU) - SCHOBEL, men considered. For this reason the tubercles reduce p. 119, pi. 4, figs.1, ?2, 3; pi. 5, figs.1-4. progressively, until they disappear in the later mem­ 1982 Pa ravascoceras cauvini (CHUDEAU) - CARRETERO­ bers of V.durandi. These observations are similar to MORENO, p. 253. LuG those of BERTHOU et al. (1986), because the Portu­ 1989 Vascoceras cauvini CHUDEAU - ER & GROSCHKE, guese specimens of V.du randi, which are older, show p. 374, pi. 40, figs. 3, 6, 8-9; pi. 41, figs. 1-4; pi. 42, fig. 1, text-figs. 6 g-h, 8 c. stronger umbilical tubercles than the African ones, cf. 1989 Vtzscoceras cf. cauvini CHUDEAU - LuGER & which are earlier. GROSCHKE, p. 376, pi. 42, fig. 2; pi. 43, fig.3, text-figs. 6 f. 8 b. 1989 Vtzscoceras cauvini CHUDEAU - KENNEDY et al., p. 82, figs.9 g, 20 c-g. 1990 Va scoceras cauvini CHUDEAU - ZABORSKI, figs.8 a-b, 12-15.

A B

Te xt-fig. 6. Suture lines. A, Vtzscoceras durandi (THOMAS & PERON 1889), TA-S-575, from the Ch ojfoticeras ( Chojfoticeras) quaasi Zone ofTamaj6n. B, Vtzscoceras kossmati CHOFFAT, 1898, TA-R-624, from the Sp athites (Ingrideffa) maffadaeSubzone ofTamaj6n. Both are x 3/2. The ammonite genus Va scoceras CHOFFAT, 1898 215

1992 Vascoceras (Paravascoceras) cauvini CHUDEAU - 2006 Va scoceras cauvini CHUDEAU- EL QoT, p. 1 17, pi. 25, MEISTER et al., p. 71, pi. 4, fig.6; pi. 5, figs. 1-3; pi. 6, figs.2- 3, 5. figs. 1- 4; pi. ?7, figs. 1- 2, 4-5, text-fig. 16. 2006 Va scoceras cauvtm CHUDEAU BARROSO­ 1996 Paravascoceras cauvini (CHUDEAU) - ZABORSKI, BARCENILLA, p. 238, pi. 33, fig.a. p. 65, figs.2- 8. Type: The lecrotype designated by KENNEDY et al. (1989) 1996 vascoceras cauvini CHUDEAU- AMEDRO et al., p. 215, is a specimen of CHUDEAU (1909), from Gj adjidouma, Niger, fig. 18. and held in the MNHN with the number R52488. 2002 vascoceras cauvini CHUDEAU - EL-HEDENY, p. 406, Material: 1 specimen; PU-S-312. figs.4 b-c, 7f.

Dimensions: D H (%) E (%) 0(%) 690 316 (46) �245 (58-46) 228-129 (26-19)

Description: Moderately involute and small of the whorl section and of the ornamentation. LUG ER specimen, with a compressed and subelliptical whorl & GROSCHKE (1989) considered Vt:zscoceras rumeaui section. It presents rounded ventral region, flat or a slightly different species from V.cauvi ni. MEISTER slightly convex flanks,and subvertical umbilical walls et al. (1992) differentiated in V. cauvini the fo rms with blunt margins. Its visible surface is smooth and its "cauvini s.s."; "forme lisse", which is smooth or nearly suture lines have a reduced number of elements with smooth and includes the "nigeriensis" and "ta vense" clearly divided saddles. styles; "forme comprimee", which is compressed and Discussion: CHUDEAU (1909) described this involute; and "forme crassum", which is globose. They species in comparison with Vt:zscoceras durandi (THO­ also recognised a transitional fo rm between the "forme MAS & PERON 1889). He stated that the firstwhorls crassum" and the species Vascoceras (Paravascoceras) of both taxa are very similar, and emphasised that, in proprium (REYMENT 1954b). AMEDRO et al. (1996) contrast to V.ca uvini, none of the closer forms, such as considered the taxon P.rumeaui to be conspecificwith Vascoceras douvillei CHOFFAT, 1898, and Vt:zscoceras V.ca uvini, whereas ZABORSKI ( 1996) included with­ amieirensis CHOFFAT, 1898, retain their ornamenta­ in the synonymy of this species V.(P.) chevalieri, V.(P.) tion into the maturity. CHUDEAU (1921) asserted chudeaui, V.(P.) cauvini var. semigla bra, P.cauvini var. that Vt:zscoceras nigeriense WooDS, 1911, may possibly evoluta, P.ca uvini var. injlata and Vt:zscocerasdep ressum be a synonym of V. cauvini whilst, in contrast, BARBER, 1957. He also stated that P. rumeaui and REYMENT ( 1954b) affirmed that the second species Vt:zscoceras costellatum AMARD et al., 1981, could be could be a synonym of the firstone. SCHOBEL (1975) regional variants of V cauvini. The same author only considered Vt:zscoceras (Pachyvascoceras) crassum considered as conspecific with V. cauvini the taxa FURON, 1935, Vt:zscoceras (Paracanthoceras) chevalieri assigned to the forms "cauvini s.s.", "lisse" and FURON, 1935, Vt:zscoceras (Paravascoceras) chudeaui "comprimee" by MEISTER et al. (1992). FuRON, 1935, Vt:zscoceras (Paravascoceras) cauvini var. The relatively compressed whorl section and the semiglabra FuRON, 1935, Nigericeras jacqueti notably strong adult ribbing of this species make its SCHNEEGANS, 1943, Pa ravascoceras crassum var. but­ identificationeasier. Nevertheless, at least V.(P. ) chev­ lata SCHNEEGANS, 1943, Pa ra vascoceras cauvini var. alieri and V.(P.) chudeaui seem to be indistinguishable evoluta SCHNEEGANS, 1943, Pa ravascoceras cauvini from V.ca uvini. With regard to the specificseparation var. inflata ScHNEEGANS, 1943, Broggiiceras hum­ between V. barcoicense and V cauvini it should be boldti BENAVIDEs-CAcERES, 1956, Broggiiceras emphasised, as indicated by ZABORSKI ( 1996) and ofssoni BENAVIDES-CACERES, 1956, and Pa ravas­ detailed above, that the fo rmer lacks the mature rib­ coceras rumeaui COLLIGNON, 1957, as synonyms of bing ofthe latter. The Paravascoceras tavense (FARAUD V.cauvi ni. 1940) ofFREUND & RAAB (1969) should be assigned BERTHOU et al. ( 1986) indicated that V.cauvini to V.cauvini, as observed by SCHOBEL (1975) and could be a synonym of Vt:zscoceras barcoicense CHOF­ BERTHOU et al. (1986), because even the smallest of FAT, 1898, but maintained their specificseparation on them has no ornamentation. The V. cauvini of the basis of their different ontogenetic development KENNEDY et al. (1989) were considered as a possible 216 Barroso-Barcenilla & Goy

V.barcoicense exile COBBAN et al., 1989, by ZABOR­ vascocerasamieirense CHOFFAT, 1898 SKI ( 1996). These ammonites lack ornamentation, de­ (Plate 4, figs.D-E) spite being immature specimens, and seem to corre­ 1898 Vascoceras amieirensis CHOFFAT, p. 61, pi. 12, spond to V.cauvini. Regarding its phylogeny, ZABOR­ figs. 1-2; pl. 13, figs. 1-2; pi. 21, figs. 17-21. SKI ( 1996) indicated that V.cauvini could be derived cf. 1914 rascoceras sp. cf. amieirensis CHOFFAT - EcK, p. 201. from N gadeni by progressive loss of the juvenile orna­ cf. 1957 rascoceras (Discovascoceras) cf.amieirense CHOFFAT - mentation, simplification ofthe suture lines and de­ COLLIGNON, P· 124. velopment of the adult ribbing. cf. 1960 Discovascoceras cf. amieirense (CHOFFAT) Distribution: This species has been identified in WIEDMANN, pp. 712,714, 721. cf. 1964 Discovascoceras cf. amieirense (CHOFFAT) the upper Cenomanian of Niger, the Sudan, Israel, WIEDMANN, pp. 111, 115. Spain, Egypt, the USA, Nigeria and Algeria, in bios­ cf. 1969 rascoceras cf. V amieirense CHOFFAT - FREUND & tratigraphic units equivalent to the M. geslinianum RAAB, p. 32, text-fig.6 k-1. and N juddii standard zones. However, ZABORSKI ? 1972 rascoceras sp. COBBAN & SCOTT, P· 83, pi. 36, figs.3-4, (1996) observed that the ammonites collected from text-fig. 41 a. the M geslinianum Zone of Israel and classified as cf. 1975 Discovascoceras cf. amieirense (CHOFFAT) V.ca uvini by LEWY et al. (1984) present many simi­ WrEDMANN, p. 142. 1984 Pa ra vascoceras sp. 2 MELENDEZ-HEVIA, p. 102, pi. 7, larities with the members of the genus Nigericeras fig. 1 a-c [only]. SCHNEEGANS, 1943, and that they may be transition­ 1984 Pa ravascoceras sp. 3 MELENDEZ-HEVIA, p. 102, pi. 6, al specimens. In the Iberian Trough, the ammonite fig.3. presented here has been obtained at the top of the S. 2006 rascoceras amieirense CHOFFAT BARROSO­ (])subcon ciliatus Zone of the North-Ebro Area, and BARCENILLA, p. 242, pJ. 33, figs.b-e. other specimens were recorded by CARRETERO­ Type: The holotype is specimen 825 ofthe SGP, the original MORENO (1982) in undetermined levels of the Guad­ ofCHOFFAT (1898, pJ. 12, fig. 1 a-b; pi. 13, fig. 1; pi. 21, figs. 17, arrama Area. 20), from the lower Turonian of Amieira, Portugal. Material: 6 specimens; CA-R-598, CB-R-542, CC-S-211, TA-S-570, TA-S-571 and TA-S-578.

Dimensions: D H (%) E(%) 0 (%) Range 1400-968 649-380 (46-39) 550-435 (50-37) 382-258 (35-26) Mean 1105 461 (42) 489 (45) 322 (30)

Description: Relatively compressed ammonites durandi (THOMAS & PERON 1889), maintained their with subtriangular whorl section and without orna­ specific separation. On the basis of morphometric mentation. They have rounded ventral area, slightly analysis, BERTHOU et al. (1986), and other authors arched and convergent flanks and not too wide um­ such as CHANCELLOR et al. (1994), considered bilici with subvertical walls and rounded margins. V.amieirense as a synonym of V.durandi. Discussion: CHOFFAT (1898) highlighted the Nevertheless, as can be seen in the diagram by subtriangular whorl section and the lack of umbilical BERTHOU et al. (1986, p. 61, fig. 5), the dimensions of tubercles as the main fe atures of his new species Vtzs­ the holotype of V.amieirense are at the morphological coceras amieirensis. He also stated that it presents extreme of the group designated V.durandi by these higher whorls and broader umbilici than Vtzscoceras authors. We therefore agree with the interpretation harttiformis CHOFFAT, 1898. CoLLIGNON (1957) made by BERTHOU et al. ( 1975) concerning the com­ included V.amieirense within his new genus Discovas­ parison of the specimens of V.amieirense and V.d u­ coceras, although this interpretation was rejected by randi. Accordingly, we have maintained the specific COBBAN & SCOTT (1972). The latter authors also division of V amieirense and of V durandi, as they dif­ noticed that the specimens they had classified as Vtzs­ fer in their whorl section and in their umbilical width. coceras sp. have features in common with V.amieirense. This conclusion is supported by the fact that V. amiei­ BERTHOU et al. (1975), although recognising the rense occurs in higher levels than V durandi in the great similarity between V.amieirense and Vtzscoceras Iberian Trough, an observation also made by BERTHOU The ammonite genus Vascoceras CHOFFAT, 1898 217

et al. ( 1975) in Portugal, who remarked that the strati­ 1937 Va scoceras hartii (HYATT) - MAURY, p. 247, pi. 22, graphical distribution of V douvillei coincides with figs. 1-2. that of V. subconciliatus, and is lower than that of cf. 1940 Vascoceras (Pachyvascoceras) cf. hartii (HYATT) - FARAUD, p. 48, pi. 4, fig. 2; pi. 5, fig. 2; pi. 9, V.amieirense, V.kossmati. which occurs with figs. 1-2. With regard to the specimens classified as Vas­ cf. 1960 Vascoceras (Pachyvascoceras) cf. hartii (HYATT) - coceras sp. by COBBAN & SCOTT (1972), BERTHOU WIEDMANN, P· 712. et al. (1986) stated that these ammonites probably be­ cf. 1964 1/ascoceras (Pachyvascoceras) cf. hartii (HYATT) - long to V. durandi, and CHANCELLOR et al. (1994) WIEDMANN, p. 111. added that they may be compressed variants of the 1969 Pachyvascoceras harttii (HYATT) - DE OLIVE!RA & same species. In our opinion, although the specimen B RITO, p. 220, pi. 2, figs. 1-2 (only). 1977 Paravascoceras hartti (HYATT) - CHANCELLOR et of CoBBAN & SCOTT (1972, pl. 36, figs. 1-2) is too al., p. 96, fig. 20. deformed to permit a precise taxonomic classification, 1982 Paravascoceras hartti (HYATT) - CHANCELLOR, in general, the ammonites classified by these authors p. 98, figs.2 b, 29-33. as Vascoceras sp. exhibit several fe atures close to those ? aff. 1982 Paravascoceras aff. hartti (HYATT) - CHANCELLOR, of V.amieirense. p. I 0 1, fig.34. Phylogenetically, V. amieirense seems to have 1984 Paravascoceras hartti (HYATT) - MELENDEZ­ originated from some of the latest members of V.du­ HEVIA, p. 99, pi. 4, figs.1 a-b, 2 a-b, 3 a-b. 1985 1/ascoceras (Paravascoceras) harttii (HYATT) - ra ndi, by modifications in the width and the contour HOWARTH, P· 100, fig. 25. of the whorl section. In fact, PERVIN�IERE ( 1907) 1989 1/ascoceras hartti (HYATT) - COBBAN et al., p. 49, had already suggested the possible existence of transi­ figs.49, 91 a-d, g-k. tional specimens between both species, and LUG ER & ? 1989 Fagesia superstes var. lev is RENZ - MEISTER, p. 37, GROSCHKE (1989) highlighted the fact that their pi. 16, fig. 2, text-fig. 26. representatives of V.durandi have several fe atures close ? 1992 Vascoceras gr. globosum (REYMENT) ou Fa gesia to those of the members of V.amieirense. Likewise, the sp. CouRVILLE, pi. 9, fig. 1. V.durandi and V.cf. amieirense of FREUND & RAAB 1996 Vascoceras hamii (HYATT) - ZABORSKI, p. 81, figs. 58, 60. (1969) seem to be examples of these ammonites with 2006 Vascoceras harttii (HYATT) - EL QoT, p. 118, pi. 26, intermediate fe atures. fig. 2a-b. Distribution: V.amieirense has been obtained in 2006 Va scoceras harttii (HYATT) BARROSO­ the lower Turonian of Portugal, Spain and, possibly, BARCENILLA, p. 244, pi. 34, figs. a-f; pi. 35, Egypt, Algeria, Israel and the USA. In the Iberian figs.a-c. Trough, it has been fo und in the upper part of the Type: Although HYATT ( 1870) did not provide illustrations quaasi Zone and in the lower part of the of this species, WHITE (1887, pi. 19, figs. 1-2; pi. 20, fig. 3) C. (C.} S. (I. } illustrated a specimen that possibly corresponds to one of the malladaeSubzone ofthe North-Ebro and Guadarrama original syntypes. According to article 33a of the ICZN, the areas. terminations -ii and -i are allowed alternatives, which is why the specific names of harttii and hartti are both valid. In the present Vtzscoceras harttii (HYATT 1870) work, however, we have used the first one,as it coincides with the (Plate 5, figs.A-E) original denomination ofHYATT (1870). Material: 24 specimens; CA-S-509, CB-R-552, CB-R-855, 1870 Ceratites harttii HYATT, p. 386. CC-R-11, CC-R-197, CC-R-228, CC-R-8, CC-S-226, CG-R- 1875 Buchiceras harttii (HYATT) - HYATT, p. 370. 533, CS-R-591, CS-R-592, TA-R-412, TA-R-426, TA-R-574, 1887 Ammonites (Buchiceras) harttii (HYATT) - WH ITE, TA-R-617, TA-R-645, TA -R-656, TA-R-750, TA- R-751, TA-S- p. 226, pi. 19, figs. 1-2; pi. 20, fig. 3. 431, TA-S-608, TA-S-648, TA-S-649 and TE-S-587. 1903 Vascoceras hartti (HYATT) - HYATT, p. 103, pi. 14, fig. 16.

Dimensions: D H (%) E(%) 0 (%) Range 1125-516 421-200 (48-31) 795-420 (87-61) 335-155 (38-25) Mean 794 303 (39) 587 (75) 245 (31) 218 Barroso-Barcenilla & Goy

Description: Slightly involute and globose cadi­ from V. harttiforme. KENNEDY et al. ( 1987) stated cones with depressed half moon-shaped whorl section. that the species of HYATT is very similar to Vascoceras Medium-sized, very deep and steep umbilici with sub­ globosum (REYMENT 1954b) and to certain specimens vertical or oblique walls and sharp margins. Ornamen­ of Vascoceraspro prium (REYMENT 1954b ), but asserted tation is reduced to about twenty non-persistent and that V.harttii shows a more evolute coiling and a steep broad ventral juvenile plications per whorl, irregularly umbilical wall. ZABORSKI ( 1996), however, included distributed and slightly orientated towards the body within the species ofHYATT all the specimens classified chamber. The first whorls can present a subtriangular as Fagesia superstes var. levis RENZ, 1982, by MEISTER section, and the adult spires almost hide the rest of (1989, p. 37, pl. 16, fig. 2, text-fig. 26) and as V.gr. these ammonites. Simple and rounded suture lines, globosum or Fagesia sp. by CouRVILLE (1992, pl. 9, each one with fo ur wide lateral saddles. fig. 1). Discussion: After being described by HYATT Regarding the separation between V.harttii and ( 1870), the species Ceratites harttii was included by V. harttiforme, it can be deduced from the Spanish CHOFFAT (1898) in his new genus Vascoceras. HYATT material that the distributions of these species are dif­ ( 1903) indicated that this species is related to the taxa fe rent, as the fo rmer occurs in higher levels than the Vascoceras harttiformis CHOFFAT, 1898, Vascoceras latter. These facts are consistent with those observed amieirensis CHOFFAT, 1898, and Vascoceras kossmati by other authors, such as CHANCELLOR (1982) in CHOFFAT, 1898. PERVIN�IERE (1907) observed Mexico, and support the specificdivision of these two great similarities between Vascoceras harttii and Vas­ taxa. The high depression of the whorl section of coceras durandi (THOMAS & PERON 1889), although V.harttii enables its differentiation from other fo rms he added that both species can be distinguished by the of the genus with higher whorls, such as V.durandi presence of umbilical tubercles and the complexity of and Vascoceras ellipticum BARBER, 1957. The umbili­ the suture lines of the latter. FuRON ( 1935) described cal width can also be considered an important feature a species with a similar morphology, named Vascoceras in order to differentiate V. harttii, being wider than (Pa chyvascoceras) crassum. BARBER ( 1957) remarked that of V.glo bosum, and narrower than that of V.du­ that V.harttii, as well as V.harttiforme, is more globose ra ndi. In addition to the important differences be­ than Vascoceras nigeriense Wooos, 1911, but less in­ tween V.harttii and V.gl obosum stated by KENNEDY volute, more compressed and with steeper umbilical et al. ( 1987), the Spanish specimens ofthe first species walls than Vascoceras robustum BARBER, 1957. show a more triangular and less rounded whorl sec­ CHANCELLOR ( 1982) stated that the species V.harttii tion and a flatterumbilical wall than those of the sec­ is close to some globose taxa with a narrow umbilicus ond one. Likewise, due to its globose aspect it is easy of the same genus, such as V.angerm anni BosE, 1920, to confuse V.harttii with some members of the genus and V.carteri (BARBER 1957). He classified a speci­ Fagesia, although this species follows the typical on­ men from Mexico, with an umbilical width reaching togenetic development of the group Vascoceras, as approximately 57 % of the diameter of the shell, as emphasised by ZABORSKI ( 1996), and exhibits a sim­ Paravascoceras aff. harttii. This ammonite was consid­ pler outline of the septa. The F. superstes var. levis of ered by BERTHOU et al. ( 1986) as a possible specimen MEISTER (1989) and the V.gr.gl obosum or Fagesia sp. of V.durandi, and excluded by HoWARTH (1985) and of CoURVILLE (1992) show important similarities ZABORSKI (1996) from the synonymy of V.harttii. with V.harttii. Likewise, the F. superstes var. levis of CHANCELLOR (1982) also indicated that the V.hartii MEISTER (1989) seems to lack the umbilical tubercles ofWILLARD ( 1966, pl. 41, fig.2; pl. 42, fig. 1) is close of the taxon of RENZ. Phylogenetically, V.harttii was to the V.amieirensis of LISSON (1908). This ammo­ apparently originated fr om the more depressed, invo­ nite was subsequently considered by BERTHOU et al. lute and globose members of V.durandi, such as the ( 1986) as a possible specimen of Vascoceras cauvini specimen CC-R-199 (Plate 4, figs.A-B). CHUDEAU, 1909. BENGTSON (1983) and HOWARTH Distribution: Cited in the lower Turonian of (1985) assigned the specimens of Pa chyvascoceras Brazil, Mexico, Spain, Angola, the USA, Nigeria, harttii of DE 0LIVEIRA & BRITO (1969, p. 220, pl. 2, Egypt and, possibly, France. The members of V.harttii fig. 3) and REYMENT & TAIT (1972) to the genus from the Iberian Trough have been located in the low­ Fagesia PERVIN�IERE, 1907. HOWARTH (1985) er part of the S. (1 ) malladae Subzone of the Central also emphasised that V.harttii is only slightly different Sector. The ammonite genus Vascoceras CHOFFAT, 1898 219

Jiascoceras kossmati CHOFFAT, 1898 1975 Pa chyvascoceras (Pachyvascoceras) harttiforme (Place 5, figs. F-H; Te xt-figure6 B) (CHOFFAT) - WIEDMANN, p. 142. cf. 1979 Pachyvascoceras cf. harttiforme (CHOFFAT) - 1898 Vascoceras harttiformis CHOFFAT, p. 61, pi. 12, fig. 3; WI EDMANN, P· 203. pi. 13, figs.3-6; pi. 21, figs.22-24. 1984 Paravascoceras cf. carteri (BARBER) - MELENDEZ­ 1898 Vascoceras kossmati CHOFFAT, p. 63, pi. 13, figs.8-9; HEVIA, p. 100, pi. 5, figs. 1 a-c, 2 a-b. pi. 14, figs. 1-2; pi. 21, figs.26-27. 1986 Vascoceras kossmati CHOFFAT - BERTHOU et al., 1914 Vascoceras kossmati CHOFFAT - EcK, pp. 182, 202. p. 76, pi. 5, figs. 1 -9. 1934 Vascoceras harttiformis CHOFFAT - FARAUD, p. 9, 1992 Pa ravascoceras kossmati (CHOFFAT) - THOMEL, fig. 6. p. 225, pi. 119, figs. 2-3. aff. 1935 V. aff. harttiformis CHOFFAT - K.ARRENBERG, ? aff. 1992 Pa ravascoceras aff.kossmati (CHOFFAT) - THOMEL, p. 140, pi. 31, fig.11. p. 225, pl. 118, figs. 1-2. 1960 Vascoceras (Pachyvascoceras) harttiforme (CHOFFAT) ? 2005 Va scoceras durandi (THOMAS & PERON)- MEISTER - WIEDMANN, pp. 712, 723. & ABDALLAH, p. 135, pi. 26, fig. 1; pi. 27, fig.1. cf. 1960 Vascoceras (Pachyvascoceras) cf. harttiforme 2006 Vascoceras kossmati CHOFFAT - BARROSO­ (CHOFFAT) - WIEDMANN, pp. 712,714. BARCENILLA, p. 248, pi. 35, figs. d-f, text-fig. 70. 1960 Vascoceras (Pachyvascoceras) crassum FuRON - Type: Among the syncypes, BERTHOU ec al. (1986) WIED.MANN,P· 712. designated as lectotype the specimen 827-1 of CHOFFAT ( 1898, 1960 Va scoceras (Pachyvascoceras) kossmati (CHOFFAT) - pi. 13, fig. 8; pi. 14, fig. 1 ), collected from Costa d 'Arnes, WIEDMANN, p. 723. Portugal, and held in the SGP. In spite of recognising that page 1964 Vascoceras (Pachyvascoceras) harttiJorme (CHOFFAT) priority corresponds eo Va scoceras harttiformis CHOFFAT, 1898, - WIED.MANN,pp. Ill' 116. BERTHOU et al. (1975) used the name Vascoceras kossmati cf. 1964 Vascoceras (Pachyvascoceras) cf. harttijorme CHOFFAT, 1898, for this species, afterobserving that the types of (CHOFFAT) - WIED.MANN, P· 111. this taxon are in a better state of preservation. 1964 Vascoceras (Pachyvascoceras) crassum FURON - Material: 5 specimens; CB-R-21, CB-R-554, PU-R-455, WIEDMANN, P· Ill. TA-R-644 and TE-S-588. 1964 Vascoceras (Pachyvascoceras) kossmati (CHOFFAT) - WIEDMANN, P· 116.

Dimensions: D H (%) E (%) 0 (%) Range 1193-685 546-334 (47-41) 840-500 (73-64) 225-141 (24-14) Mean 896 426 (45) 619 (69) 175 (19)

Description: Globose and highly involute am­ of the types of V. kossmati described by CHOFFAT monites with depressed whorl section. They show (1898), and demonstrated that this species cannot be rounded or slightly subtriangular ventral region, and distinguished from V. harttiforme. BERTHOU et al. deep and narrow umbilici with vertical walls and sharp (1986) considered that, among the species of the ge­ margins. Juvenile specimens may show weak and nus Vascoceras CHOFFAT, 1898, occurring in Portugal, slightly prorsiradiate ribs that become stronger as they V.kossmati is the most globose and involute and has reach the ventral region. Adult specimens lack orna­ the deepest umbilici. They specifiedthat the cadicones mentation, and have a body chamber that gradually of Vascoceras angermanni BosE, 1920, which to­ reduces in width and becomes slightly less involute. tally lacks ornamentation, and of Vascoceras carteri They exhibit suture lines with short saddles. (BARBER 1957) present an even more spherical aspect Discussion: CH OFF AT (1898) justifiedestablish­ than those of V.kossmati. ing V.hartti.formis as a new species on the grounds Besides the impossibility of determining a mor­ that it exhibited wider whorl section and narrower phological limit between V. kossmati and V. hartti­ umbilici than his new taxon Vascoceras amieirensis. forme, in the Iberian Trough it has been observed that FREUND & RAAB ( 1969) emphasised the strong re­ the geographical and stratigraphical distributions of semblance between V.hartti.forme, Vascoceras durandi both taxa are coincident. Due to the subspherical (THOMAS & PERON 1889) and Vascoceras harttii shape, the small umbilical width and the highly de­ (HYATT 1870). BERTHOU et al. (1975) remarked that pressed whorl section of V.kossmati, this species can they were not able to observe the umbilical tubercles easily be differentiated from other related members of 220 Barroso-Barcenilla & Goy

the genus, such as V.durandi and V.harttii. Neverthe­ V. kossmati. Although all these taxa had already been less, there are certain doubtful specimens, such as the cited, with the logical exception of the new species, ammonite classifiedas V.durandi by MEISTER & AB­ V. barcoicense, V. durandi, V. amieirense and V. koss­ DALLAH (2005, pl. 26, fig. l; pl. 27, fig. 1). Although mati had never previously been properly classifiednor this specimen exhibits seemingly eroded umbilical ar­ illustrated for this palaeogeographical region. During eas, it shows notably wide and involute whorls and the developed revision, the presence of specimens at­ presents dimensions close to those of V.kossmati. A tributable to V. silvanense and, possibly, V. globosum close phylogenetic relationship might be established and V. crassum has been observed in the collections of between the latest and most involute members of the UT and the UCM. On the other hand, the speci­ V.harttii and the earliest and the most evolute speci­ mens of these research centres regarded as V. triangu­ mens of V.kossmati. lare, a taxon that may be a synonym of Cho..lfaticeras Distribution: Lower Turonian of Portugal, ( Cho..lfaticeras) douvillei, should be assigned to the Egypt, France and Spain. In the Iberian Trough, in the Pseudotissotiidae. Therefore, it can be assumed that upper part of the S. (I ) malladae Subzone of the Inner the Vascoceratidae are represented, among others, by Castilian Platform. the genus Vascoceras in the Iberian Trough, where the species V.ga mai, V. charoni, V. barcoicense, V. cauvini, V. durandi, V.amieirense, V.harttii, V.kossmati, V.sil­ Conclusions vanense and, probably, V. globosum and V. crassum In the present work, new specimens from the Ibe­ have been properly identified. rian Trough have been described and assigned to the Furthermore, the distribution of the new mem­ species Vascoceras ga mai, V. charoni, V. barcoicense, bers of the family Vascoceratidae presented here has V. cauvini, V. durandi, V. amieirense, V. harttii and been determined (Text-fig. 7). The records of the ge-

Iberian Trough, Spain. Stratigraphic ranges, phylogenetic BARROSO-BARCENILLA et al. (2009) relationships and evolutionary phases

Grffatiams (Grff atiams) q£nSi

• � !p:thite> (J�) u juilii szlxrniliatus � Q) c. c. Jv11ciaxHrE 1v11oiaxx:ras � g:sliniamm gslinicnun

Te xt-fig.7. Stratigraphical ranges of the species of Vt:tscoceras identifiedin this work, and their inferred phylogenetic relationships and evolutionary phases in the Iberian Tr ough. The ammonite genus Vt:tscoceras CHOFFAT, 1898 221

nus Vascoceras extend from the V.gamai to S. (!.) mal­ of Vascoceras without umbilical tubercles, such as the ladae subzones. Among its species, Vascocerasga mai is typical specimens of V. durandi, V. amieirense, V. hart­ the first taxon of the genus, and of the family, estab­ tii and V. kossmati. Their most abundant members also lished in this palaeogeographical region. Many speci­ exhibit depressed sections and involute coilings, and mens ofthis species have been obtained in the V.ga mai their morphologies can be considered as "evolved". Subzone. In the upper part of the V. gamai Subzone, These two evolutionary phases seem not only to coin­ and the lower part of the S. (!.) subconciliatus Zone, cide with the evolutionary intervals fo llowed by the V. charoni and V. barcoicense have been identified.Al­ family in other palaeogeographical regions, but also though these two species show partially coincident by any other groups, as indicated by BARROSO-BAR­ vertical distributions, that of the latter is slightly high­ CENILLA & GOY (2005). er than that of the fo rmer. V. durandi has been ob­ tained in the upper part of the S. (!.) subconciliatus Acknowledgements Zone and the lower part of the C. (C.) quaasi Zone. The occurrence of V. cauvini seems to be restricted to The first author appreciates the help offered by Dr. A. the top of the S. (!.)subcon ciliatus Zone. V. amieirense Liebau and Dr. H. Schulz in the Universitat Ttibingen, Germany, and V. harttii have been obtained from the upper part by Pro£ Dr. M. Brandao and Mrs.]. Pachucka in the Museu do Instituto Geol6gico e Mineiro de Lisboa, Portugal, and by Dr. A. of the C. (C.) quaasi Zone and the lower part of the S. . Cornee, Dr. D. Merle, Dr. ].M. Pacaud and Pro£ Dr. JP. Saint (I ) malladae Subzone. Finally, within the upper part Martin in the Museum National d 'Histoire Naturelle de Paris, of the S. (I.) malladae Subzone, V. kossmati has been France, in order to study the types from the upper Cenomanian identified. This is the most recent species of Vascoceras and lower Turonian which are held in their respective institutions. collected in the Iberian Tr ough. He also expresses his gratitude to Prof. Dr. WJ Kennedy from In terms of the phylogeny of the group, one pos­ the Oxford University Museum ofNatural History, UK, and Mr. F. Amedro from Calais, France, for their valuable remarks sible evolutionary lineage joining V.ga mai, V. duran­ concerning the identificationof some of the specimens presented di, V. harttii and V. kossmati has been identifiedwithin here. We want to thank Pro£ Dr. M. Segura Redondo from the the genus Vascoceras (Text-fig. 7). It becomes progres­ Universidad de Alcala de Henares, Spain, for his help lending us sively more involute and depressed, with less orna­ some cephalopods belonging to his own collection, Mr. C. Alonso mented fo rms. Likewise, the representatives of V. cha­ Recio, for his photographs, Mrs. M.I. Martinez Martfnez and roni and of V. barcoicense seem to be phylogenetically Miss M.M. Martinez Martinez, fo r their translations, and Mr. I. close to the wider or more involute specimens of Melendez Hevia for his work carried out during his final project V. ga mai with persistent ornamentation, from which for graduation. We also recognize the work of Dr. ]. Lehmann from the Universitat Bremen, Germany, who has notably they could respectively be derived. The members of improved the original content of the present paper. This study V. amieirense possibly originated from some of the has been carried out within the projects 01!1003/2001 of the more compressed specimens of V. durandi. Comunidad de Madrid, CGL2005-01765/BTE and CGL2008- After observing the distribution of the family in 03112/BTE of the Ministerio de Educaci6n y Ciencia, and the Iberian Trough, two main evolutionary phases can PAIOS-0204-1312 of the Junta de Castilla-La Mancha, and be distinguished. These are characterised by the suc­ financed bythese institutions of Spain and by the Social Fund of cessive dominance of the "primitive" Vascoceras and of the European Union. the "evolved" Vascoceras (Text-fig.7). The first one has been identifiedin the V.gamai Subzone and the S. (!.) References subconciliatus Zone. During this phase the family is AMARD, B., COLLIGNON, M. & ROMAN,]. (1981): Etude strati­ mainly represented by upper Cenomanian species of graphique et paleontologique du Cretace Superieur et Vascoceras with umbilical tubercles, at least during ear­ Paleocene du Tinrhert-W et Tadema.it-E (Sahara algerien). ly ontogeny, such as V. gamai, V. charoni, V. barcoi­ - Doe. Lab. Geol. Fac. Sci. Lyon, 6: 15-173. cense and the first specimens of V. durandi. Their most AMEDRO, F.,B ussoN, G. & CoRNEE, A. (1996): Revision des numerous members also show compressed sections ammonites du Cenomanien superieur et du Turonien in­ and evolute coilings, and their morphologies can be fe rieur du Tinrhert (Sahara algerien): implications bios­ considered as "primitive". The second phase has been tratigraphiques. - Bull. Mus. Nat. d 'Hist. Natur., SCr. IV, identified in the C. (C.) quaasi Zone and the S. (I.) 18: 179-232. malladae Subzone. During this phase the family is BARBER, W. (1957): Lower Turonian Ammonites from north­ predominantly represented by lower Turonian species eastern Nigeria. - Geol. Surv. Nigeria, Bull., 26: 1-87. 222 Barroso-Barcenilla & Goy

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Plate 1

1/ascoceras gamai CHOFFAT, 1898 - A, B, C, D, E, F, G, H, I, J, K, L, ventral, lateral and apertural (complete), ventral, lateral and apertural (without a fragment of body chamber), ventral, lateral and apertural (without body chamber), and ventral, lateral and apertural (without body chamber and a fragment of phragmocone) views of CA-S-151, from the Vascoceras gamai Subzone of Condemios, x 3/4. Theammonite genus Vascoceras CHOFFAT, 1898 227 228 Barroso-Barcenilla & Goy

Plate 2

Vascoceras charoni sp. nov. - A, B, lateral and apertural views ofMS-3, holotype, from the Vascoceras gamai Subzone ofTamaj6n, x I. C, D, E, F, lateral, apertural, lateral and ventral views ofTA-S-556, paratype, from the Vascocerasgamai Subzone ofTamaj6n, x 1. Vascoceras barcoicense CHOFFAT, 1898 - G,H, lateral and ventral views ofTA- R-1 06, from the Vascoceras gamai Subzone ofTamaj6n, x 1. I,J, K, apertural, lateral and ventral views ofTA-R-634, from an unknown level ofTamaj6n, x 1. The ammonitegenus Vascoceras CHOFFAT, 1898 229 230 Barroso-Barcenilla & Goy

Plate 3

Vascoceras durandi (THOMAS & PERON 1889) - A, B. lateral and apertural views of MS-4, from the Sp athites {jeanrogericeras) subconciliatus Zone ofTamaj6n. C, D, ventral and lateral views ofTA-S-568, from the Sp athites {jeanrogericeras) subconciliatus Zone ofTamaj6n. E, F, G, apertural, lateral and ventral views ofTA-S-573, from the Chojfaticeras ( Chojfaticeras) quaasi Zone ofTamaj6n, X 5/6. The ammonite genus Vczscoceras CHOFFAT, 1898 23 1 232 Barroso-Barcenilla & Goy

Plate 4

Vtl Scoceras durandi (THOMAS & PERON, 1889) - A, B, ventral and lateral views of CC-R-199, from an unknown level of Cantalojas, X S/6. Vtl Scoceras cauvini CH UD EAU, 1909 - C, lateral view ofPU-S-312, from the Sp athites (jeanrogericeras) subconciliatus Zone ofPuentedey, X S/6.

Va scoceras amieirense CHOFFAT, 1898 - D, E, apertural and lateral views of CA-R-598, from the Sp athites (Ingridella) malladae Subzone ofCondemios, x S/6. Theammonite genus Vtzscoceras CHOFFAT, 1898 233 234 Barroso-Barcenilla & Goy

Plate 5 Vascoceras harttii (HYATT 1870) -A, B, apertural and lateral views of CC-S-226, from the Sp athites (Ingridella) malladae Subzone of Cantalojas, x 1. C, D, E, ventral, lateral and apertural views of CB-R-855, from the Sp athites (Ingridella ) malladae Subzone of Condemios, x 1. Va scoceras kossmati CHOFFAT, 1898- F, G, H, apertural, lateral andventral views ofPU-R-455, from the Sp athites (Ingridella) malladat Zone of Puentedey, x 1. The ammonite genus Vascoceras CHOFFAT, 1898 235 This paper or parts thereof may not be reproduced in any form without permission from the publishers

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