Profil 16: 95-106, 2 Figs.; Stuttgart 1999

The Lower Kimmeridgian Section at Gallipuen. Ammonite Biochronostratigraphy and Ecostratigraphic Remarks

L. MOLINER, ZARAGOZA & F. OLORIZ, GRANADA*

Abstract The precise analysis of ammonite ranges in the Kimmeridgian section at Gallipu_n permits the recognition of the Platynota, Hypselocyclum and Divisum Zones of the Lower Kimmeridgian, but the presence of the lowermost Upper Kimmeridgian in the area is doubtful. From bottom to top, Ihe Platynota Zone is subdivided in the „Orthosphinctes interval", the middle Desmoides Subzone (with the enayi and desmoides biohorizons), and the upper Guilherandense Subzone (with the Guilherandense (G-1) interval and Ihe guilherandense biohorizon). The first appearance of crussoliceratids is the easier way to recognise Ihe Hypselocyclum/Divisum Zone boundary. In general, the correlation with standard biochronostratigraphic units proposed for Submediterranean Europe is acceptable, but Ihe identification of minor differences deserves future research. The composition of fossil assemblages of macroinvertebrates, including megabenthos, fits well with the assumption of changes in ecospace forced by sea-level fluctuations in low-energy environments. The analysis of changes in the composition of ammonite assemblages complements that of the benthic : pelagic ratio, hence improving ecostratigraphic interpretations at the zone level. The epitome of ammonites correlates with a significant decrease in Ataxioceratinae (the highest specialised group), increasing haploceratids and especially aspidoceratids (less specialised), and the epibole of benthic fauna The most balanced ammonite assemblages occur in the Divisum Zone, in accordance with the maximum flooding that affected the area.

1 INTRODUCTION sedimentology, stratigraphy, tectonics and regional geology. However, the analysis of fossil assem- The knowledge on the Kimmeridgian from the blages in their stratigraphic context, i.e. the basis eastern Iberian Chain improved significantly during for ecostratigraphic interpretations, is poorly de- seventies with contributions such as those made by veloped on Kimmeridgian materials. Preliminary Bulard et al. (1971), Felgueroso Coppel & Ramirez data from the Platynota Zone were reported by del Pozo (1971), Canerot (1974), El Khoudary OI6riz et al. (1988) on the basis of precise ammon- (1974) and G6mez (1979). Concerning precise ite biostratigraphy. Finkel (1992) showed the com- Kimmeridgian ammonite biostratigraphy and the position of ammonite assemblages recovered from composition of macroinvertebrate fossil assem- Kimmeridgian outcrops northeastern from Gal- blages (megabenthos included) in the eastern Iberian lipuen. The aim of the present paper is to show Chain, the valuable contribution by Geyer & some general traits of the ecostratigraphic analysis Pelleduhn (1979) was pioneer. Later, data on the applied to, the Lower Kimmeridgian in the Gallipuen Kimmeridgian in the region increased and diversi- area, supported by precise ammonite bio- fied from eighties (Moliner, 1983; Atrops & Mel§- stratigraphy. ndez, 1984; Moliner & OI6riz, 1984; OI6riz et al., 1988; Aurell, 1990; Aurell & Melendez, 1990; Melendez et al., 1990; Finkel, 1992; Bddenas et 2 LOCATION AND GEOLOGICAL al., 1993; Salas & Casas, 1993; Atrops et al., SETTING 1997; Aurell et al., 1997; Badenas, 1997; Badenas & Aurell, 1977; and G6mez & Goy, 1997; among Upper Jurassic outcrops in Las Umbri'as are lo- others). Hence, a wide spectrum exists of updated cated close to the village of Alcorisa, province of information about Kimmeridgian palaeontology, Teruel, between coordinates 0°25'6" -0°24'56"

* Addresses of the authors: Prof. Dr. Federico Oloriz S£ez, Depto. Estratigraffa y Paleontologfa, Fac. Ciencias, Univ. Granada. 18002, Granada, Spain, e-mail: [email protected] -- L. Moliner, ENTECSA, Poligono Argualas, nave 52-A, c/ Argualas s/n, 50012 Zaragoza, Spain, e-mail: [email protected] 96

Fig. 1: Location and geologic sketch of the Gallipuen area. and 40°52'39"- 40°52'46" in the sheet of ridgian, we investigated sections showing the Calanda, no 494 (scale 1:50.000) in the National Platynota, Hypselocyclum and Divisum Zones. Topographic Map (Fig. 1). The Upper Jurassic shows a W-E trend about 3 Km in the northern limb 3 THE LOWER KIMMERIDGIAN of the Cerro de las Umbr'as, from the Gallipuen reservoir to the neighbouring of Alcorisa. The SECTION AT GALLIPUEN Gallipuen sections are located north from the dam In the Gallipuen area, the bottom of the Upper and were favoured by quarry works in Kimme- Jurassic sections studied (Fig. 2) shows ca. 70cm ridgian limestones during reservoir construction. thick Oxfordian deposits composed of nodular pe- The outcropping of Mesozoic materials in Las loid-oolitic-algal limestones indicating low rates of Umbr'as shows an anticline with the Upper Jurassic deposition and probably hiatuses during late Oxfor- underlying the uncomformable Lower Cretaceous. dian times. However, these bioclastic wackestones, Favourable sections of Upper Jurassic deposits and locally packstones, with frequent sponge spi- exist towards the western extreme of this cules and irregular bedding (horizons 5 to 20cm structure. In the Gallipuen sector, the Upper Juras- thick), show energy conditions sporadically higher sic is composed of a Small section belonging to the that background conditions assumed for the typical Oxfordian, and a better-developed Kimmeridgian sponge limestones of the Yatova Mb. of the Chelva section including a thin stratigraphic interval at the Fm. Identifiable ammonites are rare, but Trimar- top with possibility to shows lowermost Upper ginites sp., as well as Epipeltoceras treptense from Kimmeridgian deposits. An erosive unconformity neighbouring areas, were reported by Moliner separates Kimmeridgian deposits from the over- (1983), thus indicating the possibility for the local lying upper Lower Cretaceous sandstones. There- recognition of the upper Bimammatum Zone. As re- fore, there is low possibility, if anyone, to recog- cognised by Moliner (1983), the Trimarginites sp. nise Upper Jurassic deposits younger than the up- mentioned could indicates even the lower Planula permost Lower or the lowermost Upper Kimme- Zone according to data in Karve-Corvinus (1966), ridgian in the Gallipuen area. Within the Kimme- Wierzbowski (1978), Garcia-Hernandez et al. (1979) and Sequeiros & Oloriz (1979).

Fig. 2: Lithological columns showing the Kimmeridgian section at Gallipuen. Pie-diagrams give the composition of fossil assemblages of macroinvertebrates (megabenthos included) at the ammonite-zone level, based on the material recovered from particular sections. Note different magnification for the particular sections analysed. Vertical ruling for indeterminate Divisum or Acanthicum Zones. 97

THE LOWER KIMMERIDGIAN AT GALLIPUEN -MTG.3-26 INVERTEBRATES

111 3 conglomerate

silty dolostone sample size ammonitea, I „_ataxioceratins bioclastic limestone belemnitids AMMONITINA & nautilidr -/\other ammonites j limestone brachiopods • •Jl • y haploceratins //•* • * • * • marly limestone gastropods + • bivalves indeterminate ammonites marl ~r T

0 nodular oolitic limestone

i ' i n-MTns--s J -MTG.2-40

EZE INVERTEBRATES f i

i r

-MTG.2-20

AMMONITINA

>• • \ / • • *\ * • • • • 4• •

r-:

11.77m

I Om -MTG.2-1 98

Overlying the condensed deposits are 70cm thick densely packed fossil-remains and bioturbation. greyish, fine siliciclastics, which include ferrugi- The Lower Kimmeridgian section at Gallipuen is nous ooids at the base (lowermost 20cm) and some 51,40m thick and belongs to the lower part of the intercalations of centimetres-thick marly lime- Ritmita Calcarea de Loriguilla Fm. (Calanda Mb.; stones, but no sandy horizons (Fig. 2). These de- Melendez et al., 1990). These deposits have been posits belong to comparatively basinward records interpreted to represent deposition in an exten- of the Margas de Sot de Chera Fm., the deposition sional lagoon (Aurell & Melendez, 1990), under of which was related to pulses of basin instability high rate of carbonate production within photic (Aurell et al., 1997; Gomez & Goy, 1997; Badenas depths in outer ramp settings (Aurell, 1990). & Aurell, 1997). These pulses were related to a Badenas (1997) typified the Kimmeridgian ramp as tectonic phase widely identifiable in the eastern- a flat-topped ramp, and calculated a maximum most Iberia (Salas & Casas, 1993), as well as in depth higher than 40-60m but lower than 100m. southern and western Iberia (Marques et al., 1991; Rodr'guez-Tovar, 1993; Oloriz & Rodriguez- Tovar, 1998), and northern Africa (Atrops & Ben- 4 PALEONTOLOGICAL STUDY est, 1986; Benzaggagh, 1988; arques et al., 1991, Previous researches in the Gallipuen sections among others). Ammonites are scarce and not well were reported by Bulard (1972) and Marin et al. preserved in the sections studied at Gallipuen, but (1977), who identified the Lower Kimmeridgian and Atrops & Melendez (1984) identified the lowermost indicated the Gallipuen section as a fossiliferous Platynota Zone (Orthosphinctes Subzone) with site in the geological map of Calanda (geological- Orthosphinctes (Orthosphinctes) polygyratus sheet 494, 1:50.000, IGME), respectively. Moliner (REINECKE), Orthosph. (Orthosphinctes) polygyra- (1983) made the first detailed analysis on Upper tus (REINECKE) morph. pseudopolyplocoides GEYER, Jurassic ammonites in the Alcorisa-Berge sector and Nebrodites hospes (NEUMAYR) at the top of (Gallipuen sections included) providing the major slightly thicker sections of the Margas de Sot de traits of the composition of fossil assemblages. At- Chera Fm. in the area. North-westwards, in the rops & Melendez (1984) included data from the Moyuela section, Aurell et al. (1997) recovered Gallipuen area in their revision of Upper Jurassic Sutneria galar (OPEL) and Orthosphinctes castroi ammonites from the Calanda-Berge sector. Oloriz & (CHOFFAT) revealing deposits belonging to the Moliner (1984) reported the precise ranges of am- upper Planula Zone. However, Aurell (1990) re- monite species in the Platynota Zone at Gallipuen. ported Orthosph. (Orthosphinctes) polygyratus Oloriz et al. (1988) interpreted the composition of (REINECKE) morph. pseudopolyplocoides GEYER from fossil assemblages from the Platynota Zone at Gal- the lower part of the Margas de Sot de Chera Fm. lipuen, and made comparisons with those obtained In general, these data confirm the time-transgres- by the authors in the southern Iberian paleomargin sive character of this formation on shelves of ir- to identify evidences of the Platform Effect (Oloriz, regular bottoms, and/or its hiatal record. In addi- 1985) in epicontinental shelves of the Iberian Pen- tion, they reinforce previous hypotheses inter- insula during the Early Kimmeridgian. In 1998, the preting the Oxfordian/Kimmeridgian boundary (as Gallipuen sector has been included in the "Archivo traditionally considered in Submediterranean Eu- del Patrimonio Aragones de la Diputacion General de rope) to be placed within the Margas de Sot de Aragon". Recent research on fossil assemblages Chera Fm. in the Gallipuen area (Moliner, 1983; recovered from the Gallipuen area allows the im- Atrops & Melendez, 1984; Moliner & Oloriz, 1984; provement of published ammonite biostratigraphy OI6riz et al., 1988; Melendez et al., 1990). (see above), and gives valuable data about general A rather monotonous section of wackestones and trends in the evolution of fossil assemblages of mudstones with variable content in pellets and bio- macroinvertebrates, including megabenthos. clasts, and secondary intercalations of marls, cha- racterises the major part of the Lower Kimme- 4.1 AMMONITES AND BIO- ridgian in the Gallipuen area. Bedding is typically well expressed with decimetres-thick beds showing CHRONOSTRATIGRAPHY ferruginized top-surfaces locally. Marly intercala- The analysis of a total of 530 ammonites col- tions in the Gallipuen area are only significant and lected in situ supports the recognition of the typical in the upper Lower Kimmeridgian, where Platynota, Hypselocyclum and Divisum Zones in the bioclastic packstones and silty dolostones are found Lower Kimmeridgian. Doubts persist about the pos- episodically (Fig. 2), probably showing the in- sible existence of lowermost Upper Kimmeridgian creasing influence of the Montalban-Ejulve High at deposits belonging to the lower part of the Acan- the time. In this upper part of the Kimmeridgian thicum Zone, due to the scanty record of ammonites section, condensed horizons show ferruginization in obtained from the uppermost Kimmeridgian horizons 99 below sandy deposits of the Lower Cretaceous and relatives. Ataxioceratinae are dominant and (Bedulian). The following bio-chronostratigraphic permit the subdivision of the Desmoides Subzone in interpretations are based on the standard proposed a lower enayi horizon and an upper desmoides hori- by the Group Fran_ais du Jurassic as shown by zon, which are correlated with those defined by At- Hantzpergue et al. (1991, see column nr. 3) for rops (1982) in SE France. Thus, the Desmoides Submediterranean Europe, which fits the definition Subzone recognised in the Gallipuen area by Moliner of the Hypselocyclum/Divisum Zone boundary ac- (1983), Moliner & Oloriz (1984), and Atrops & cording to Geyer (1961) and OI6riz (1978). To note Melendez (1984) are correlated with the same sub- that in the schema of Hantzpergue et al. (1991), the zone proposed by Atrops (1982). base of the Divisum Zone in column number 1 was - The enayi horizon. The enay biohorizon identified placed higher than considered by OI6riz (1978). The is 2,50m thick, embraces the range of Ardescia interpretation and correlation of the enayi ATROPS accompanied by large forms of the Hypselocyclum/Divisum Zone boundary in the group of desmoides WEGELE, and yielded Ortho- northeastern Iberian Chain made by Finkel (1992) sphinctes polygyratus (REINECKE), Orthosph. sp., did not take into account the different interpreta- Ardescia sp. aff. Ardescia enayi ATROPS, tion of this zone boundary by previous authors. Ardesc. desmoides quenstedti ATROPS, Ardesc. desmoides desmoides (WEGELE), Ardesc. sp. gr. 4.1.1 PLATYNOTA ZONE Ardesc. desmoides (WEGELE), Ardesc. sp., Litha- The Platynota Zone is based on the range of the cosphinctes sp.1, Lithacosph. sp. gr. Lithacosph. index species in the Gallipuen area where frequent evoiutus (QUENSTEDT), Lithacosph. sp., Ataxio- Ataxioceratinae accompanied it. On the basis of the ceratinae (celtiberic form A), Presimoceras latter, the Platynota Zone is subdivided in subzones hossingense (FISCHER), Sutneria platynota and biohorizons. In the Gallipuen area the Platynota (REINECKE), Pseudowaagenia micropla (OPPEL), section is 14,40m thick. Ataxioceratinae over- Glochiceras nimbatum (OPPEL), and Lingulaticeras whelmingly dominate. Among the remainder am- sp. gr. Ling, lingulatum (QUENSTEDT). Within the monites, Glochiceratidae are more abundant than Desmoides Subzone recognised at Gallipuen, this Aspidoceratidae, Passendorferiinae-"ldoceratids", enayi biohorizon was labelled as "Ardescia sp. and Taramelliceratinae. 1" horizon by Moliner (1983), and named intra- subzone interval, D-1 horizon by Moliner & The lowermost "Orthosphinctes" interval OI6riz (1984). The enayi biohorizon is equivalent This is the lowermost stratigraphic interval to the enayi horizon in Atrops (1982). recognisable within ammonitiferous deposits of the - The desmoides horizon. This biohorizon expands Platynota Zone. The lowermost "Orthosphinctes" from the upper boundary of the enayi horizon to interval is 1,20m thick, yielding incomplete speci- the FAD of Schneidia guilherandense ATROPS. men of Orthosphinctes polygyratus (REINECKE) and This biohorizon, is 2m thick and yielded Ardescia Orthosphinctes sp., together with Presimoceras desmoides desmoides (WEGELE), Ardesc.sp. gr. hossingense (FISCHER), Taramelliceras (Metahaplo- Ardesc. desmoides (WEGELE), Ardesc. sp., Litha- ceras) litocerum (OPPEL), Glochiceras sp. Among cosphinctes sp. 2., Lithacosph. sp., Ataxiocerati- indeterminate Ammonitina one specimen could nae (celtiberic form A), Sutneria platynota belong to Passendorferiinae (see discussion about (REINECKE), Pseudowaagenia sp., Physodoceras the record of Passendorferiinae within the Platy- sp., and Glochiceras sp. From the top of the des- nota Zone in Caracuel et al.; this volume). Moliner moides biohorizon was recovered Ardesc. sp. gr. (1983) proposed the "Orthosphinctes Subzone" for Ardesc. desmoides debelmasi ATROPS and Ard- the lowermost Platynota Zone in the Gallipuen area. escia sp. gr. Ardesc. thieuloyi ATROPS. In the Moliner & Oloriz (1984) labelled this stratigraphic Gallipuen area, the desmoides biohorizon was interval as "Orthosphinctes" interval. The strati- proposed by Moliner (1983) and confirmed by graphic interval analysed is correctable with the Moliner & Oloriz (1984). This biohorizon is cor- Orthosphinctes Subzone defined by Atrops (1982) relatable with the homonymous defined by Atrops and proposed by Atrops & Melendez (1984) for the (1982). lowermost Platynota Zone at the surroundings of Gallipuen The Guilherandense Subzone This stratigraphic interval is 8,70m thick em- The Desmoides Subzone bracing the range of Schneidia guilherandense AT- This subzone is registered in the overlying ROPS in the Gallipuen area. This subzone shows the 4,50m and corresponds to the stratigraphic inter- range of Schneidia with no records of Schneidia lus- val between the first appearance datum (FAD) of sasense ATROPS, although Atrops and Melendez Ardescia spp. and Schneidia guilherandense ATROPS (1984) reported relative forms from the 100

Platynota/Hypselocyclum Zone boundary in the 4.1.2 HYPSELOCYCLUM ZONE northeastern Val de la Piedra section. The major Just above the LAD of Schneidia guilherandense component in ammonite assemblages from the Guil- ATROPS and Sutneria platynota (REINECKE) there is herandense Subzone recognised in Gallipuen are a significant impoverishment in ammonites in the Gallipuen area, a fact previously reported by Ataxioceratinae, the record of others ammonites Moliner (1983) and Atrops & Melendez (1984). being occasional. Based on Ataxioceratinae, two Hence, the precise identification of the Platynota/ biohorizons were identified in the Guilherandense Hypselocyclum Zone boundary is difficult, this being Subzone. reinforced by the scanty record of the youngest - The Guilherandense-1 (G-1) horizon. This strati- Sutneria platynota (REINECKE). Since no Schneidia graphic interval is 3,25m thick ranging from the lussasense ATROPS has been recorded, the FAD of Schneidia to the last appearance datum Platynota/Hypselocyclum Zone boundary inter- (LAD) of Ardescia sp. gr. Ardescia schaireri AT- preted coincides with the LAD of Schneidia. The up- ROPS. Other ammonites recovered are Ardesc. per boundary of the Hypselocyclum Zone is marked sp. gr. Ardesc. desmoides (WEGELE) from the by the FAD of Crussoliceras, Garnierisphinctes, and base, and from higher levels Ardescia sp. gr. Huguenisphinctes, which is considered as a valuable Ardesc. schaireri ATROPS, Ardescia sp., biostratigraphic datum in such a context of scarce Schneidia guilherandense ATROPS, Schn. sp. cf. ammonite record. Hence, the Hypselocyclum Zone at Schn. guilherandense ATROPS, Schn. sp., Lithaco- Gallipuen is 23,55m thick. The Ataxioceratinae are sphinctes pseudoachilles (WEGELE), Lithacosph. dominant, but the record of other Ammonitina in- sp. aff. Lithacosph. subachilles (WEGELE), Litha- creases, especially in aspidoceratids and haplo- cosph. sp. 2, Lithacosph.sp., Ataxioceras stri- ceratids. From the Hypselocyclum Zone have been atellum ? SCHNEID, Ataxioceratinae (celtiberic recovered Ardescia inconditus (FONTANNES), Ard- forms A and B), Presimoceras or Trenerites sp., esc. sp., Ataxioceras sp., Nebrodites sp. cf. Nebro- Sutneria platynota (REIN.), and Glochiceras sp. dites hospes minor (QUENSTEDT), Nebrod. sp., The stratigraphic interval Guilherandense-1 (G-1 Pseudowaagenia micropla (OPPEL), Physodoceras horizon) recognised corresponds to the thieuloyi sp. cf. Physod. wolfi (NEUMAYR), Taramelliceras horizon proposed by Moliner (1983) and to the (Metahaploceras) sp. aff. subnereus (WEGELE), 7a- intra-subzone interval G-1 horizon identified by ram. (Metahaploceras) sp., Glochiceras sp. gr. Moliner & Oloriz (1984) in the lower part of the Gloch. nimbatum (OPPEL), and Gloch. sp. From the Guilherandense Subzone. The precise correlation upper levels of this zone was registered Ataxio- with the thieuloyi horizon in Atrops (1982) is not ceras sp. gr. Atax. hypselocyclum (FONTANNES). conclusive. The scanty ammonite record at Gallipuen im- - The guilherandense horizon. This biohorizon cor- pedes to identify the Hypselocyclum Zone as the responds to the upper 5,45m of the Guilheran- stratigraphic interval that shows the epibole dense Subzone at Gallipuen and ranges from the (acme) of Ataxioceras s.l., as proposed by Geyer LAD of Ardescia schaireri ATROPS to the LAD of (1961) for southern Germany. Without doubt, envi- Schneidia guilherandense ATROPS. The latter co- ronmental factors forced variation of acme zones in incides with the LAD. of Sutneria platynota (REIN- different areas. In addition, the impoverishment in ECKE) that determines the upper boundary of the ammonites during the Hypselocyclum Chron has Platynota Zone in the Gallipuen area. Within the been identified as a common feature in the Hypselo- guilherandense biohorizon have been recovered cyclum Zone recognisable in epicontinental shelves Lithacosphinctes pseudoachilles (WEGELE) from southwards from ' Germany (Atrops, 1982; Ro- the base, and from higher levels Ardescia sp., dri'guez-Tovar, 1993; Oloriz & Rodri'guez-Tovar, Schneidia guilherandense ATROPS, Schn. sp., 1993; Oloriz et al., 1993; Benzzaghag & Atrops, Lithacosphinctes sp., Sutneria platynota (REIN- 1997). ECKE), Physodoceras altenense morph. altenense Moliner (1983) interpreted the subdivision of (D'ORBIGNY), Physodoceras altenense (D'OR- the Hypselocyclum Zone at Gallipuen in a lower BIGNY) morph. circumspinosum (OPPEL), and "Hypselocyclum-I" and an upper "Hypselocyclum- Glochiceras sp. The guilherandense horizon II" section. Ammonites reported from the lower described is slightly thicker than the homonymous Hypselocyclum-I" section were Schneidia sp. aff. biohorizon in Moliner (1983) and Moliner and collignoni ATROPS, Schn. sp., and rare Glochiceras Oloriz (1984). More research is necessary to sp. Later findings of Sutneria platynota (REINECKE) evaluate its precise correlation with the guilher- in these levels determined their reinterpretation as andense horizon proposed by Atrops (1982). belonging to the uppermost Platynota Zone. The stratigraphic interval labelled as "Hypselocyclum- II" by Moliner (1983) is here retained as repre- 101 senting the Hypselocyclum Zone without subzone ceras sp., Sutneria sp. gr. Sutn. cyclodorsata divisions, at least preliminary, and could be cor- (MOESCH) - batalleri GEYER, Aspidoceras sp. gr. related with part of the Hippolytense and Lothari binodum. (OPPEL) - longispinum (SOWERBY), Glochi- Subzones defined by Atrops (1982). ceras sp., and Glochiceras (Lingulaticeras) sp. gr. Due to scarcity in ammonites in the extended Gloch. (Lingulaticeras) crenosum (QUENSTEDT). middle Lower Kimmeridgian section studied at Gal- Overlapping the uppermost records obtained for lipuen, it was difficult to identify the Hypselocy- crussoliceratids and in the overlying three lime- clum Zone as proposed by Atrops (1982), who paid stone-beds was found Progeronia (Hugueni- special attention to the relative record of Ataxio- sphinctes) sp. gr. Proger. (Huguenisphinctes) bre- ceras and Crussoliceras. In contrast, the base of viceps (QUENSTEDT). Taramelliceras sp was reco- the Divisum Zone placed at the FAD of crusso- vered from the limestone-bed just above the youn- liceratids seems easier to apply in the Gallipuen gest crussoliceratids registered, and Streblites le- area, such as usually has been made in condensed vipictus (FONTANNES) from a limestone-bed one epioceanic deposits of the Mediterranean Tethys meter above. (Sapunov, 1977; Oloriz, 1978; Sarti, 1985, The biochronostratigraphic interpretation ap- 1988a-b, 1990, 1994). However, an equivalent to plied to the upper 13,45m in the Kimmeridgian sec- the Hypselocyclum/Divisum Zone Boundary pro- tion at Gallipuen cannot be conclusive due to limita- posed by Atrops (1982) could be recognised in the tions induced by the ammonite record obtained. In interpretations made by Pavia et al. (1987) and fact, the lower 10,80m belong to the Divisum Zone, Sarti (1993) on ammonitico rosso sections in the but the upper 2,66m, which are above the youngest Venetian Alps, NE Italy. Notwithstanding, these Crussoliceras and Garnierisphinctes registered, authors worked on condensed deposits, and their yielded rare ammonites among which the most si- records reported from horizons below the Divisum gnificant for biochronostratigraphy were Progero- Zone, or equivalent stratigraphic intervals assum- nia (Huguenisphinctes) group of breviceps QUEN- edly deposited during the Divisum Chron, alluded to STEDT and Orthaspidoceras group of lallierianum specimens of rare and/or less typical crusso- OPPEL. The holotypes of these two species have liceratids. Caracuel et al. (1998) showed the sig- been assigned to the Upper Kimmeridgian, but these nificance of the Divisum Zone based on the range of species and/or their relatives could exist in the up- Crussoliceras for correlation in The Mediterranean permost Lower Kimmeridgian Divisum Zone as envi- Tethys. saged by Atrops (1982) for Huguenisphinctes. In addition, Finkel (1992) reported Progeronia sp. ex. 4.1.3 DIVISUM ZONE gr. breviceps (QUENSTEDT), as well as Proger. A relative increase in ammonites occurs in the lictor (FONTANNES) and relatives from the Lower Galllipuen area just from the FAD of crussolicera- Kimmeridgian slightly north-eastern from Galli- tids upward in the Lower Kimmeridgian section. puen. Other records of interest from North Italy This trend is interpreted to be not an artefact since are those of Progeronia (Huguenisphinctes) bre- this was registered despite of the fact that lime- viceps (QUENSTEDT), Prog. (Huguenisphinctes) sp. stone-beds, the ammonite bearing deposits at Galli- aff. breviceps (QUENSTEDT), Prog. (Hugue- puen, decreased in number and thickness (Fig. 2). In nisphinctes) sp. cf. ernesti (LORIOL), Proger. (Hu- contrast, siliciclastic deposition increased notably guenisphinctes) spp. (sp1 and sp2 in Pavia et al., within the range of crussoliceratids, and carbona- 1987), Proger. (Progeronia) pseudolictor (CHOF- tes registered episodic deviations recognised as FAT), and Orthaspidoceras sp. cf. lallierianum (OP- sandy dolostone-horizons. Thus, marly and shaly PEL) from the Divisum Zone at northern Monti Les- deposits poor in ammonites overwhelming domina- sini (Pavia et al., 1987). Moreover, Progeronia sp. ted during the late Early Kimmeridgian, and pro- gr. pseudolictor (CHOFFAT) was collected from the bably also during the earliest Late Kimmeridgian, Divisum Zone in the Venetian Alps and Progeronia providing the typical appearance of the upper sp. cf. eggeri (AMMON) and Prog. sp. aff. ernesti 13,45m of the Kimmeridgian section at Gallipuen. (LORIOL) from the Acanthicum Zone in the same Among ammonites, the record of Ataxiocerati- area (Sarti, 1993). In addition, the significant ab- nae was nearly counterbalanced by other Ammoni- sence of Orthaspidoceras uhlandi (OPPEL), a com- tina (aspidoceratids) by the first time. The ammo- mon species in the upper Divisum Zone from both nites recovered were Crussoliceras divisum epicontinental and epioceanic deposits (OI6riz, (QUENSTEDT), Crussol. sp. cf. Crussol. crussoliense 1978; Pavia et al., 1987; Marques & OI6riz, 1992; (FONTANNES), Crussol. sp. gr. Crussol. almolaense Oloriz & Rodr'guez-Tovar 1993; Sarti, 1993; and (GEYER & OL6RIZ), Garnierisphinctes championneti Caracuel at al., 1998; among others) is significant. (FONTANNES), Garnierisph. sp., Progeronia This absence could be interpreted as related to (Huguenisphinctes) sp., Nebrodites sp., Mesosimo- sampling hazards or to erosions in the area before 102 the preserved Lower Cretaceous (Bedulian). Both casses, affecting especially to great-size speci- two hypotheses could be admitted since the record men, and finally to that ammonite assemblages of this species is known from close areas of the might be considered a mixture of in-situ and Iberian Chain (Atrops & Melendez, 1984; Finkel, slightly drifted shells. Since no cases of reworking 1992) and could be used to identify the Uhlandi with biostratigraphic incidence were identified to Subzone in the area, as envisaged by Oloriz & Ro- affect to ammonites, the studied ammonite assem- dri'guez-Tovar (1993). blages offer a very common picture of the ammo- nite record in low energy deposits. 4.2 FOSSIL ASSEMBLAGES AND The averaged composition of fossil assemblages ECOSTRATIGRAPHIC REMARKS per ammonite zone is shown in Fig. 2. Pie-diagrams in Fig. 2 include, separately, the representation of Based on precise ammonite biostratigraphy and the composition of ammonite assemblages, but the then the biochronostratigraphic interpretation, the total amount of ammonites and other cephalopods approach to ecostratigraphic interpretations is (belemnitids, nautiloids) are included as distinct available to envisage long-term paleoecologic components in pie-diagrams showing invertebrates trends according to the fossil record of macroin- other than ammonites. Thus, can be analysed the vertebrate assemblages (megabenthos included). main traits of the evolution of the record obtained The following comments will be necessarily re- of pelagic vs. benthic macroinvertebrates (including stricted to the data obtained at Gallipuen and, the- megabenthos), as well as fluctuations in the compo- refore, their ecostratigraphic interpretation should sition of ammonite groups. The latter were selected be complemented with data recovered from other according to the total amount of ammonites registe- areas in the Iberian Chain before to be conclusive. red per stratigraphic interval of reference (here, An early approach to the interpretation of the the zone level) and the relative significance assu- composition of fossil assemblages of macroinverte- med for different groupings, which was deduced brates in the Gallipuen area was given by Oloriz et from previous research (for an extended treatment al. (1988). Finkel (1992) provided pie-diagrams see references in Oloriz & Rodri'guez-Tovar; this with the composition of ammonite assemblages at volume). the genus level according to data recovered north- Two major features shown in Fig. 2 are the ab- eastern from Gallipuen. In the Lower Kimmeridgian, sence of Phylloceratina and Lytoceratina in ammo- this author gave a combined spectrum for Pla- nite assemblages, and the accentuated increase of tynota+Hypselocylum Zones separately from the invertebrates other than ammonites in the Hypse- ammonite spectrum belonging to the Divisum Zone. locyclum Zone. The exclusive record of Ammonitina The Kimmeridgian section analysed at Gallipuen points to ammonite assemblages representatives of yielded a total of 4732 specimens and fragments inhabitants of relatively proximal ammonite ecos- recovered in situ, bed-by-bed, following sampling paces, certainly unconnected from (and sporadi- methods and observations widely applied to Upper cally connected with) open sea environments and Jurassic deposits by OI6riz and collaborators du- their communities. Given the assumed potential for ring more than a decade (see references in OI6riz & post-mortem transport in Phylloceratina, among Rodri'guez-Tovar, this volume). Main traits of pre- which existed ubiquitous taxa (i.e. Sowerbyceras), servation include dominant concave-up position of the non-record of these ammonites reinforces our valves in epibenthic bivalves, some of them show- interpretation of a inland, epeiric, sea colonised by ing open and articulated valves or complete au- the Ammonitina registered, even assuming some ricles; although separate valves including those of degree of post-mortem transportation of cephalo- endobenthic bivalves were found in horizons with pod shells. This hypotheses fits well paleoecologic dense packing of fossils. Moreover, ammonites with and paleobiogeographic data and interpretations in preserved lappets and/or peristomal and pre- Oloriz et al.(1988) and does not contradict the en- peristomal structures are frequent, and the majo- vironmental reconstruction as proposed by Aurell rity of those that show epibionts are usually of (1990), Aurell & Mel6ndez (1990), Badenas great size. Body-chamber preservation is common, (1997), and Badenas & Aurell (1997). and isolates phragmocones are secondary, as they The accentuated increase of invertebrates other are fragments of body chambers that belong to than ammonites in the Hypselocyclum Zone separa- great-size specimens that usually show epibionts tes the lower (Platynota Zone) and the upper (Divi- (serpulids and briozoa, among others). Aptychi are sum Zone) parts of the Kimmeridgian section at scarce, but some were found within the body cham- Gallipuen. The averaged record registered in the ber in aspidoceratids. On the whole, these data Hypselocyclum Zone might be related with ecospace point to dominant settling in low energy conditions, deterioration for ammonites. This agrees with the with episodic exceptions, to the existence of varia- long-time assumed picture of proximal environ- ble post-mortem transportation of cephalopod car- 103

merits enriched in benthics, and impoverished to cognition of transgressive deposits, with or with- devoid in ammonites, while increasing distance out signs of relative condensation, in southern Eu- from shore would reverse the assumed ratios rope and northern Africa. landwards. In general, this hypothesis is acceptable The comparison with data on ammonite assem- and supported by empirical data elsewhere. On the blages in Finkel (1992) is limited since this author other hand, since vagrant organisms track favou- did not separate data from the Platynota and Hyp- rable environments as far as possible, following selocyclum Zones [its UKK (l/ll) ammonite spec- shifting ecological conditions, ammonites should trum], UKK (l/ll) ammonite spectrum in Finkel have a similar behaviour despite of their low capa- (1992) differs from the averaged spectrum of the bility for swimming. Therefore, pattern of currents Platynota plus Hypselocyclum Zones in Fig. 2. UKK and movements of water masses essentially drove (l/ll) ammonite spectrum in Finkel (1992) shows ammonites. In accordance, major fluctuations in the higher number of Ataxioceratinae and Sutneria re- record of ammonites might be interpreted, finally, covered from the area studied by Finkel (1992), in terms of shifting ecospaces. In accordance, the the latter genus being fortuitous component significant decrease in ammonites registered in the throughout the Kimmeridgian section studied at Gal- Hypselocyclum Zone at Gallipuen might correspond lipuen. The UKK (III) ammonite spectrum in Finkel to unfavourable life-conditions for ammonites. (1992) corresponds to the Divisum Zone and is OI6riz et al. (1993) identified this feature as more similar to that shown in Fig. 2 for the same general one registered in Lower Kimmeridgian stratigraphic interval. In fact, Finkel' spectrum deposits from southern epicontinental shelves in shows a comparatively balanced ammonite assem- Europe and North Africa, in relation with ecospace blage respect to the lower UKK (l/ll) ammonite reduction during low sea levels and deposition under spectrum. However, Sutneria and "simoceratins" shelf-margin-wedge conditions. This interpretation are more abundant while haploceratids are rarer in agrees with proposals of sequence stratigraphy Finkel' spectrum of the Divisum Zone. Anyway, that worked on the southern paleomargin of Iberia does the ammonite record studied by Finkel (1992) (Marques et al., 1991; Rodriguez-Tovar, 1993; to share a feature with that analysed in this paper Oloriz et al., 1994; among others), but contrasts is the comparatively balanced ammonite assemblage with those applied in the Iberian Chain (Aurell, recovered from the Divisum Zone. This is a fact of 1990; Badenas, 1997; among others). We interpret significance revealing enlarging ecospaces for am- this disagreement to results from different orders monites in north-eastern Iberian shelves (Iberian of observations and analyses, resulting in the Chain) during the late Early Kimmeridgian, the time recognition of sequences of different order, that for major flooding in epicontinental shelves sur- does not contradict the hierarchical model of rounding Iberia (Aurell, 1990; Marques et al., sequence stratigraphy. 1991; Marques & Oloriz, 1992; Rodriguez-Tovar, Concerning the composition of ammonite assem- 1993; Badenas, 1997; among others). Other diffe- blages, pie-diagrams in Fig. 2 show the Ataxio- rences between Finkel' ammonite assemblages and ceratinae as the dominant group, and its progres- those discussed in this paper could be the evidence sive decrease throughout the Lower Kimmeridgian. of minor ecological (and taphonomic?) differences The latter determines an increasing balance in the for epicontinental (neritic) ammonites in different composition of ammonite assemblages. An inte- sectors of the flat-topped ramp developed during resting feature is registered in the Hypselocyclum the Kimmeridgian in eastern Iberia. Zone where the ammonite epitome combines de- Recent paleoenvironmental reconstruction (Au- creased Ataxioceratinae and increased haplocera- rell & Melendez, 1990; Badenas, 1997) agrees tids and others Ammonitina, especially aspidocera- with our data and taphonomic and ecostratigraphic tids. In paleoecologic terms, it is compatible with interpretations. All of this reinforce previous in- relatively long-term processes of ecological turn- terpretations by Oloriz et al. (1988) on factors over during maximal reduction of ecospace (i.e. the controlling the composition of fossil assemblages, highly specialised Ataxioceratinae giving place to especially ammonites, from the Platynota Zone in other more generalists ammonites). Thus, the the Alcorisa-Berge area, which includes the lower higher amount of aspidoceratids together with less part of the Kimmeridgian section studied at Galli- specialised Ataxioceratinae were registered in the puen. In fact, the extensional lagoon interpreted by Divisum Zone, the stratigraphic interval that shows Aurell & Melendez (1990) and the external ramp of the most balanced composition in the averaged am- the flat-topped-ramp proposed by Badenas (1997) monite assemblage. These facts are in accordance as depositional models for the Kimmeridgian Lori- with enlarging ecospaces related to higher sea le- guilla Fm. does not contradict the interpretation by vels during the late Early Kimmeridgian, the latter Oloriz et al. (1988) of a turbid, shallow-water, and being a widely accepted hypothesis based on the r e - nutrient-rich environment, with depths rather 104 shallower than deeper in the range of 30-80m, and ammonite assemblage, at the zone level, in accor- with restricted connection with open seas during dance with the maximum flooding in the area. the Platynota Chron. In such a depositional envi- ronment, Oloriz et al. (1988) concluded that the ecological scenario determined the catching of am- ACKNOWLEDGEMENTS monite assemblages through adaptation. Hence, This research was made within the frame of their composition was ecologically determined, activities of Project PB97-0803 (DGCYT) and the which was interpreted in terms of the so-called EMMI Group (RNM-178, Junta de Andaluc'a). Platform-Effect (OI6riz, 1985; Oloriz et al., 1988). The ecological dynamics of the model applied by OI6riz et al. (1988) fit well with ecostra- REFERENCES tigraphic interpretations in this paper, which are based on the assumption of shifting ecospaces du- Atrops, F. (1982): La sous-Famillie des Ataxioceratinae ring fluctuations of sea level as modelled by OI6riz (Ammonitina) dans le Kimmeridgien inferieur de Sud- et al. (1995) within the conceptual template of Se- Est de la France: syst6matique, evolution, chrono- quence stratigraphy. stratigraphie des genres Orthosphinctes et Ataxio- ceras. Doc. Geol. Lyon, 83: 463 p. Atrops, F. & Benest, M. 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