Palaeogeography and relative sea-level history forcing eco-sedimentary contexts in Late Jurassic epicontinental shelves (Prebetic Zone, Betic Cordillera): An ecostratigraphic approach

Earth-Science Reviews 111 (2012) 154–178

Contents lists available at SciVerse ScienceDirect

Earth-Science Reviews

j ourna l homepage: www.e lsev ie r .com/ locate /earsc i rev

Palaeogeography and relative sea-level history forcing eco-sedimentary contexts inLate Jurassic epicontinental shelves (Prebetic Zone, Betic Cordillera): Anecostratigraphic approach

Federico Olóriz a,⁎, Matías Reolid b, Francisco J. Rodríguez-Tovar a

a Dept. Estratigrafía y Paleontología, Facultad de Ciencias, Universidad de Granada. Campus Fuentenueva, 18071 Granada, Spainb Dept. Geología, Universidad de Jaén, Campus Las Lagunillas, 23071 Jaén, Spain

⁎ Corresponding author. Tel.: +34 958 243 345; fax:E-mail addresses: [email protected] (F. Olóriz), mreolid

0012-8252/$ – see front matter © 2011 Elsevier B.V. Alldoi:10.1016/j.earscirev.2011.11.004

a b s t r a c t
a r t i c l e i n f o
Article history:Received 22 March 2010Accepted 8 November 2011Available online 19 November 2011

Keywords:PalaeogeographySea-levelEco-sedimentary conditionsMacroinvertebratesForaminiferaUpper Jurassic

The analysis of macroinvertebrate and foraminiferal assemblages from Upper Jurassic (Middle Oxfordian toLower Kimmeridgian) epicontinental shelf deposits in the Prebetic (Betic Cordillera, southern Spain) revealsthe influence of environmental changes. They are expressed as selected parameters in palaeogeographic andstratigraphic trends (litho- and microfacies, faunal composition, taphonomy), which are interpreted in thecontext of relative sea-level histories.Middle Oxfordian to early Kimmeridgian (Transversarium to Planula Chrones) rocks and faunal assemblagesin comparatively distal sectors (distal shelf) show lower sedimentation rates (lumpy lithofacies), and higherproportions of ammonoids, planktic foraminifera, corrasion degree, microboring and encrustation. Land-wards, towards the mid-shelf, eco-sedimentary conditions resulted in spongiolithic limestones and marl-limestone rhythmites with local development of microbial-sponge buildups.Greater distance from shore during relative sea-level highs accords with greater: (1) stratigraphic condensa-tion; (2) abundance in ammonoids, planktic foraminifera and nubeculariids; and (3) degrees of corrasion,microboring and encrustation. These trends in faunal composition and taphonomy agree with backsteppingphases, increasing ecospace and a longer exposition of shelly remains on the sea bottom.Decreasing distance from shore during relative sea-level lows relates to opposite trends, as evidenced by: (4)increasing terrigenous input and decreasing stratigraphic condensation; (5) impoverishment in ammonoidsand planktic foraminifera; and (6) diminution of corrasion, microboring and encrustation. Phases of forestep-ping/progradation and aggradation, a reduction of ecospace for nekto-planktic organisms, and comparativelyrapid burial of shell remains are interpreted to force the recorded trends.An ecostratigraphic approach is used here to correlate and characterise sea-level changes, applying high res-olution stratigraphy to sections where the identification of relevant surfaces is more difficult. The changes indistance from shore and ecospace, triggered by relative sea-level fluctuations, are considered prime factorsforcing trade-offs in faunal communities of the studied fossil assemblages. Ecostratigraphy was used as atemplate for the characterization, correlation and interpretation of relative sea-levels and associated sedi-mentary packages in a time span from just above the Milankovitch band to the million-year scale.

© 2011 Elsevier B.V. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1551.1. Remarks on sequence stratigraphy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1551.2. Ecostratigraphy — from HIRES to the record of ecological dynamics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155

2. Geological setting. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1593. Material and methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1594. Foraminiferal assemblages. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161

4.1. Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1614.2. Taphonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1624.3. Stratigraphic distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162

+34 958 248 [email protected] (M. Reolid), [email protected] (F.J. Rodríguez-Tovar).

rights reserved.

http://dx.doi.org/10.1016/j.earscirev.2011.11.004

mailto:[email protected]



http://dx.doi.org/10.1016/j.earscirev.2011.11.004

http://www.sciencedirect.com/science/journal/00128252

155F. Olóriz et al. / Earth-Science Reviews 111 (2012) 154–178

5. Macroinvertebrate fossil assemblages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1625.1. Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1625.2. Taphonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1625.3. Stratigraphic distribution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163

6. Facies, stratigraphic intervals and ecostratigraphy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1666.1. Stratigraphic Interval I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1676.2. Stratigraphic Interval II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1676.3. Stratigraphic Interval III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1686.4. Stratigraphic Interval IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168

7. Sequence stratigraphic interpretation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1687.1. Stratigraphic Interval I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1697.2. Stratigraphic Interval II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1707.3. Stratigraphic Interval III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1727.4. Stratigraphic Interval IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172

8. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174

1. Introduction

The southeastern palaeomargin of the Iberian subplate became dif-ferentiated in epicontinental and epioceanic environments (the PrebeticZone and the Subbetic Zone and lateral equivalents, respectively) early inthe Jurassic, and drowning of the epicontinental shelf took place in theLate Jurassic (Olóriz et al., 2002a; García-Hernández and López-Garrido,2004). Studies of regional geology in the Prebetic Zone during the se-venties (Behmel, 1970; Foucault, 1971; Jerez-Mir, 1973; Azéma, 1977;García-Hernández, 1978) provide data on Upper Jurassic deposits. Re-cent approaches to interpret Late Jurassic deposits in the Prebetic Zoneentail precise biostratigraphy, taphonomyandpaleoenvironmetal recon-struction, with preliminary ecostratigraphic interpretations (Rodríguez-Tovar, 1993; Olóriz et al., 1999, 2002b, 2003b; Reolid, 2003; Olórizet al., 2004a,b, 2006; Reolid, 2007; Olóriz et al., 2008; Reolid et al.,2008a,c). Herewe propose ecostratigraphic analyses to interpret Late Ju-rassic deposits from the Prebetic Zone in terms of sequence stratigraphy.

1.1. Remarks on sequence stratigraphy

An integrative approach to sedimentary packages can be con-ducted through sequence stratigraphy analysis. However, even afterthirty years of development, such an approach can give rise to debateregarding differences between subsurface and outcrop level observa-tions. There are disparate models of sequence and surface definition,interpretation and correlation, as well as conceptual revisions forthe evaluation of surface dyachroneity (for a recent, general picturesee Miall and Miall, 2001; Miall, 2004; Schlager, 2005; Christie-Blicket al., 2007; Embry et al., 2007; Catuneanu et al., 2009, 2010; Bhatta-charya, 2011; and references therein). At present, sequence strati-graphic interpretations cover a wide array of unconformity-boundedand unconformity-plus-relative-conformity-bounded sedimentarypackages. Moreover, assumptions about the chronostratigraphic po-tential of reference surfaces and subsequence units can vary (i.e., se-quence boundaries and other intra-sequence surfaces of reference,systems tracts and parasequences). Early on, the so-called “procedureinverse”was supported in sequence stratigraphic interpretation, withcorrelation on the basis of assumed surface isochroneity regardless ofthe depositional environment – e.g., Mitchum and Vail (1977), Vail etal. (1987, 1991), Haq et al. (1988), Van Wagoner et al. (1987, 1988),Galloway (1989), Hunt and Tucker (1992 pro parte), Plint andNummedal (2000), Posamentier and Morris (2000) – and identifica-tion of surface dyachroneity and use of physical surfaces minimisingdiachrony (e.g., Nummedal and Swift, 1987; Hunt and Tucker, 1992pro parte; Helland-Hansen and Gjelberg, 1994; Catuneanu et al.,

1998; Posamentier and Allen, 1999; Anderson, 2005; Catuneanu,2006; Embry et al., 2007; Catuneanu et al., 2010; Bhattacharya,2011; but see Christie-Blick et al., 2007). Siliciclastic and carbonatesystems, as well as mixed carbonate–siliciclastic systems, are knownto represent scenarios forcing different, autocyclic sedimentarytrends for a given allocyclic process of variable magnitude — indeed,Davaud and Lombard (1973), Hallam (1986), Sarg (1988), Dromart(1989), Galloway (1989, 1998, 2002), Einsele and Ricken (1991),Schlager (1991, 1992, 2005), Hunt and Tucker (1993), Leinfelder etal. (1993), Pomar (1993, 2001a,b), Pomar and Ward (1995), Pittetand Strasser (1998a,b), Brandano and Corda (2002), Catuneanu(2002), Kerans and Loucks (2002), Anderson (2005), and Pomarand Kendall (2008) all describe records of sedimentary packages asdisparate responses to factors controlling local siliciclastic vs. carbon-ate and mixed carbonate–siliciclastic systems.

We agree that sequence stratigraphy analysis is a valuable tem-plate for interpreting stratigraphic architectures, and coincide withEmbry (2002), amongst others, on focusing sequence stratigraphy in-terpretation on changes in sedimentary trends and their identifiablebounding surfaces, while recognising limitations for surface identifi-cation in distal and/or basinal settings (Embry, 1995; Posamentierand Allen, 1999; Plint and Nummedal, 2000; Embry, 2002;Posamentier and Kolla, 2003; Posamentier and Walker, 2006;Christie-Blick et al., 2007; Catuneanu et al., 2009, 2010), especiallywhen monotonous low-energy conditions persist. In such a context,we conducted ecostratigraphic analysis based on the selected fossilassemblages described below.

1.2. Ecostratigraphy — from HIRES to the record of ecological dynamics

Ecostratigraphy assumes that sedimentary successions record in-formation about environmental dynamics affecting the ecologicalconditions for diverse organisms and/or species assemblies, with fluc-tuations of key environmental factors forcing ecological tradeoffs'readjustment in the local community. From the understanding ofecostratigraphy as the stratigraphy of ecosystems (Martinsson,1973) to the relevance of the ecostratigraphic analysis to identifybioevents within the framework of “High Resolution Event Stratigra-phy” (HIRES in Kauffman, 1986, 1988) as promoted by Boucot (1986),ecostratigraphy appears as a tool of great potential in basin analysisthat is environmentally oriented (ecology included; e.g., Brett andBaird, 1997; Brett, 1998; Brett et al., 2007a,b). One major avenue forecostratigraphy is to improve stratigraphic subdivisions by providingmore detailed stratigraphic intervals than those obtained from classicbiostratigraphy (Waterhouse, 1976; Sokolov, 1988). Thus,

Fig. 1. Location and geological setting. Note studied sections at Riogazas-Chorro-II (RGCHSP); El Chorro (CHO); Puerto Lorente (PL); Pozo Cañada (PC); Chinchilla (CH); FuenteÁlamo (FA); Navalperal (NV); and Río Segura (RS).

156 F. Olóriz et al. / Earth-Science Reviews 111 (2012) 154–178

ecostratigraphy can reveal palaeoecological frameworks in close rela-tion with the stratigraphic intervals used traditionally (Rabe andCisne, 1980; Boucot, 1982; Cisne and Chandlee, 1982; Berry, 1983;Cisne et al., 1984; Martin, 1991; amongst others), which applies to re-cent studies (e.g., Holland and Patzkowsky, 2004) including recogni-tion of the incidence of stratigraphic artefacts (e.g., Holland andPatzkowsky, 2009). From the 1970s (“Proyect Ecostratigraphy”IGCP) to the nineties, ecostratigraphy evolved as a methodology foranalysing community–environment interactions (“Palaeocommu-nities — A case study from the Silurian and Lower Devonian, Final Re-port IGCP, Project 53 Ecostratigraphy, 1996”; Boucot and Lawson,1999). Raffi and Serpagli (1993) proposed the term Ecobiostratigra-phy to distinguish biostratigraphy from a merely descriptive practicein stratigraphy (e.g., Catuneanu et al., 2010). Later approaches useecostratigraphy in terms of records of long-term ecological dynamics(e.g., Olóriz, 2000 and references therein), and valuable contributionsabout changes in fossil assemblages and sea-level fluctuations in epi-continental areas can be included in this approach (e.g., Brett andBaird, 1986; Brett, 1988a,b; Brett and Baird, 1997; Brett, 1998; Brettet al., 2006, 2007a,b).

Also important in this context is the understanding of two distinctbut related concepts: accommodation and ecospace. Accommodation(Jervey, 1988 but see Muto and Steel, 2000 and Catuneanu et al., 2009for conceptual fine-tuning) refers to a physical space available for po-tential sedimentation. Biotically induced changes in bottom profiles,and therefore in accommodation, are rightly interpreted as ecologicalaccommodation, in turn related to carbonate factory dynamics(Pomar, 2001a; Pomar et al., 2005; Pomar and Kendall, 2008). Accom-modation dynamics in marine environments influence abiotic–bioticenvironmental complexes and biofacies distribution, with conse-quent fluctuations in the composition of fossil assemblages. In thissense, accommodation entails a complex multidimensional ecologicalvolume known as ecospace (Fig. 2; see Olóriz et al., 1993, 1995 andreferences therein). Accordingly, ecospace is a multidimensionalspace for biotic interactions amongst biological entities and the phys-ical–chemical environment surrounding them (e.g., from Valentine,1973 onwards). When ecospace is projected on a given physical(three-dimensional) space such as accommodation in marine envi-ronments, it may refer to either the total ecospace for species assem-blies inhabiting a given environment, or the ecospace for a particulargroup of organisms under study (Olóriz, 2000). Thus, the simplestunit of ecospace in a given area would correspond to the water

column plus the sediment–water interphase — i.e., the Taphonomi-cally Active Zone or TAZ.

Awide array of ecospace situations is possible inmarine, natural en-vironments. They range fromminimal ecospace resulting from zero ac-commodation (only infaunal forms living in final phases of basininfilling) to maximal ecospace (maximum accommodation coupledwith favourable ecological conditions). Yet no linear relationship be-tween ecospace and accommodation is common, even when only oneparticular biological group is investigated. Ecospace dynamics is indi-rectly related to the physical environment since it will be mediated byecological tolerance to changing environmental conditions. Changingaccommodation will force shifting environmental conditions, and viceversa, resulting in changes in ecospace (e.g., Olóriz, 2000 and referencestherein). Stratigraphic records of changing ecospace are registered asecostratigraphic events and trends (Olóriz et al., 1995, 2008).

Ecostratigraphic events are significant changes in community struc-ture forced by environmental changes (Kauffman, 1986). In addition tothe identification of ecostratigraphic events for improving high resolutionstratigraphic subdivision (HIRES), the recognition of ecostratigraphictrends (Olóriz et al., 1995) and their combined interpretation with ecos-tratigraphic events (Olóriz et al., 2008) is favoured from a theoreticalpoint of view (Olóriz, 2000). Thus, shifting environmental conditionsthat force changes in accommodation and ecospace necessarily modifythe stratigraphical signature of biotic–abiotic interactions. This appliesto particular cases of given organisms and depositional conditions.

As stated by Olóriz (2000), ecostratigraphic trends can be consid-ered as the stratigraphic signature of dynamic adjustments of multi-species groupings interacting with the physical environment overmuch longer time intervals than those involved in ecostratigraphicevents (i.e., over a thousand years). Ecostratigraphic interpretationsapplied to long-term palaeoecology are based on the assumptionthat no major taphonomic distortion occurred in a context ofwithin-habitat time averaging (Olóriz, 2000). Furthermore, a com-plex, combined signal is expected due to the disparate ecologies ofparticular organisms under study. No matter how intricate the recordof ecological responses may be, the coherence of ecological responsesmust be proven for a reliable ecostratigraphic interpretation.

The following remarks on the ecospace concept apply to the casestudy: (1) the accommodation-ecospace template; and (2) the eco-space within the substrate.

(1) Ecostratigraphic interpretations of faunal assemblages call fora knowledge of depositional dynamics. Relative sea-level fluctuations,

Fig. 2.Model of relationships between accommodation and ecospace and the response of foraminiferal and macroinvertebrate assemblages from mid- to distal-shelf environments.Other environmental parameters fluctuate according to distance from shore.


changing accommodation and shifting facies are fundamental forinterpreting stratal patterns and depositional environments withinthe template of systems tracts and depositional sequences (e.g.,Brett and Baird, 1986; Brett, 1988a,b, 1995, 1998; Brett et al., 2006).Thus, the stratigraphic architecture is associated with the accommo-dation history, which in turn affects ecospace.

When only subtle changes in lithofacies result, fossil assemblageanalysis can identify potential fluctuations in accommodation and eco-space (e.g., Holland et al., 2000). The latter would result from biotic in-teractions in each particular case under study (e.g., ecological dynamicsof benthos vs. ecological dynamics in nektonic and planktic organisms).

In epicontinental shelves, enlarging/contracting ecospace dynam-ics expose planktic, nektonic and benthic faunas to different degreesof ecological stress. In the Upper Jurassic deposits investigated, ben-thic and nekto-planktic organisms were the most sensitive faunabased on their stratigraphic record (Fig. 3). Ammonoids were espe-cially susceptible to environmental fluctuations. The neritic oneswere commonly captured – perched – through adaptation to localconditions (Platform Effect in Olóriz, 1985, 1990) and unable to aban-don the epicontinental environments during high or low sea-levels,as reflected by the lack of neritic assemblages in epioceanic deposits.In contrast, epioceanic ammonoids colonised neritic shelves duringflooding episodes (Olóriz, 1985, 1990; Olóriz et al., 1993, 1995;Olóriz, 2000). Hence, changes in ammonite assemblages, amongstother susceptible organisms in the water column, are reliably relatedto relative sea-level fluctuations undetected through comparativelypersistent epibenthos, especially for epiocenic fringes such as thoseknown form Tethyan areas (citations above).

(2) With the above assumption of relative sea-level fluctuations asrelevant controlling factors of the composition of fossil assemblages,through dynamic relationships amongst accommodation, ecospace

and distance to the shoreline. Hence, environmental parameters canbe said to mainly relate to distance from shore in the case study.Thus, sedimentation rate, terrigenous input, nutrient influx, water en-ergy, oxygenation degree and organic content in sediments are primefactors determining ecospace conditions within the substrate and therelated ecological dynamics for endobenthos. Infaunal microhabitatdepth (as an expression of ecospace for endobenthos) is related tonet sedimentation rates and the redox boundary, the latter dependingon organic matter content and oxygenation degree within the sedi-ment–water interphase (i.e., the TAZ vs. the sediment/water surface;Fig. 3). Jorissen et al. (1995) and Van der Zwaan et al. (1999) mod-elled microhabitats for infaunal forams based on food availabilityand oxygenation as the limiting factors for within-substrate ecology.However, a more intrincate scenario can be envisaged when the re-gional variation of species microhabitat depth is known (Jorissen,1999), along with the influence of infaunal bio-irrigation influx onmicroenvironmental conditions and redox patterns (Gobeil et al.,1997; Aller et al., 1998; Jørgensen et al., 2005) and subsequent distri-butions of endobenthic foraminifera (Schmiedl et al., 2004).

In a neritic-shelf environment, organic matter consumption cantake place directly or indirectly, involving: 1) autochthonous or para-utochthonous material and nutrients proceeding largely from prima-ry photosynthetic production; and/or 2) particulate material andnutrients exported from other sectors (shallower or emerged areas),in relation to water mass hydrodynamics and/or detritic influxes. Pri-mary photosynthetic production depends on light availability andtakes place close to shallow sea-bottoms and/or in the water column,the productivity driven by phytobenthos and phytoplankton, respec-tively. Primary production in mid- and distal-shelf areas is lower thanin the proximal shelf, and a major part of the organic matter inputcomes from influxes from emerged and more proximal areas,

image of Fig.�2

Fig. 3.Model of distribution for Late Jurassic vagile benthic foraminifera modified from Reolid (2003). Oxygenation degree and nutrients availability are the main controlling factorsof infaunal ecospace. Based on the “TROX model” of Jorissen et al. (1995) and the “TROX-2 model” of Van der Zwaan et al. (1999).


diminishing with distance from shore when fertilisation under up-welling influence is discarded. Phytodetritus inputs re-distribute or-ganic matter from the shallow proximal euphotic zone towardsmore distal and commonly deeper areas (Gooday, 1996). Offshore de-creasing inputs of siliciclastics and nutrients have been recently de-scribed in Late Jurassic epicontinental platforms (Pittet and Gorin,1997; Insalaco, 1999; Pittet and Mattioli, 2002; Bartolini et al., 2003;Olóriz et al., 2003a, 2006; Reolid et al., 2008a,b).

The organic matter content in sediments, hence the trophic re-sources for detritivorous, depends onnet sedimentation rate and organ-ic matter accumulation. In hemipelagic settings, the sedimentation ratedepends on climatic, tectonic and accommodation dynamics, and relat-ed hydrodynamic effects. In carbonate shelves, sedimentation dependslargely on active biogenic production, while accommodation is sub-jected to regulation close to the carbonate factory (i.e., ecological ac-commodation in Pomar, 2001a; Pomar et al., 2005; Pomar andKendall, 2008) and its influence varies in the short- and long-term(Pomar, 2001a; Pomar et al., 2005). The combination of terrigenous in-puts and accommodation dynamics affects photosynthetic primary

productivity through changing water transparence and hydrodynamics(energy) on the proximal shelf.

Within the seabed, the redox boundary location determines theavailable ecospace and ecological structuring for endobenthos, takinginto account local influences of potential irrigation downwards. In-creasedwithin-sediment organicmatter content raisesmetabolic activ-ity in the microbiota, consuming pore-water oxygen. Gooday (1996)described the close relationship between organic matter input and sed-iment oxygenation. According to Kuhnt et al. (1996), the redox bound-ary location under oligotrophic conditions may be several metres deepand thus lie below the zone of benthic activity; whereas under condi-tions of very high organic influx, the redox boundary layer can beclose to the sediment–water interface (an unfavourable situation fordeep substrate colonisation by endobenthos).

In light of the above, we favour ecostratigraphic interpretations de-rived through integrating data from different disciplines (e.g., Boucot,1982, 2005) combining taphonomy, palaeoecology, microfacies andlithofacies, elaborated under precise biostratigraphic control, to contrib-ute to sequence stratigraphic interpretations in the case at hand.

image of Fig.�3


Examples of palaeontological information, including biostratigraphy forprecise recognition of sedimentary packages interpreted in terms of se-quence stratigraphy, stem from the late eighties to early nineties. Theydocument:

– Species abundance and diversity, morphogroups and compositionof the foraminiferal assemblages (e.g., Cubaynes et al., 1989, 1990;Bonnet et al., 1991, 1992; Magniez-Jannin, 1992; Haig and Lynch,1993; Rey et al., 1994; Hylton and Hart, 2000; Nagy et al., 2001;Olóriz et al., 2003a; Reolid and Nagy, 2008; amongst others).

– Changes in composition and diversity of benthic macroinvertebrateassemblages related to type of substrate, environmental instabilityand sedimentation rate, controlled by transgressive–regressive cy-cles (e.g., Fürsich et al., 1991; McRoberts and Aberhan, 1997;Neagu et al., 1997), and the potential forcing of benthic multispeciesmacro–megainvertebrate assemblages through oxygenation levels(e.g., De la Mora et al., 2000).

– Ammonoid assemblage composition as a reflection of sea-level fluc-tuations given the ecological requirements of each group in the ab-sence of taphonomic distortion (e.g., Donovan, 1985; Gygi, 1986;Hantzpergue, 1991; Olóriz et al., 1991; El Hariri et al., 1992; Olórizet al., 1995, 1996; Yin et al., 1997; amongst others).

– Taphonomic features and taphofacies of macroinvertebrate assem-blages, have been used for interpreting skeletal exposure time and re-lated effects, sedimentary rate, water energy and relative distancefrom shore for depocenters, as evidence of sea-level fluctuations(e.g., Brett and Baird, 1986; Brett, 1988a,b; Fürsich and Oschmann,1993; Fernández-López and Meléndez, 1994; Brett, 1995;Fernández-López, 1997, 2000; Courville and Collin, 2002; Olóriz etal., 2002b, 2004a; Brett et al., 2006).

Thus, over the past twenty years, ecostratigraphic analyses haveproved useful in interpreting macrofossil assemblages and their rolein sequence stratigraphic interpretations of Upper Jurassic depositsin the South-Iberian palaeomargin (Fig. 1), in epicontinental (e.g.,Olóriz et al., 1991; Marques et al., 1993; Rodríguez-Tovar, 1993;Olóriz et al., 1994; Marques et al., 1996; Olóriz and Rodríguez-Tovar,1999; Reolid, 2003; Olóriz et al., 2004a, 2008) and epioceanic(Olóriz et al., 1993, 1995, 1996; Caracuel, 1996) environments.

2. Geological setting

The Prebetic (Betic Cordillera) is largely made up of epicontinentalshelf deposits accumulated on the South-Iberian palaeomargin duringthe Mesozoic. The External and Internal Prebetics are, respectively,the proximal and distal areas of this neritic, carbonate and carbon-ate–siliciclastic shelf system. Geographically, outcropping areas inthe Prebetic are divided into the central (Sierra de Cazorla and Sierrade Segura) and eastern (Altos de Chinchilla) sectors. The successionstudied here corresponds to Middle Oxfordian–Lower Kimmeridgian(Upper Jurassic) sections of the External and Internal Prebetics intheir central and eastern sectors (Figs. 1 and 4).

Oxfordian rocks indicate the first hemipelagic–pelagic conditionsin the Prebetic. They overlie a discontinuity surface truncating whiteoolitic limestones and dolomites related to the Early–Middle Jurassicshelf that developed along the South-Iberian palaeomargin (García-Hernández et al., 1981). The upper boundary of the studied succes-sion is biostratigraphically recognised either within a marl-limestone rhythmite that includes the first Kimmeridgian deposits,or in an omission surface or hardground preceding this rhythmite(Fig. 4).

Four groups of lithofacies are distinguished from the Transversar-ium (Middle Oxfordian) to Planula zones (Lower Kimmeridgian): (1)a lumpy lithofacies group (lumpy limestone, lumpy-oncolitic lime-stone and condensed lumpy-oncolitic limestone), (2) a spongiolithiclithofacies group (spongiolithic limestone and spongiolithic marl

and peloidal limestone), (3) a marl-limestone rhythmite lithofaciesgroup, and finally (4) a marl lithofacies group (Olóriz et al., 2002b).

The lumpy limestones are well-bedded, made up of wackestonewith abundant lumps, iron oxides, and glaucony. The lumpy-oncolitic limestones are nodular-like beds of wacke–packstoneswith abundant lumps and oncoids. The condensed lumpy-oncoliticlimestones consist of nodular-like beds rich in ammonites, made bywacke–packstones of lumps, oncoids and bioclasts. Spongiolithiclimestones are characterised by richness in siliceous sponges andwackestones of bioclasts. The alternation of spongiolithic marl andpeloidal limestones featured wackestones with bioclasts and peloids,and sponge-rich intercalated marls. Marl-limestone rhythmites con-sist of wackestones rich in peloids and glaucony. Marls contain localpeloidal and silty levels, and occasional sandy to marly-limestonehorizons.

In general terms, the lumpy lithofacies group, with microfaciesfeaturing a greater content of oncoids with nubeculariids, is palaeo-geographically limited to comparatively distal areas (Internal Pre-betic). The remaining lithofacies (spongiolithic limestone, marl-limestone rhythmite and marl lithofacies, the microfacies havingmore siliciclastic grains and peloids) are registered in the compara-tively proximal sectors (External Prebetic). The spongiolithic lithofa-cies predominate in the eastern sector, while the marl-limestonerhythmite lithofacies group is more abundant in the central sector.

3. Material and methods

The study was performed on eight profiles (Figs. 1, 4), six in theExternal Prebetic (central sector: RGCHSP, CHO, PL; eastern sector:CH, PC, FA) and two in the Internal Prebetic (NV and RS in the centralsector). Fine-resolution ammonite analyses (Olóriz et al., 1999;Reolid, 2003) allowed for a detailed biostratigraphic characterizationat the zone and subzone level (even some biostratigraphic horizonswere identified) from the Transversarium Zone (Middle Oxfordian)to the Planula Zone (Lower Kimmeridgian).

Microfossil study focused on foraminiferal assemblages, throughthin-section analysis. A total of 370 thin sections were obtainedfrom 146 sampling stations, providing around 31,000 specimens. Instudying foraminiferal assemblages, two types of faunal spectrawere characterised: 1) Spectrum A for the total assemblage, distin-guishing between planktics, vagile benthics and sessile benthics;and 2) Spectrum B for the relative composition of vagile benthic fora-minifera: ophthalmidiids, spirillinids, nodosariids, agglutinatedforms, and other rare forms.

The foraminiferal assemblages were differentiated by test mor-phology (e.g., Bernhard, 1986; Corliss, 1991; Nagy, 1992; Tyszka,1994; Nagy et al., 1995; Fugagnoli, 2004; Reolid et al., 2008a,b;Nagy et al., 2009), and the benthic foraminiferal assemblages weresegregated into three groups depending on their microhabitat depthin the substrate:

(1) Epifaunal foraminifera: Up to one centimetre deep (Corliss,1991). Includes all the sessile foraminifera, ophthalmidiids,miliolids, spiral agglutinated foraminifera (Ammodiscoides, Glo-mospira and Trochammina) and spiral calcareous foraminifera(spirillinids, Epistomina, Trocholina, etc.), except Lenticulina.

(2) Shallow infaunal foraminifera: Depth in sediment less than5 cm, according to Kuhnt et al. (1996), but specified as be-tween 1 and 5 cm in this case-study. Elongated agglutinatedforaminifera, mainly uniserial and showing streptospiral orplanispiral initial stages (Ammobaculites, Ammomarginulinaand Haplophragmium), and elongated calcareous foraminifera(nodosariids such as Nodosaria, Dentalina, Planularia, etc.).

(3) Ubiquitous foraminifera: Distributed in a wide range of depthsin the substrate. Lenticulina, a highly ubiquitous genus that tol-erates a wide range of microhabitats from epifaunal to deep

Fig. 4. Studied sections showing biostratigraphic correlation (based on Olóriz et al., 1999, 2006), distribution of lithofacies, location of omission surfaces and hardgrounds. Sectionlabelling as in Fig. 1.


infaunal (Tyszka, 1994), and uniserial (Reophax) and biserialagglutinated foraminifera (undifferentiated textularids),which vary between shallow and deep infaunal microhabitats.

The methodology applied for the analysis of macroinvertebratefossil assemblages is based on a detailed bed-by-bed study of the suc-cession, with sampling of complete specimens and fragments, and apreliminary taphonomic analysis; sponges, epibionts and crinoidswere only analysed qualitatively to avoid counting problems (Olórizet al., 1993, 1994, 1995, 2006).

A total of 13,500 macroinvertebrate specimens were analysed, dif-ferentiating three types of faunal spectra: (1) The general spectrumfor macroinvertebrates refers to the relative abundance in ammo-noids, belemnoids and benthos; (2) the ammonoid spectrum differ-entiates between perisphinctoids, Sowerbyceras, Phylloceratina

+Lytoceratina, haploceratids and other Ammonitina (Olóriz et al.,1995, 1996); and (3) the faunal spectrum for benthic macroinverte-brates differentiates between brachiopods, bivalves, regular echi-noids, irregular echinoids and others (gastropods, ahermatypiccorals and crustaceans).

Complementary parameters of palaeoecological interest were alsostudied: relative abundance, alpha diversity index at the genus level(Fisher et al., 1943) and relative dominance (equitability).

Stratigraphic analysis of the macro- and microfossil assemblageswas based on the differentiation of four intervals revealing a strati-graphic template that satisfied two requirements: (1) These strati-graphic intervals result from integrating palaeontological(composition of fossil assemblages, biochronostratigraphy) and strat-igraphic information (macro- and microfacies analysis); and (2) theintervals allow interpretation in terms of sequence stratigraphy by

image of Fig.�4


taking into account taphonomic and ecostratigraphic approaches.Thickness distribution in these stratigraphic intervals varies amongstthe Internal (more distal) and the External (more proximal) Prebetic,being more homogeneous in the former. Greater thickness was foundin eastern areas within the Internal Prebetic and in western areaswithin the External Prebetic. Thus we established: 1) Stratigraphic In-terval I (5.2–6 m in the Internal Prebetic and 6.7–7 m in the ExternalPrebetic) for the Transversarium Zone and the lower part of the Bifur-catus Zone, 2) Stratigraphic Interval II (5.6–6.4 m in the Internal Pre-betic and 3.9–5 m in the External Prebetic) for the upper part of theBifurcatus Zone, 3) Stratigraphic Interval III (2.7–3.1 m in the InternalPrebetic and 3–10.1 m in the External Prebetic) for the BimammatumZone, and 4) Stratigraphic Interval IV (1.3–1.6 m in the Internal Pre-betic and 3.3–4.6 m in the External Prebetic) for the Planula Zone.On this basis, ecostratigraphy was used to support sequence strati-graphic interpretations through a consensus approach assumingequivalent duration for Oxfordian biozones (e.g., Hardenbol et al.,1998; but also see Strasser, 2007 and Boulila et al., 2008). An

Fig. 5. Selected genus of Middle Oxfordian-Lower Kimmeridgian foraminifera from the PrebeF. Trocholina, G. Epistomina, H. Redmondoides, I. Ammodiscus, J. Ammobaculites, K. Reophax a

approximate time frame of 400 to 800 kyr for each of the studiedstratigraphic intervals is the reference order-hierarchy for corre-sponding sedimentary packages and related sequences.

4. Foraminiferal assemblages

4.1. Composition

Foraminifera recorded in the studied successions are mainly ben-thic (agglutinated, calcareous perforate and porcelaneous forms;Fig. 5) belonging to the suborders Textulariina, Lagenina, Spirillinina,Involutinina, Milionina and Robertinina. Planktic foraminifera belongto Globigerinina (Globuligerina).

The average foraminiferal assemblage is made up of vagile ben-thics (68%), followed by planktics (21%) and sessile benthics (11%).Vagile benthic foraminifera are principally agglutinated foraminifera(41%) and spirillinids (26%). Nodosariids and porcelaneous forms(mainly ophthalmidiids) are generally secondary. Sessile benthic

tic Shelf. A. Globuligerina, B. Spirillina, C. Laevidentalina, D. Lenticulina, E. Ophthalmidium,nd L. Thurammina.

image of Fig.�5


foraminifera are mainly nubeculariids, together with scarce siliceousagglutinated forms and other calcitic foraminifera.

Distal and proximal foraminiferal assemblages are differentiated,characterising relatively distal- (Internal Prebetic) and middle-shelf(External Prebetic) areas (Olóriz et al., 2003a):

The comparatively distal assemblage contains the greatest abun-dance of Globuligerina, Epistomina and encrusting nubeculariids. Vag-ile benthic foraminifera are dominated by uniserial agglutinatedforms, mainly Ammobaculites, and Ophthalmidium. The diversity ofbenthic foraminifera is greatest, and the largest average size isfound for Globuligerina, Lenticulina, uniserial nodosariids, Ammobacu-lites and Reophax. In terms of microhabitat, benthic foraminifera aredominated by epifaunal (46%) and shallow infaunal forms (31%),whereas potentially deep infaunal forms are scarcer (23%).

The relatively proximal assemblage contains lower proportions ofGlobuligerina, Epistomina and encrusting nubeculariids. The vagilebenthic assemblage is dominated by spirillinids, followed by uniserialagglutinated forms, mainly Reophax. Diversity is lower than in thedistal assemblage, while the size of Spirillina is greater than in the dis-tal assemblage. With respect to the microhabitat depth, epifaunalforms are clearly dominant (57%) and potentially deep infaunal(27%) and shallow infaunal forms (16%) are secondary.

4.2. Taphonomy

Taphonomic analysis focused on the relative significance of taph-onomic features such as fossil size, fragmentation and corrasion,microencrustation, microboring, sedimentary infilling and cementa-tion, and recrystallisation and mineralization of tests. Their relationto the lithofacies allows us to characterise three major taphofacies.

1) Foraminiferal Taphofacies I: Characterised by high microencrusta-tion and microboring in the biggest shells of nodosariids. Fragmen-tation is low and micritic sedimentary infilling of shells dominates.This taphofacies is registered in the condensed lumpy-oncoliticlimestone and lumpy-oncolitic limestone lithofacies.

2) Foraminiferal Taphofacies II: Characterised by common microen-crustation and microboring in the biggest shells of nodosariids.Fragmentation is higher than in foraminiferal taphofacies I, mainlyaffecting Lenticulina. Micritic infilling predominates. This taphofa-cies is registered in the spongiolithic limestone lithofacies.

3) Foraminiferal Taphofacies III: Characterised by a low incidence ofmicroencrustation and microboring as well as the highest valuesof fragmentation, especially in nodosariids and locally spirillinids(Reolid, 2008). Inside the chambers, cementation by sparite pre-dominates over the micritic infilling. This taphofacies, also charac-terised by frequent glaucony within chambers, is typical amongstthe marl-limestone rhythmite, spongiolithic marl-peloidal lime-stone and lumpy limestone lithofacies.

4.3. Stratigraphic distribution

Stratigraphic Interval I is characterised by the abundance of Globu-ligerina, mainly in the Internal Prebetic — i.e., relative distal-shelfareas (Fig. 6). The comparative abundance of sessile foraminifera in-creases in relation to the record of spongiolithic limestones (easternsector) and sponge bioherm-microbial lithoherm complexes (centralsector). Vagile benthic foraminifera in the External Prebetic – i.e., rel-ative middle-shelf areas – are dominated by spirillinids and aggluti-nated forms (mainly Reophax), whereas seawards – InternalPrebetic – they are dominated by agglutinated forms (mainly Ammo-baculites) and ophthalmidiids. Amongst vagile benthics, epifaunaltaxa present higher values in the Internal Prebetic (55%) than in theExternal Prebetic (47%) — i.e., a seaward-increasing record. Genus-level α-index of diversity for benthics is higher in the Internal Pre-betic (4.3) than in the External Prebetic (3.4), the lowest values

corresponding to the marl-limestone rhythmite. Throughout Strati-graphic Interval I, foraminiferal diversity diminishes in the ExternalPrebetic and increases in the Internal Prebetic.

Stratigraphic Interval II is mainly characterised by an accentuateddecrease in Globuligerina (Fig. 6). Compared with the previous strati-graphic interval, vagile foraminifera feature: 1) increasing values of ag-glutinated forms (Ammobaculites and Reophax) in the Internal Prebetic;2) decreasing proportions of ophthalmidiids in the Internal Prebetic;3) a near absence of Epistomina; 3) decreasing proportions of uniserialnodosariids; and 4) epifaunal forms diminishing in the Internal Prebetic(35%) while increasing in the External Prebetic (52%). Genus-level α-index diversity for the benthic forms is highly variable.

Stratigraphic Interval III contains vagile benthic foraminifera char-acterised by: 1) a lesser abundance in ophthalmidiids; 2) nodosariidsdominated by Lenticulina with respect to uniserial forms (except inthe RGCHSP); and 3) decreasing epifaunal forms (29% in the InternalPrebetic and 45% in the External Prebetic; Fig. 6). Proportions of ses-sile forms decrease landwards – External Prebetic – whereas nubecu-lariids are common seawards – in the Internal Prebetic. Benthicforaminiferal diversity depends on the genus-level α-index values,which are lower in the central sector of the External Prebetic (2.5)and higher in the Internal Prebetic (4.3).

Stratigraphic Interval IV comprises dominant vagile benthic fora-minifera while sessile benthics and planktics are scarce. In relativelyseaward sections – Internal Prebetic – proportions of Globuligerina in-crease close to the top of the succession, especially in the omissionsurface that terminates the succession (Fig. 6). Vagile benthic formsdecrease in proportions of spirillinids and agglutinated foraminiferawith respect to Stratigraphic Interval III. This trend is compensatedin the External Prebetic by increasing proportions of nodosariids(mainly Lenticulina) and a greater content of ophthalmidiids in the In-ternal Prebetic. As compared to the previous stratigraphic interval,the proportion of epifauna increases in the Internal Prebetic (33%)and the central sector of External Prebetic (53%), whereas in the east-ern sector of the External Prebetic it decreases (38%). Genus-level α-index diversity is 3.4 in the External Prebetic and 3.9 in the InternalPrebetic.

5. Macroinvertebrate fossil assemblages

5.1. Composition

The recorded assemblage of fossil macroinvertebrates (Fig. 7) isclearly dominated by seaward-increasing proportions of ammonoids(56% in the External and 78% in the Internal Prebetic), followed bybenthics showing a reversed trend (30% in the External and 16% inthe Internal Prebetic). Cephalopods such as belemnites and nautiloidsare secondary. Amongst ammonoids, there is a predominance of theperisphinctoid group (38%), followed by haploceratids (29%). Thescarce Phylloceratina+Lytoceratina are slightly more abundant inthe Internal Prebetic. The benthic macroinvertebrates are dominatedby brachiopods (47%), followed by bivalves (31% in the External and20% in the Internal Prebetic). Echinoids make up 12% in the ExternalPrebetic (7% irregular and 5% regular echinoids) and 24% in the Inter-nal Prebetic (20% irregular and 4% regular echinoids). Other benthicmacroinvertebrates are scarce. Sponges are mainly recorded in theExternal Prebetic and are characteristic of the spongiolithic lithofaciesgroup, where hexactinellids (Hexactinosida, Lyschniskosida and Lys-sacinosida) and lithistids were recognised (Olóriz et al., 2003b;Reolid, 2007, 2011).

5.2. Taphonomy

According to Olóriz et al. (2002b) the relative significance of tapho-nomic features (state of preservation in terms of size, within-bed posi-tion, corrasion, fragmentation, epibionts/encrustation, disarticulation,

Fig. 6. Stratigraphic distribution and relative abundance of main foraminiferal groups from Middle Oxfordian to Lower Kimmeridgian deposits in the Prebetic Shelf.


uncoupling and deformation) and their relation to the lithofacies pointsto three major taphofacies.

1) Taphofacies I (TF-I) is characterised by the greatest average size ofspecimens, the highest values of corrasion, encrustation and colonisa-tion, and a high proportion of specimens in a quasi-horizontalposition.

2) Taphofacies II (TF-II) shows a higher proportion of smaller speci-mens and a greater variety of azimuthal orientations (but thequasi-horizontal position is predominant); corrasion values are low.

3) Taphofacies III (TF-III) can be characterised by a predominance ofsmall specimens, the highest proportion of quasi-horizontal orien-tation, fragmentation and deformation, and the absence of corra-sion and encrustation.

5.3. Stratigraphic distribution

In each of the four stratigraphic intervals differentiated above, thedocumented trends show difference in taphonomy (Fig. 8) and

composition of macroinvertebrate assemblages (Fig. 9) that may re-flect a variable response to environmental conditions:

Stratigraphic Interval I is characterised by the upward increase ofproportions of ammonoids with respect to benthos (Fig. 9). Ammo-noid assemblages display high proportions of Sowerbyceras and Hap-loceratoidea throughout the interval, while the values of theperisphinctoid group increase. In the Internal Prebetic, benthos isdominated by brachiopods (Nucleata and Lacunosella; Reolid, 2005),with progressively increasing proportions of irregular echinoids(Holectypus and Collyrites) and secondary bivalves. In more proximalareas (External Prebetic), brachiopods dominate (Monticlarella andPlacothyris; Reolid, 2005) in spongiolithic limestones, decreasing pro-gressively with respect to bivalves (Isoarca, Spondilopecten and Pseu-dovola). In the marl-limestone rhythmite from the central sector ofthe External Prebetic, the benthic assemblage is dominated by infau-nal and vagile epifaunal bivalves (e.g., Procardia and Entolium). Sessileforms are locally abundant in association with sponge-microbialitebuildups (Olóriz et al., 2003b, 2006). Locally, this interval shows in-creasing corrasion, encrustation and colonisation by epibionts. In

image of Fig.�6

Fig. 7. Selected macroinvertebrates from the Middle Oxfordian-Lowermost Kimmeridgian in the Prebetic Shelf. A. Right-side view of inner-cast of incomplete ammonite phragmo-cone (Perisphinctes or Sequeirosia) from the lower Bifurcatus Zone, showing edge modification by lateral damage, surface alteration and pitting [condensed lumpy-oncolitic lime-stone; NV section], B. Left-side view of inner-cast of incomplete, slightly eroded ammonite phragmocone with slightly edge rounding (Taramelliceras [Metahaploceras]) from thePlanula Zone [lumpy-oncolitic limestone; NV section], C. Left-side view of inner-cast of ammonite phragmocone (Epipeltoceras) from the Bimammatum Zone, showing potentiallateral damage and local pitting [lumpy-oncolitic limestone; NV section], D. Right-side view of inner-cast of ammonite phragmocone with preserved beginning of the body-chamber (Ochetoceras) from the Bimammatun Zone, showing no significant macroscopic damage but limited edge modification and chalky-like appearance [lumpy-oncolitic lime-stone; NV section], E and F. Right lateral views of belemnite rostra (Hibolithes) from the Planula Zone [marl-limestone rhythmite; PC section], G, H and I. Dorsal and anterior views ofneomorphic shells of Bifurcatus Zone brachiopods showing deeply to slightly sulcate commissures (G. Nucleata; lumpy-oncolitic limestone; PL section), (H. Placothyris showingencrustating foraminifera [white arrows]; spongiolithic limestone; CH section) and (I. Monticlarella; spongiolithic limestone; CH section), J to M. Bivalves (J. Neomorphic, disarti-culated right valve of Arcomytilus from the Bifurcatus Zone; lumpy-oncolitic limestone; RS section), (K. Neomorphic, disarticulated and bored [white arrow] right valve of Isoarcafrom the Bifurcatus Zone; spongiolithic limestone; PC section), (L. Left view of Pholadomya [Procardia] with damaged anterior and posterior margins, from the Bimammatum Zone;marl-limestone rhythmite; RGCHSP section), (M. Right view of Goniomya showing broken anterior margin from the Planula Zone; lumpy limestone; PL section), (N and O. Moder-ately abraded spine (N) and fragment of interamb plates (O) of Plegiocidaris from the Bimammatum Zone; spongiolithic marl-peloidal limestone; PC section), P. Oral view of Diplo-podia from the Bimammatum Zone (spongiolithic marl-peloidal limestone; PC section), Q. Aboral view of Holectypus showing local collapse from the Bifurcatus Zone (lumpy-oncolitic limestone; NV section), R. Oral and apical views of locally broken test of Collyrites from the Bifurcatus Zone (lumpy-oncolitic limestone; PL section), S. Calcite epigenizedsiliceous sponge Verrucocoelia from the Bimammatum Zone (spongiolithic marl-peloidal limestone; PC section), T. Serpula (Cycloserpula) and Neovermilia (black arrow) colonisingsiliceous sponge from the Bifurcatus Zone (spongiolithic limestone; FA section, scale bar=1 mm). Section labelling as in Fig. 1.


image of Fig.�7

Fig. 8. Stratigraphic distribution of taphonomic features and taphofacies recognised in macroinvertebrate assemblages from Middle Oxfordian to Lower Kimmeridgian deposits inthe Prebetic Shelf.


the Internal Prebetic, TF-II is replaced by TF-I, whereas in the ExternalPrebetic TF-II is exclusive (Fig. 8).

Stratigraphic Interval II shows upwardly decreasing proportions ofammonoids in the assemblages, and the opposite trend for benthicforms (Fig. 9). The values of perisphinctoids diminish throughoutthe interval while percentages of Haploceratoidea and Sowerbycerasincrease. The composition of the benthic assemblage features a signif-icant diminution of brachiopods in relatively distal sectors, whereasproportions of Mytiloidea (Falcimytilus) and irregular echinoids(Holectypus) increase. In the spongiolithic limestone of the easternsector of the External Prebetic, the benthic assemblage records ahigher percentage of brachiopods (mainly Placothyris and Monticlar-ella) and a lesser amount of bivalves such as Isoarca, Procardia andEntolium. In the marl-limestone rhythmite of the central sector ofthe External Prebetic, bivalves dominate with Pectinoidea (Entolium,Propeamussium, Aequipecten) and Pholadomyoidea (Procardia).

Stratigraphic Interval II is also characterised by decreasing values ofcorrasion, encrustation and colonisation by epibionts, and TF-II is ex-clusive in both the Internal and the External Prebetic (Fig. 8).

Stratigraphic Interval III has noteworthy different trends related tochanges in lithofacies. Ammonoids show increasing proportions ofHaploceratoidea, and locally there are even high proportions of theperisphinctoid group. Aspidoceratidae become common in somelevels. Benthic assemblages from the Internal Prebetic are dominatedby brachiopods (Nucleata and Lacunosella) and irregular echinoids(Holectypus), while taxa typical of landward environments such assponges and Placothyris also occur (Reolid, 2005). Landwards, in theeastern sector of the External Prebetic, increasing terrigenous con-tents determine substitution of spongiolithic limestones by spongio-lithic marl-peloidal limestones, with a predominance of brachiopods(Dictyothyris and Monticlarella), irregular echinoids (Plegiocidaris)and more common cup-shaped sponges. The benthic assemblage of

image of Fig.�8

Fig. 9. Stratigraphic distribution and relative abundance of macroinvertebrate assemblages showing selected groups according to life-habit and trophic behaviour across the Pre-betic Shelf.


the marl-limestone rhythmite from the central sector of the ExternalPrebetic is dominated by bivalves such as Procardia and Entolium.Taphonomic traits indicate Taphofacies II.

Stratigraphic Interval IV shows an upward increase in proportionof ammonoids and belemnites. Amongst ammonoids, the increase ofHaploceratoidea is generalised, whereas the relative abundance ofthe perisphinctoid group is higher landwards (External Prebetic),and Sowerbyceras seawards (Internal Prebetic). Benthic assemblagesfrom outer sectors record diminution in brachiopods and increasingnumbers of bivalves and irregular echinoids. In relatively proximalsectors, characterised by the record of the marl-limestone rhythmite

and marls, Pectinoidea and irregular echinoids dominate, without oc-currence of sessile epifaunal forms (Fig. 9). Taphonomic features inthe Internal Prebetic correspond to TF-II, whereas in the External Pre-betic TF-II is replaced by TF-III (Fig. 8) with increasing proportions ofpyrite moulds, collapse fragmentation and plastic deformation, andthe absence of corrasion and encrustation.

6. Facies, stratigraphic intervals and ecostratigraphy

As widely recognised, the degree of time-averaging (e.g., Walkerand Bambach, 1971) is crucial for interpreting fossil assemblages,

image of Fig.�9


especially in biostratigraphy and palaeoecology. Fossil assemblagesstudied in Upper Jurassic epicontinental shelf deposits of the South-Iberian palaeomargin are considered to show primarily within-habitat time averaging, and less commonly environmental condensa-tion as deduced from biostratigraphic and taphonomic analyses(Rodríguez-Tovar, 1993; Olóriz et al., 1999, 2002b; Reolid, 2003;Olóriz et al., 2004a; Reolid, 2008; see Olóriz, 2000 for comprehensivediscussion). That is, fossil assemblages should represent skeletal accu-mulations derived from local subtropical communities averaged inthe range of one to some 102 years to 104 years according to estima-tions proposed by Kidwell and Bosence (1991) for within-habitattime-averaging and environmental condensation. Data relevant to es-timations of mean time-averaging in shelly assemblages from recentshelves (ca. 104 years according to Flessa and Kowalewsky, 1994;widely variable ranges even for close settings are reported byBarbour-Wood et al., 2003; Carroll et al., 2003; Olszewski, 2004;Tomašových and Zuschin, 2009) suggest that palaeoecologic analysisbe aimed at long-term ecology. This is especially interesting when weare interpreting macroinvertebrate fossil assemblages includingmegabenthos from epicontinental shelf environments. According toOlóriz (2000), long-term palaeoecology and biostratigraphy can beapproached through fossil assemblages made up of ammonites andbenthic macroinvertebrates in Upper Jurassic deposits. On the as-sumption of ecological fidelity for skeletal remains (e.g., Kidwell,2002) and homotaxy, long-term palaeoecological interpretationsbased on ecostratigraphic trends would be particularly appropriatein contexts where time-averaging reflects both the long-term envi-ronmental conditions (e.g., Olszewski, 1999) and the relative abun-dance of species-level and higher taxa (e.g., Kidwell, 2001).

6.1. Stratigraphic Interval I

Deposits belonging to Stratigraphic Interval I rest, alternatively, onLower-to-Middle Jurassic shallow carbonates with thin iron-ooliticcrusts that represent plinthitic palaeosoils developed in coastal plainswhen the Middle Jurassic shallow shelf was emerged (Reolid et al.,2008c). The oldest record of marine Late Jurassic deposits is locallyrepresented by a cephalopod-rich shell bed revealing rapid floodingof the shelf under high energy conditions (Olóriz et al., 2008; Reolidet al., 2008c).

The lower part of Stratigraphic Interval I (Transversarium to lowerpart of Bifurcatus zones) is characterised by lumpy lithofacies in sea-ward settings – Internal Prebetic – whereas it is dominated land-wards by spongiolithic limestones (eastern sector of the ExternalPrebetic) and marl-limestone rhythmites with microbial-spongebuildups (central sector of the External Prebetic). Sponge-microbialbuildups are registered throughout the Transversarium and lowerpart of Bifurcatus Zone in outcrops of the central sector, and locallyin the eastern sector of the External Prebetic (Reolid, 2007, 2011). Ex-cept for the marl-limestone rhythmite, increasing condensation up-wards is typical, especially seawards, where condensed lumpy-oncolitic limestones are present in comparatively distal areas.

The foraminiferal assemblage is characterised by high values ofencrusting and planktic forms as well as the occurrence of Epistominain relatively distal-shelf areas. Amongst benthic foraminifera, infaunalspecimens decrease seawards (Internal Prebetic). Previous evidencefor correlation between higher proportions of Globuligerina and Epis-tomina and longer distance from shore and larger ecospace has beeninterpreted by Henderson and Hart (2000), Oxford et al. (2000,2002), Reolid and Nagy (2008) and Reolid et al. (2008a), amongstothers.

Upwards in Stratigraphic Interval I, the macroinvertebrate assem-blage shows increasing proportions of ammonoids (more marked inrelatively distal settings), and progressively decreasing values of ben-thic infauna and semi-infauna. This situation agrees with enhancedconditions for nektonic as well as for sessile epifaunal forms

(enlarged and/or ameliorated ecospace), the latter being favouredby a higher cohesion of the substrate while within-substrate ecospaceshrank.

Taphofacies II dominates landwards. Seawards, the upper part ofStratigraphic Interval I shows the shift from Taphofacies II to Taphofa-cies I through increasing corrasion, encrustation and colonisation ofskeletals by epibionts; this trend is related to longer exposure of skel-etal remains on the sea-bottom and lower sedimentation rates. Olórizet al. (2004b), Reolid and Gaillard (2007) and Reolid (2008) found in-creasing values of microborings and microencrusters in analysis ofmicrotaphonomic traits in this interval.

Two reference horizons serve to identify the upper part of Strati-graphic Interval I. The lower one is revealed by an increased occur-rence of both cephalopods and sessile epifaunal suspension-feeders.These features are better recorded seawards, where a relative impov-erishment in sedimentivores points to changes in substrate condi-tions, most probably reflecting the change to sea-bottoms enrichedin shelly remains and microbial encrustations (Reolid et al., 2005).The second reference horizon is a boundary horizon separating Strat-igraphic Interval I from the overlying Stratigraphic Interval II, identi-fied through inflection points in the registered trends forsedimentary and substrate conditions, taphonomic traits, a relativeabundance of major components of ecologic groups, and ecospacethroughout the sectors studied in the Prebetic shelf system (Figs. 8and 9). This inflection point in registered trends is coincident withthe last appearance of sponge-microbial buildups in the central sectorof the External Prebetic (Olóriz et al., 2003b; Reolid, 2011).

6.2. Stratigraphic Interval II

Stratigraphic Interval II (upper part of Bifurcatus Zone) is charac-terised by decreased stratigraphic condensation, which is well markedin seawards sections (Internal Prebetic), where the condensed lumpy-oncolitic limestones is replaced by lumpy-oncolitic limestone. Themarl-limestone rhythmite evidences a progressive disappearance offavourable conditions for sponge-microbialite buildup growth.

Foraminiferal assemblages feature a diminution in sessile forami-nifera, planktic forms and Epistomina, revealing shifting ecospaces.The record of infaunal foraminifera shows opposing trends, theirabundance increasing seawards (Internal Prebetic) and decreasinglandwards (External Prebetic).

The presence of ammonoids is seen to decrease with respect to ben-thic macroinvertebrates (Fig. 9). Amongst ammonoids, the percentageof the perisphinctoid group diminishes upwards, whereas Haplocera-toidea and Sowerbyceras increase. The macrobenthic assemblage ischaracterised by a slight decrease in sessile forms (brachiopods andsponges) as opposed to infaunal and semiinfaunal bivalves and irregu-lar echinoids. This picture is coherent with a higher sedimentationrate, as well as with lower corrasion, encrustations and epibionts, indi-cating minor exposure of skeketals on the sea-bottom. Comparativelysoft substrates are also revealed by a greater degree of plastic deforma-tion and variable orientation of skeletals within the sediment, which iscongruentwith increasing amounts of infaunal forms. In Stratigrahic In-terval II, Taphofacies II is exclusive in all the sectors studied within thePrebetic shelf system (Fig. 8).

The boundary horizon separating Stratigraphic Interval II fromthe overlying Stratigraphic Interval III is comparatively subtle. Closeexamination reveals changes affecting substrate rather than water-column conditions. Ameliorated conditions for both infaunal sedi-mentivores in landwards settings and epifaunal suspensivores sea-wards – most probably forcing a relative inhibition of the infauna inrelatively distal substrates – may be indicative of a breaking-pointin the eco-sedimentary framework. The absence of records revealingparallel effects in water-column ecospace, such as fluctuations incephalopods vs. benthics or relevant fluctuations in proportions ofammonoid groups (Fig. 9), points to rather unconnected dynamics,


which would result from a lower level of environmental change af-fecting the shelf system. The lack of relevant changes in taphonomictraits reinforces the interpretation of a comparatively low level of en-vironmental change; hence, the assumption of minor change in thetotal ecospace across the sectors studied in the Prebetic shelf system.This horizon separating Stratigraphic Intervals II and III is thereforeinterpreted as an ecostratigraphic event of special significance forsubstrate ecology.

6.3. Stratigraphic Interval III

Stratigraphic Interval III (Bimammatum Zone) is characterised byincreased terrigenous content, mainly in more proximal settings.The spongiolithic limestone of the External Prebetic is generallyreplaced by spongiolithic marl-peloidal limestones and the marl-limestone rhythmite.

Clear differences in foraminiferal assemblages are recognised be-tween the relatively proximal and distal areas. Foraminifera are scarcerlandwards, and planktics and sessile benthics are rare to absent. Sea-wards, microfauna are more abundant than in the underlying Strati-graphic Interval II, and characterised by a greater percentage of sessileforaminifera and agglutinated forms (Ammobaculites and Reophax).Amounts of infaunal foraminifera increase across the shelf, as evidencedby their record in both the External and Internal Prebetics.

Macroinvertebrate assemblages are characterised by significantfluctuations in the ammonoid/benthos ratio, pointing to changes inenvironmental conditions and lithofacies. Ammonoids show higheroccurrence of Haploceratoidea and Aspidoceratidae. Benthic assem-blages reflect decreasing sessile suspension-feeders landwards. Indistal-shelf areas (Internal Prebetic), the typical macrobenthic assem-blage (e.g., Nucleata, Lacunosella, Falcimytilus) includes taxa commonin proximal settings (e.g., Monticlarella, Placothyris and siliceoussponges), thus indicating a relatively homogenising phase and com-parative eurytopy.

Lower levels of corrasion, encrustation and colonisation by epi-bionts are characteristic of landward settings, which received highersedimentation and burial rates resulting from increased inputs in sili-cilastics. Taphofacies II is exclusive in the sectors studied in the Pre-betic shelf system.

Identification of the boundary horizon separating Stratigraphic In-terval III from Stratigraphic Interval IV is reinforced by the definitivedisappearance of environmental conditions that had previouslyfavoured the persistence of lumpy-oncolitic sediments in seawardsettings (Internal Prebetic), and of sponge meadows and the resultingspongiolithic limestones in eastern settings landwards (eastern Ex-ternal Prebetic). Complementary information revealing this boundaryhorizon between Stratigraphic Intervals III and IV seawards is therecorded increase in proportions of nekto-planktic and bottom-levelinvertebrates, including infauna. Since the registered increase affect-ed water column inhabitants as well as substrate colonisers, this isinterpreted as an ecostratigraphic event related to ecospace enlarge-ment in the distal shelf.

Landwards in the central sector of the External Prebetic, no partic-ular horizon is well recorded, and no relevant changes in water col-umn conditions can be inferred, at least in terms of their potentialincidence on the abundance of common, major nekto-planktic inver-tebrate components. Again, the only identifiable difference would bein substrate conditions, determining ecospace amelioration for infau-na and the reverse for sessile epifauna in these landward settings.

Differential ecostratigraphic signals according to distance from theshore would indicate environmental variations that were betterexpressed in water-column ecospace. In contrast, the comparatively in-distinctive ecostratigraphic signal registered at the substrate levelacross the shelf, with no difference in the relative abundance of thetwo major trophic groups (suspension-feeders and sediment-feeders),is of particular importance (Olóriz et al., 2006). Environmental

dynamics therefore caused some amelioration of within-substrate con-ditions throughout the shelf, togetherwith subtle changes in burial con-ditions and preservation.

6.4. Stratigraphic Interval IV

The upper Stratigraphic Interval IV (Lower Kimmeridgian, PlanulaZone) is characterised by a generalised and strong increase in terrig-enous content. Landwards, the marl-limestone rhythmite and marlsdominate in the External Prebetic, while a comparatively subtle in-crease in siliciclasts was registered seawards, entailing quartz grainsand the disappearance of oncoids of nubeculariids in the Internal Pre-betic (Reolid et al., 2005). At the top of Stratigraphic Interval IV, anomission surface – locally a hardground – is present, underlying sev-eral metres of dark-grey marls.

Foraminiferal assemblages are characterised by severe diminu-tion, or even absence, of nubeculariids and planktic foraminifera. Re-lated to the omission surface, the proportions of Globuligerina andEpistomina increase. The occurrence of thick-shelled specimens ofLenticulina with high degrees of fragmentation indicates transportfrom landward settings (Reolid, 2008).

There is a greater relative abundance of ammonoids and belemnites,while the proportion of benthos diminishes. Amongst ammonoids, theproportion of Haploceratoidea increases overall, perisphinctoids in-crease in proximal settings, and Sowerbyceras in distal-shelf areas. In-faunal bivalves and irregular echinoids are abundant at the expense ofsessile epifauna.

The state of preservation in macroinvertebrates would point toTaphofacies II and Taphofacies III (absence of corrasion, encrustationsand epibionts, and abundance of fragmentation by collapse) as de-scribed by Olóriz et al. (2002b).

The sharp upper boundary of Stratigraphic Interval IV is a clear,reference horizon for the sections studied. In ecostratigraphic terms,it corresponds to a sudden increase in siliciclastics that affected envi-ronmental conditions of the substrate and the water column. There-fore, a distinct environmental shift is envisaged in relation to atectonic pulse (e.g., Marques et al., 1991), registered as an ecostrati-graphic event entailing ecological deterioration across the shelf.

7. Sequence stratigraphic interpretation

Local physiography and tectonics, climate, hydrography, sedimenta-ry input, and the interaction of such forcing factors determine local sed-imentary packages as well as the stratigraphic record of a particulardepositional sequence in shelves under dominant siliciclastic deposi-tion. Carbonate productivity and sediment production (exportation)proves to be a relevant factor in carbonate shelves, but its influence inmixed carbonate–siliciclastic shelves can be minor depending on shelfphysiography and hydrography. In cases of comparatively low potentialfor recording changes in depositional conditions (e.g., relatively distaland/or deeper settings, and settings subjected to persistent fine grainedclastic-carbonate deposition), the macroscopic, physical aspect andcharacterization of stratigraphic surfaces of value for sequence stratigra-phy can be meagre. In addition, variability in time duration assumed forso-called 3rd cycles mainly driven by global-eustasy and/or other forc-ing (e.g., Posamentier and James, 1993;Miall, 1997) are alsomost prob-ably combined with, and/or forced, by potential difficulties for precisetime evaluations (e.g., Nummedal, 2004; Boulila et al., 2011). Hence, aprecise outcrop level correlation of depositional sequences is commonlyobscured by the wide range of duration assigned to these 3rd order cy-cles (0.1 to 10 My; e.g., from Vail et al., 1977; Haq et al., 1988 and Rey etal., 1988 to Kendall et al., 1995; Strasser et al., 2000; Olóriz et al., 2003c;and Ogg and Przybylski, 2006) and the differential interpretation ofbounding discontinuities and correlative conformities (e.g., fromMitchum et al., 1977 to Posamentier and Vail, 1988; Posamentier et al.,1988; Van Wagoner et al., 1988; Galloway, 1989; Mitchum and Van


Wagoner, 1991; Embry and Johannessen, 1992; Hunt and Tucker, 1992;Embry, 1993; Helland-Hansen and Gjelberg, 1994; Embry, 1995;Catuneanu et al., 1998; Posamentier and Allen, 1999; Plint andNummedal, 2000; Catuneanu, 2006; Christie-Blick et al., 2007; Embryet al., 2007; Catuneanu et al., 2010; Bhattacharya, 2011). In generalterms, potential correlation of a particular depositional sequence relieson distance from shore and differential eco-sedimentary scenarios re-garding the geological histories of depositional areas under analysis. Ac-cordingly, correlation is more limited amidst smaller cycle orders (i.e.,related directly to shorter duration processes). While beyond thescope of the present study, the correlation potential of smaller-scale se-quences interpreted as derived from allocyclic remote forcing in theMilankovitch band (e.g., Pittet and Strasser, 1998a,b; Pittet et al., 2000;Rodríguez-Tovar et al., 2010) clearly deserves attention in the future.

In the nearby areas of the S-SE palaeomargin of Iberia studied, theMiddle Oxfordian to Lower Kimmeridgian succession shows uncom-mon, distinct expressive stratigraphic surfaces prone to correlation.Nonetheless, a strict biostratigraphic control of lateral and stratigraphicdistribution of lithofacies, taphofacies and micro- and macrofossil as-semblages led us to differentiate the four stratigraphic intervals de-scribed above. Their interpretation in terms of sequence stratigraphyis founded on the ecostratigraphic interpretation of their boundary ho-rizons, which are distinctive ecostratigraphic events separating ecostra-tigraphic trends represented by the bulk of the described intervals. Theduration of these events is difficult to determine, but it may be less thanone tenth the timespand of ecostratigraphic trends discerned accordingto taphonomic and biostratigraphic information.

Olóriz et al. (2008) interpreted the beginning of Late Jurassic pe-lagic–hemipelagic deposition in the central sector of the PrebeticShelf as related to short-time, rapid eco-sedimentary changesrecorded as high-energy ecostratigraphic events, demonstrating thefirst occurrence of pelagic macro-invertebrates resting on Lower–Middle Jurassic shallow carbonate-shelf deposits. Later and persistentlower-energy conditions in the rest of the Middle and throughout theLate Oxfordian were interpreted as the onset of a long-lasting ecostra-tigraphic trend, with a sharp change in lithofacies revealing a severedrop in environmental energy; that is, a low-energy ecostratigraphicevent.

The Middle Oxfordian to Lower Kimmeridgian stratigraphic inter-vals analysed above can be associated with ecostratigraphic trendsand their fluctuations, and so lead to a better understanding of theMiddle Oxfordian to Lower Kimmeridgian succession in terms of se-quence stratigraphy.

Fig. 10. Stratigraphic interpretation and correlation. (A) Standard biochronostratigraphy forareas], (C) Standard sequences [Tethyan areas], (D) 3rd order cycles, (E) 3rd order sequenceof backsteeping [Bck] and foresteeping [fst: initial; FST: later] phases, ecostratigraphic eventing; bwc: bottom and water column], (G) Stratigraphic intervals described, (H) System trabased on Hardenbol et al., 1998 [brackets for approximate ages based on Haq et al., 1988Ages according to Gradstein et al. (2004). Tectonic pulse, forced foresteeping phase and pr(1988), J and K from Marques et al. (1991).

7.1. Stratigraphic Interval I

Sediments belonging to this stratigraphic interval rest on ferrugi-nous oolitic limestone related to the widespread drowning that char-acterised the beginning of Late Jurassic sedimentation on Tethyanshelves (Ziegler, 1962; Goy et al., 1973; Gygi, 1981; Gygi andPersoz, 1987; Huber et al., 1987; Aurell et al., 1994; Courville andCollin, 1997; Ramajo and Aurell, 1997; Scouflaire et al., 1997; Aurellet al., 1999; Ramajo et al., 2002; Lorin et al., 2004; Collin et al.,2005; Meléndez et al., 2005; Ramajo and Aurell, 2008). In the studiedarea, its base is related to a complex unconformity (García-Hernándezet al., 1989; Marques et al., 1991; Reolid et al., 2008c) and interpretedas directly overlying a context of high-energy ecostratigraphic eventsand related facies (Olóriz et al., 2008). In the absence of intercalated,distinctive surfaces, the turnover to lower-energy characterising theonset of Stratigraphic Interval I conditions (in contrast to underlyingdeposits) was a comparatively rapid event. The stratigraphic recordof a sharp replacement of high-energy conditions clearly underlinesan ecostratigraphic event.

Lithological and palaeontological features characterising Strati-graphic Interval I are compatible with the combined effect of increas-ing distance from shore, together with progressively lowersedimentation rates. Such a scenario points to a backstepping phaseearly on in the Late Jurassic (Middle Oxfordian), well documentedelsewhere in Europe as transgressive conditions within a major trans-gressive–regressive cycle (e.g., from Haq et al., 1988 to Hardenbol etal., 1998). The Transversarium proparte and part of the Bifurcatuszones would correspond to a lower part of the OX.II sequence pro-posed by Marques et al. (1991) and included in the 3rd-order cycle4.3 interpreted by Haq et al. (1988) (Fig. 10). Increasing stratigraphiccondensation upwards in distal-shelf settings (Internal Prebetic)throughout Stratigraphic Interval I is interpreted as evidence of a con-densed section, resulting in reduced thickness of lower BifurcatusZone deposits. As expected, local environmental conditions repre-sented by Stratigraphic Interval I in the studied Prebetic Shelf contrastwith those reported from European shelves as a result of the variableorder of proposed sequences (Hardenbol et al., 1998; Pittet et al.,2000; Allenbach, 2001; Aurell et al., 2003).

The higher abundance of ammonoids, planktic foraminifera andEpistomina throughout Stratigraphic Interval I would reflect increasingecospace, whereas lower nutrient levels could be responsible for theprogressive impoverishment in infaunal foraminifera seawards (Inter-nal Prebetic; Figs. 11 and 12). This interpretation is compatible with

southwestern Europe, (B) Standard transgressive-regressive facies cycles [T-R; Tethyans, (F) Long-term sea-level curve and interpreted local relative sea-level with indications [low-energy: ; high-energy: Y], and ecospace fluctuations [E/e: increasing/decreas-cts [grey shading for condensed sections; e-: early; l-: late], (I) Geochronology in My], (J) Wide-correlation discontinuities in Iberia, (K) Depositional sequences, and (L)ogradation [* FFst]. B, C and I from Hardenbol et al. (1998), D, E and F from Haq et al.

image of Fig.�10

Unlabelled image


increasing distance from shore and the consequent lower nutrient inputfromnearshore environments. The sedimentation ratewas lowwith re-spect to accommodation, resulting in increasing condensation sea-wards. The reduced thickness of this stratigraphic interval in theInternal Prebetic agrees with the course of the maximum of the poten-tial transgressive pulse on-shore. Landwards, local occurrences ofhigher terrigenous inputs (e.g., central sector in the External Prebetic)have been related to preferential accumulation onmore depressed bot-toms that resulted in marl-limestone rhythmites. Upwards in this strat-igraphic interval, the comparatively lower sedimentation rate favouredsponge-microbialite buildups' growth in these relative low-energyareas (Olóriz et al., 2003b). Leinfelder et al. (1994) and Reitner and

Fig. 11. Simplified stratigraphic distribution of lithofacies and fossil assemblages (large pietransect, according to the stratigraphic intervals identified.

Neuweiler (1995), report that sponge-rich facies and buildups are com-mon during 3rd-order transgressive pulses.

7.2. Stratigraphic Interval II

The stratigraphic horizon that serves to separate Stratigraphic In-tervals I and II reveals changes in eco-sedimentary conditions affect-ing previous trends in substrate traits, taphonomy, and the relativeabundance of major ecologic groups (Figs. 11, 12), signalling ecospacefluctuation across shelf areas of the sections studied. Ammonite bio-chronostratigraphy allows correlation of this ecostratigraphic eventwith the correlative conformity corresponding to the discontinuity

diagrams: macroinvertebrates; small pie diagrams: foraminifera) in a proximal-distal

Fig. 12. Evolution of palaeoenvironmental parameters and relative sea-level changes according to lithofacies, composition of foraminiferal and macroinvertebrate assemblages, andtaphonomic features. Relative sea-level curve as in Fig. 10.


surface dV interpreted by Marques et al. (1991) (Figs. 10 and 13). Themaximum condensation degree shown in the upper part of the un-derlying Stratigraphic Interval I in distal-shelf environments pointsto condensed section deposits (Fig. 13), evidencing final maximumflooding relative to initial forestepping conditions for the beginningof Stratigraphic Interval II. These changes would therefore constitutean ecostratigraphic event.

Lithologic and palaeontologic features in Stratigraphic Interval IIagree with a relative landward shift of depocenters, under increasinginputs and sedimentation rates. This suggests initially limited forest-epping and progradation, a context compatible with the developmentof early-highstand system tract conditions (highstand normal regres-sive deposits sensu Catuneanu et al., 2009) during the late BifurcatusChron (Figs. 11–13).

Decreasing distance from shore and shallowing led to ecospace dim-inution and ecologic stress for ammonoids and planktic foraminifera

(impoverished records; Figs. 11 and 12). In addition, it gave rise to in-creasingfine terrigenous input upwards in the Bifurcatus Zone deposits,as evidenced by shalier sediments and the diminution of microbialites,sponges and sessile suspension-feeders, marking ecological stress inthe bottom level community as well. The increased record of infaunalforaminifera landwards implies more organic matter within the sub-strate (Reolid et al., 2008a,b), coming from the proximal shelf factorywhere high primary photosynthetic production took place (Olóriz etal., 2006). The relationship between terrigenous inputs and nutrientshas been established for Late Jurassic epicontinental shelves elsewhere(Insalaco, 1999; Bouhamdi et al., 2001; Pittet and Mattioli, 2002;Bartolini et al., 2003; Olivier et al., 2004). All of this is consistent withthe relative sea-level fall during Highstand System Tract (HST) stages.

Olóriz et al. (2002b) interpreted progressing HST conditions (i.e.,progressive shallowing) within the OX.II sequence as proposed byMarques et al. (1991), as a part of the 3rd-order cycle 4.3 (Haq et

image of Fig.�12

Fig. 13. Sequence interpretation. Section labelling as in Fig. 1.


al., 1988). The interpreted relative decrease in accommodation fitswell with the context described by Haq et al. (1988) and Hardenbolet al. (1998) for the upper Bifurcatus Zone deposits. The latter authorsproposed a smaller-order sequence boundary, analogously identifiedby Pittet et al. (2000); yet local conditions reported from otherEuropean shelves point to transgressive pulses (e.g., upper part ofthe Bifurcatus Zone in the Swiss Jura; Allenbach, 2001).

7.3. Stratigraphic Interval III

The relatively subtle boundary horizon separating StratigraphicInterval II from the overlying Stratigraphic Interval III reveals changesthat affect substrate rather than water-column conditions. The turn-ing point in the eco-sedimentary framework would therefore havebeen a widespread bottom-level process with no major change inburial and preservation conditions. Accordingly, limited environmen-tal changes affected the studied areas of the Prebetic shelf system. Itsrecord, then, is interpreted as an ecostratigraphic event related tolocal smoothing (masked appearance) of the type II discontinuityDVI envisaged by Marques et al. (1991) for the Iberian subplate(Figs. 10 and 13), with no evidence of shelf-margin sedimentarypackages in the area of study. Lacking traces of flooding conditions,the interpreted ecostratigraphic event is more suggestive of the se-quence interpretation by Haq et al. (1988) than that of Hardenbol etal. (1998) and other proposals, yet there is a noteworthy overall cor-relation of lower relative sea-level conditions between transgressive

peaks. Variable regressive conditions related with BimammatumChron tectonic instability have been postulated for NE Iberia (Aurellet al., 2003) and the Swiss Jura shelves (Allenbach, 2001).

Lithologic and palaeontologic data in Stratigraphic Interval III maybe interpreted within the context of increasing terrigenous inputs,even greater in landward settings. High abundance of sponge remainsin the Internal Prebetic would agree with biofacies progradation. Ter-rigenous inputs likewise transported organic matter from emergedand shallower areas to the studied sectors (Fig. 11), giving rise tothe increase in infaunal foraminifera (Fig. 12). All these data are con-gruent with decreasing distance from shoreline and accentuated rel-ative sea-level fall. Developing shelf margin wedge (SMW)conditions at the beginning of the OX.III sequence, in correspondencewith the 4.4. cycle (Haq et al., 1988), were suggested by Marques etal. (1991) and Olóriz et al. (2002b). Alternatively, the scenario pro-vided by late-highstand system tract conditions adapts well to the en-vironmental dynamics (forestepping) proposed here as affectingmid- to distal-shelf areas in the Prebetic shelf system.

7.4. Stratigraphic Interval IV

The ecostratigraphic event that served to separate StratigraphicIntervals III and IV – entailing a disappearance of lumpy-oncolitic sed-iments seawards and siliceous sponge meadows landwards, accom-panied by ecospace enlargement for nekton and plankton, as well as

image of Fig.�13


benthic community reorganisation – is interpreted as indicating a re-turn to backstepping conditions.

Ammonite biochronostratigraphy allows us to correlate this ecos-tratigraphic event with the dVII transgressive surface proposed byMarques et al. (1991) and Olóriz et al. (2002b). As previously stated,the present approach favours within-shelf dynamics (late-highstandsystem tract) instead of the shelf-margin-wedge context when inter-preting underlying Stratigraphic Interval III.

Early proposals for global cycle charts (Haq et al., 1987, 1988) en-visaged Planula Zone deposits as resulting from transgressive systemtract (TST) conditions pertaining to the OX.III sequence of the 4.4 3rd-order cycle, the condensed section being placed close to the Planula/Platynota zone boundary. Later approaches also interpreted PlanulaZone deposits as compatible with backstepping within increasing ac-commodation contexts, to a variable degree, with similar interpreta-tions of maximum flooding ages (Hardenbol et al., 1998; Pittet andStrasser, 1998a; Pittet et al., 2000; Aurell et al., 2003; Ogg andPrzybylski, 2006). Furthermore, Marques et al. (1991) described in-teractions between tectonics and eustasy in the South-Iberian palaeo-margin during latest Oxfordian to earliest Kimmeridgian times, andascribed the increase of siliciclastics to reactivated topography inthe source area (see also Aurell et al., 2003 for NE Iberia, and compat-ibility with carbonate production models in Pittet et al., 2000 for SEIberia, and Allenbach, 2002 for the Oxfordian in the Swiss Jura). Evi-dence of analogous situations affecting Upper Oxfordian depositswas later reported from other areas in Iberia, epicontinental Europe,and the north-central Mexico–northern Gulf Rim, related to the evo-lution of central-North Atlantic and the West-Tethyan basins(Goldhammer, 1998; Leinfelder and Wilson, 1998; Allenbach, 2001;Bádenas and Aurell, 2001; Salas et al., 2001; Allenbach, 2002; Olórizet al., 2003c).

In mid-shelf areas (External Prebetic), the influence of tectonic re-juvenation would be more evident (Figs. 11 and 12), as revealed byincreased terrigenous input and sedimentation rates (includinghigher proportions of quartz grains). Skeletals underwent shorter ex-posure and, therefore, lower degrees of encrustation and corrasion.Relatively higher sedimentation rates favoured the occurrence ofboth hypoxic microenvironments (pyrite in the inner whorls of am-monoid remains) and geopetal infillings of shell cavities. Compara-tively soupy grounds favoured plastic deformation and collapsefragmentation of embedded carcasses. In ecological terms, increasingterrigenous input was unfavourable for sessile benthic forms.

In distal-shelf areas (Internal Prebetic), the terrigenous increasewas less evident, and the effect of eustatic sea-level rise was greater,as indicated by the record of lumpy limestones with omission sur-faces and hardgrounds (Figs. 11–13). The lower sedimentation ratefavoured the concentration of skeletals, mainly ammonoids. Decreas-ing benthic macroinvertebrate contents are congruent with enlargingaccommodation and ecospace, also recognised in the record of nekto-planktic macroinvertebrates. Omission surfaces enriched in Globuli-gerina and Epistomina would support a relative backstepping effect.

The interpretation of Stratigraphic Interval IV in terms of sequencestratigraphy fits with a widely accepted “global” rising sea-level(from Haq et al., 1988 to Hardenbol et al., 1998 and later updatedby Ogg and Przybylski, 2006). Thus, transgressive conditions andbackstepping also accord with the TST as interpreted by Marques etal. (1991) within the OX.III sequence (Fig. 10). The sharp upperboundary of the Stratigraphic Interval IV constitutes a clear referencehorizon associated with the “Final Oxfordian Crisis” interpreted byMarques et al. (1991) as widespread pulses of tectonic instabilityclose to the Oxfordian/Kimmeridgian boundary (citations above). Inecostratigraphic terms, it translates to a sudden increase in siliciclas-tics that affected environmental conditions of both the substrate andthe water column. Therefore, a distinct environmental shift is envis-aged in relation to a tectonic pulse and registered as an ecostrati-graphic event implying ecological deterioration all across the shelf.

8. Conclusions

Lithology, biostratigraphy and taphonomy aid in interpretation ofthe eco-sedimentary dynamics relative to stratigraphic intervals identi-fied in the Middle Oxfordian–Lower Kimmeridgian succession investi-gated in the Prebetic Shelf (SE Iberia). Selected macro- and microfossilassemblages provide information about environmental conditions inthe water column and of the water-sediment interphase. In additionto shedding light on the composition of fossil assemblages, taphonomyreflects a variable range of sedimentary conditions affecting skeletal re-mainsmainly derived from cephalopods (ammonites, belemnites), bra-chiopods, bivalves, echinoids, sponges, and foraminifers.

Middle Oxfordian to Early Kimmeridgian (Transversarium to Plan-ula chrones) rocks and faunal assemblages in comparatively distalsectors (distal shelf) are characterised by lower sedimentation rates(resulting in lumpy carbonate lithofacies), and higher proportions ofammonoids and planktic foraminifera, higher degrees of corrasion,microboring and encrustation. Landwards, towards the mid-shelf,eco-sedimentary conditions resulted in spongiolithic limestones andmarl-limestone rhythmites with local development of microbial-sponge buildups.

Ecospace shifts were related to changing relative sea-levels (accom-modation, backstepping, forestepping) during the Late Jurassic in theepicontinenal area analysed, and can be recognised in palaeogeographicand stratigraphic trends as expressed through selected parameters(litho- and microfacies, faunal composition, and taphonomy).

Increasing distance from shore during relative sea-level highs isreflected in greater: (1) stratigraphic condensation; (2) abundancein ammonoids, planktic foraminifera and nubeculariids; and, (3) cor-rasion, microboring and encrustation, except for landward settingssubmitted to higher than average sedimentation rates (e.g., de-pressed bottoms submitted to lower energy and/or tectonic rejuvena-tion). These trends in faunal composition and taphonomy agree withbackstepping phases, increasing ecospace and longer exposure ofshelly remains on the sea bottom. Such conditions prevailed in theTransversarium to lower Bifurcatus Zone deposits of the central Pre-betic, then returned during the Planula Chron, with a local increasein sedimentation rate landwards.

Decreasing distance from shore during relative sea-level lows isrelated to opposite trends, evidenced by: (4) increasing terrigenousinput and decreasing stratigraphic condensation; (5) impoverish-ment in ammonoids and planktic foraminifera; (6) sponge-biofaciesprogradation; and (7) diminution of corrasion, microboring and en-crustation. Phases of forestepping and/or aggradation, reduction ofecospace for nekto-planktic organisms, and comparatively rapid buri-al of shell remains are interpreted to force the recorded trends. Theseconditions dominated Upper Bifurcatus Zone deposits and were ac-centuated during the Bimammatum Chron in the central Prebetic.

Changing relative distance from shore and ecospace, triggered byrelative sea-level fluctuations, are considered prime factors forcingtrade-offs in faunal communities corresponding to the analysed fossilassemblages. When the record of well defined surfaces relevant forsequence stratigraphic interpretations is lacking, the ecostratigraphicapproach allows for recognition of ecostratigraphic events and trends.On this basis, ecostratigraphy serves as a valuable template for thecharacterization, correlation and interpretation of relative sea-levelsand their corresponding sedimentary packages in a time-span fromjust above the Milankovitch-band to the million-year scale of thiscase study.

Acknowledgements

This research was carried out with the financial support of ProjectsP08-RNM-3715, RYC-2009-04316 (Ramón y Cajal Program),CGL2008-03007, and the EMMI Group (RNM-178, Junta de Andalucía,Spain). We would like to thank to Luis Pomar (Universitat de les Illes


Balears), Carlton Brett (University of Cincinnati) and an anonymousreviewer for their careful revision of the manuscript, as well as to P.Vignall for editorial suggestions. Linguistic assistance was providedby J. Sanders.

References

Allenbach, R.P., 2001. Synsedimentary tectonics in an epicontinental sea: a new inter-pretation of the Oxfordian basins of northern Switzerland. Eclogae Geologicae Hel-vetiae 94, 265–287.

Allenbach, R.P., 2002. The ups and downs of “Tectonic Quiescence”—recognizing differ-ential subsidence in the epicontinental sea of the Oxfordian in the Swiss JuraMountains. Sedimentary Geology 150, 323–342.

Aller, R., Hall, P., Rude, P., Aller, J., 1998. Biogeochemical heterogeneity and suboxic dia-genesis in hemipelagic sediments of the Panama Basin. Deep Sea Research Part I45, 133–165.

Anderson, J., 2005. Diachronous development of late Quaternary shelf-margin deltas inthe Northwestern Gulf of Mexico: implications for sequence stratigraphy and deepwater reservoir occurrence. In: Giosan, L., Bhattacharya, J.P. (Eds.), River Deltas:Concepts, Models and Examples: Society for Sedimentary Geology (SEPM) SpecialPublication, 83, pp. 257–276.

Aurell, M., Fernández-López, S., Meléndez, G., 1994. The Middle–Upper Jurassic ooliticironstone level in the Iberian Range (Spain). Eustatic implications. Geobios M.S. 17,549–561.

Aurell, M., Bádenas, B., Bello, J., Delvene, G., Meléndez, G., Pérez-Urresti, I., Ramajo, J.,1999. El Calloviense y el Jurásico Superior en la Cordillera Ibérica Nororiental y lazona de enlace con la Cordillera Costero-Catalana, en los sectores de Sierra deArcos, Calanda y Xerta-Paüls. Cuadernos de Geologia Iberica 25, 73–110.

Aurell, M., Robles, S., Bádenas, B., Rosales, I., Quesada, S., Meléndez, G., García-Ramos,J.C., 2003. Transgressive–regressive cycles and Jurassic palaeogeography of north-east Iberia. Sedimentary Geology 162, 171–239.

Azéma, J., 1977. Étude géologique des zones externes des Cordilleres Bétiques aux con-fins des provinces d' Alicante et de Murcie (Espagne). Ph.D. Thesis Univ. Pierre etMarie Curie, Paris vi, 393 pp.

Bádenas, B., Aurell, M., 2001. Kimmeridgian palaeogeography and basin evolution ofnortheastern Iberia. Palaeogeography, Palaeoclimatology, Palaeoecology 168,291–310.

Barbour-Wood, S.L., Krause Jr., R.A., Kowalewski, M., Wehmiller, J.F., Simoes, M.G.,Goodfriend, G.A., 2003. A comparison of rates of time averaging between the bi-valve Macoma cleryana and brachiopod Bouchardia rosea on a shallow subtropicalshelf. Geological Society of America Abstracts with Programs 35 (6), 273.

Bartolini, A., Pittet, B., Mattioli, E., Hunziker, J.C., 2003. Shallow-platform paleoenviron-mental conditions recorded in deep-shelf sediments: C and O stable isotopes inUpper Jurassic sections of southern Germany (Oxfordian-Kimmeridgian). Sedi-mentary Geology 160, 107–130.

Behmel, H., 1970. Beiträge zur Stratigraphie und Paläontologie des Juras vonOstspanien. V. Stratigraphie und Fazies im präbetischen Jura von Albacete undNord-Murcia. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 137,1–102.

Bernhard, J.M., 1986. Characteristic assemblages and morphologies of benthic forami-nifera from anoxic, organic rich deposits: Jurassic through Holocene. Journal of Fo-raminiferal Research 16, 207–215.

Berry, W.B.N., 1983. On the relationship between ecostratigraphy and zonal stratigra-phy. Newsletters on Stratigraphy 12, 84–97.

Bhattacharya, J.P., 2011. Practical problems in the application of the sequence strati-graphic method and key surfaces: integrating observations from ancient fluvial–deltaic wedges with Quaternary and modelling studies. Sedimentology 58,120–169.

Bonnet, L., Cubaynes, R., Qajoun, A., Rey, J., Ruget, C., 1991. Analyse statistique des bio-cenoses de foraminifers dans les cortèges sedimentaires du Toarcien du Quercy.Comptes Rendus Academie Sciences Paris (II) 313, 1587–1593.

Bonnet, L., Cubaynes, R., Rey, J., Ruget, C., 1992. Cycles et rhythmes dans les biocenosesde foraminifers du Toarcien du Quercy. Comptes Rendus Academie Sciences Paris(II) 314, 77–83.

Boucot, A.J., 1982. Ecostratigraphic framework for the Lower Devonian of the NorthAmerican Appohimchi Subprovince. Neues Jahrbuch für Geologie und Paläontolo-gie Abhandlungen 163, 81–121.

Boucot, A.J., 1986. Ecostratigraphic criteria for evaluating the magnitude, character andduration of bioevents. In: Walliser, O.H. (Ed.), Global Bio-Events, Lecture Notes inEarth Sciences, 8. Springer, Berlin, Heidelberg, pp. 25–45.

Boucot, A.J., 2005. Ecostratigraphy's basis, using Silurian and Devonian examples, withconsideration of the biogeographic complication. In: Koutsoukos, E.A.M. (Ed.), Ap-plied Stratigraphy: Topics in Geobiology, 23, pp. 55–71.

Boucot, A.J., Lawson, J.D. (Eds.), 1999. Paleocommunities — a case study from the Silu-rian and Lower Devonian. Cambridge University Press, Cambridge, New York, Mel-bourne. xv +895 pp.

Bouhamdi, A., Gaillard, C., Ruget, C., 2001. Spirillines versus agglutinants: impact duflux organique et intérêt paléoenvironnemental (Oxfordien moyen du Sud-Est dela France). Geobios 34, 267–277.

Boulila, S., Hinnov, L.A., Huret, E., Collin, P.-Y., Galbrun, B., Fortwengler, D., Marchand,D., Thierry, J., 2008. Astronomical calibration of the Early Oxfordian (Vocontianand Paris basins, France): consequences of revising the Late Jurassic time scale.Earth and Planetary Science Letters 276, 40–51.

Boulila, S., Galbrun, B., Miller, K.G., Pekar, S.F., Browning, J.V., Laskar, L., Wright, J.D.,2011. On the origin of Cenozoic and Mesozoic “third-order” eustatic sequences.Earth-Science Reviews 109, 3–4, 94–112.

Brandano, M., Corda, L., 2002. Nutrients, sea level and tectonics: constraints for the fa-cies architecture of a Miocene carbonate ramp in central Italy. Terra Nova 14,257–262.

Brett, C.E., 1988a. Paleoecology and evolution of marine hard substrate communities:an overview. Palaios 3, 374–378.

Brett, C.E., 1988b. Taphofacies models for epeiric sea environments: Middle Paleozoicexamples. Palaeogeography, Palaeoclimatology, Palaeoecology 63, 225–262.

Brett, C.E., 1995. Sequence stratigraphy, biostratigraphy, and taphonomy in shallowmarine environments. Palaios 10, 497–516.

Brett, C.E., 1998. Sequence stratigraphy, paleoecology, and evolution: biotic clues andresponses to sea-level fluctuations. Palaios 13, 241–262.

Brett, C.E., Baird, G.C., 1986. Comparative taphonomy: a key to paleoenvironment inter-pretation based on fossil preservation. Palaios 1, 207–227.

Brett, C.E., Baird, G.C. (Eds.), 1997. Paleontological events: stratigraphic, ecological, andevolutionary implications. Columbia University Press, New York. 604 pp.

Brett, C.E., Allison, P.A., Moffat, H., Soldani, D., 2006. Sedimentology, taphonomy, andpaleoecology of meter-scale cycles from the Upper Ordovician of Ontario. Palaios21, 530–547.

Brett, C.E., Bartholomew, A.J., Baird, G.C., 2007a. Biofacies recurrence in the middle de-vonian of new york state: an example with implications for evolutionary paleo-ecology. Palaios 22, 306–324.

Brett, C.E., Hendy, A.J.W., Bartholomew, A.J., Bonelli Jr., J.R., McLaughlin, P.I., 2007b. Re-sponse of shallow marine biotas to sea-level fluctuations: a review of faunal re-placement and the process of habitat tracking. Palaios 22, 228–244.

Caracuel, J.E., 1996. Asociaciones de megainvertebrados, evolución ecosedimentaria einterpretaciones ecoestratigráficas en umbrales epioceánicos del Tethys occidental(Jurásico superior). Ph.D. Thesis, Univ. Granada, 458 pp.

Carroll, M., Kowalewski, M., Simões, M.G., Goodfriend, G.A., 2003. Quantitative esti-mates of time-averaging in terebratulid brachiopod shell accumulations from amodern tropical shelf. Paleobiology 29, 381–402.

Catuneanu, O., 2002. Sequence stratigraphy of clastic systems: concepts, merits, andpitfalls. Journal of African Earth Sciences 35, 1–43.

Catuneanu, O., 2006. Principles of Sequence Stratigraphy. Elsevier, Amsterdam. 375 pp.Catuneanu, O., Willis, A.J., Miall, A.D., 1998. Temporal significance of sequence bound-

aries. Sedimentary Geology 121, 157–178.Catuneanu, O., Abreu, V., Bhattacharya, J.P., Blum, M.D., Dalrymple, R.W., Eriksson, P.G.,

Fielding, F.R., Fisher, W.L., Galloway, W.E., Gibling, M.L., Giles, K.A., Holbrook, J.M.,Jordan, R., Kendall, C.G., Macurda, B., Martinsen, O.J., Miall, A.D., Neal, J.E., Numme-dal, D., Pomar, L., Posamentier, H.W., Pratt, B.R., Sarg, J.F., Shanley, K.W., Steel, R.L.,Strasser, A., Tucker, M.E., Winker, C., 2009. Towards the standardization of se-quence stratigraphy. Earth-Science Reviews 92, 1–33.

Catuneanu, O., Bhattacharya, J.P., Blum, M.D., Dalrymple, R.W., Eriksson, P.G., Fielding,C.R., Fisher, W.L., Galloway, W.E., Gianolla, P., Gibling, M.R., Giles, K.A., Holbrook,J.M., Jordan, R., Kendall, C.G.St.C., Macurda, B., Martinsen, O.J., Miall, A.D., Numme-dal1, D., Posamentier, H.W., Pratt, B.R., Shanley, K.W., Steel, R.J., Strasser, A., Tucker,M.E., 2010. Sequence stratigraphy: common ground after three decades of devel-opment. Stratigraphy 28, 21–34.

Christie-Blick, N., Pekar, S.F., Madof, A.S., 2007. Is there a role for sequence stratigraphyin chronostratigraphy? Stratigraphy 4, 217–229.

Cisne, J.L., Chandlee, G.O., 1982. Taconic Foreland Basin graptolites: age zonation, depthzonation, and use in ecostratigraphic correlation. Lethaia 15, 343–363.

Cisne, J.L., Gildner, R.F., Rabe, B.D., 1984. Epeiric sedimentation and sea-level: syntheticecostratigraphy. Lethaia 17, 267–288.

Collin, P.Y., Loreau, J.P., Courville, P., 2005. Depositional environments and iron ooidsformation in condensed sections (Callovian-Oxfordian, south-eastern Paris Basin,France). Sedimentology 52, 969–985.

Corliss, B.H., 1991. Morphology and microhabitat preferences of benthic foraminiferafrom the northwest Atlantic Ocean. Marine Micropaleontology 17, 195–236.

Courville, P., Collin, P.Y., 1997. La série du Callovien et de l'Oxfordien de Veuxhaulles(Châtillonnais, Côte d'Or): problèmes de datation, de géométrie et de paléoenvir-onnements dans une série “condensée”. Bulletin Sciences Bourgogne 49, 29–43.

Courville, P., Collin, P.Y., 2002. Taphonomic sequences: a new tool for sequence stratig-raphy. Geology 30, 511–514.

Cubaynes, R., Ruget, C., Rey, J., 1989. Essai de caractérisation des prismes de dépôt d'or-igine eustatique par les associations de foraminifères benthiques: exemple du Liasmoyen et supérieur sur la bordure est du Bassin Aquitain. Comptes Rendus Acade-mie Sciences Paris (II) 308, 1517–1522.

Cubaynes, R., Rey, J., Ruget, C., 1990. Renouvellement des spèces de foraminifersbenthiques et variations globales du niveau des mers. Exemples du Lias du Quercyet de l'Éocene des Corbières. Revue de Micropaleontologie 33, 233–240.

Davaud, E., Lombard, A., 1973. Statistical approach to the problem of alternating beds oflimestone and marl (Upper Oxfordian of the French Jura). Eclogae Geologicae Hel-vetiae 68, 491–509.

De la Mora, A., Olóriz, F., González-Arreola, C., 2000. ‘Autochthonous’ bivalve assem-blages and palaeoecologic interpretation in the Upper Jurassic–Lowermost Creta-ceous La Caja Formation from the Cañón de San Matías (Zacatecas, Mexico).Comptes Rendus Academie Sciences Paris (II) 331, 741–747.

Donovan, D.T., 1985. Ammonite shelf form and transgression in the British Lower Ju-rassic. In: Bayer, U., Seilacher, A. (Eds.), Sedimentary and Evolutionary Cycles. Lec-ture Notes on Earth Sciences. Springer, Berlin, pp. 48–58.

Dromart, G., 1989. Deposition of Upper Jurassic fine-grained limestones in the westernSubalpine Basin, France. Palaeogeography, Palaeoclimatology, Palaeoecology 69,23–43.


Einsele, G., Ricken, W., 1991. Limestone-marl alternation: an overview. In: Einsele, G.,Ricken, W., Seilacher, A. (Eds.), Cycles and Events in Stratigraphy. Springer-Verlag,Berlin, pp. 79–93.

El Hariri, K., Sadki, D., Dommergues, J.L., Mouterde, R., 1992. Niveaux condensés, fluc-tuations eustatiques? Implications sur les faunes d'ammonites jurassiques dansle Haut-Atlas (Maroc). Comptes Rendus Academie Sciences Paris (II) 314, 507–510.

Embry, A.F., 1993. Transgressive-regressive (T-R) sequence analysis of the Jurassic suc-cession of the Sverdrup Basin, Canadian Arctic Archipelago. Canadian Journal ofEarth Sciences 30, 301–320.

Embry, A.F., 1995. Sequence boundaries and sequence hierarchies: problems and pro-posals. In: Steel, R.J., Felt, V.L., Johannessen, E.P., Mathieu, C. (Eds.), Sequence Stra-tigraphy on the Northwest European Margin: Norwegian Petroleum SocietySpecial Publication, 5, pp. 1–11.

Embry, A.F., 2002. Transgressive–regressive (T–R) sequence stratigraphy. In: Armentr-out, J.M. (Ed.), Sequence Stratigraphic Models for Exploration and Production:Evolving Methodology, Emerging Models and Application Histories. 22nd AnnualGulf Coast Section Society for Sedimentary Geology (SEPM) Foundation Bob F. Per-kins Research Conference, pp. 151–172.

Embry, A.F., Johannessen, E.P., 1992. T–R sequence stratigraphy, facies analysis and res-ervoir distribution in the uppermost Triassic–Lower Jurassic succession, WesternSverdrup Basin, Arctic Canada. In: Vorren, T.O., Bergsager, E., Dahl-Stamnes, O.A.,Holter, E., Johansen, B., Lie, E., Lund, T.B. (Eds.), Arctic Geology and Petroleum Po-tential: Norwegian Petroleum Society Special Publication, 2, pp. 121–146.

Embry, A., Johannessen, E.P., Owen, D., Beauchamp, B., Gianolla, P., 2007. Sequencestratigraphy as a “concrete” stratigraphic discipline. Report of the ISSC TaskGroup on Sequence Stratigraphy. 104 pp.

Fernández-López, S., 1997. Ammonites, ciclos tafonómicos y ciclos estratigráficos enplataformas epicontinentales carbonáticas. Revista Española de Paleontología 12,151–174.

Fernández-López, S., 2000. Ammonite taphocycles in carbonate epicontinental plat-forms. GeoResearch Forum 6, 293–300.

Fernández-López, S., Meléndez, G., 1994. Abrasion surfaces on internal moulds of am-monites as palaeobathymetric indicators. Palaeogeography, Palaeoclimatology,Palaeoecology 110, 29–42.

Fisher, R.A., Corbet, A.S., Williams, C.B., 1943. The relations between the number of spe-cies and the number of individuals in a random sample of an animal population.Journal of Animal Ecology 12, 42–58.

Flessa, K.W., Kowalewsky, M., 1994. Shell survival and time-averaging in nearshore andshelf environments: estimates from the radiocarbon literature. Lethaia 27, 153–165.

Foucault, A., 1971. Etude géologique des environs des sources du Guadalquivir (provincesde Jaen et de Grenade, Espagne méridionale). Ph.D. Thesis, Univ. Paris, 633 pp.

Fugagnoli, A., 2004. Trophic regimes of benthic foraminiferal assemblages in Lower Ju-rassic shallow water carbonates from northeastern Italy (Calcari Grigi, Trento Plat-form, Venetian Prealps). Palaeogeography, Palaeoclimatology, Palaeoecology 205,111–130.

Fürsich, F.T., Oschmann, W., 1993. Shell beds as tool in basin analysis: the Jurassic ofKachchh, western India. Journal of the Geological Society of London 150, 169–185.

Fürsich, F.T., Oschmann, W., Jaitly, A.K., Singh, I.B., 1991. Faunal response to transgres-sive–regressive cycles: example from the Jurassic of western India. Palaeogeogra-phy, Palaeoclimatology, Palaeoecology 85, 149–159.

Galloway, W.E., 1989. Genetic stratigraphic sequences in basin analysis 1. Architectureand genesis of flooding-surface bounded depositional units. American Associationof Petroleum Geologists (AAPG). Bulletin 73, 125–142.

Galloway, W.E., 1998. Depositional processes, regime variables, and development ofsiliciclastic stratigraphic sequences. In: Gradstein, F.M., Sandvik, K.O., Milton, N.J.(Eds.), Sequence Stratigraphy — Concepts and Applications: Norwegian PetroleumSociety Special Publication, 8, pp. 117–140.

Galloway, W.E., 2002. Paleogeographic setting and depositional architecture of a sand-dominated shelf depositional system, Miocene Utsira Formation, North Sea Basin.Journal of Sedimentary Research 72, 476–490.

García-Hernández, M., 1978. El Jurásico terminal y el Cretácico inferior en las Sierras deCazorla y del Segura (Zona Prebética). Ph.D. Thesis, Univ. Granada, 344 pp.

García-Hernández, M., López-Garrido, A.C., 2004. El Prebético del sector central y aflor-amientos más occidentales, Ciclo III. In: Vera, J.A. (Ed.), Geología de España. Socie-dad Geológica de España — Instituto Geológico y Minero de España, pp. 366–368.

García-Hernández, M., López-Garrido, A.C., Olóriz, F., 1981. Étude des calcaires nodu-leux du Jurassique superieur de la Zone Prébétique (Cordillères Bétiques, SE del'Espagne). In: Farinacci, A., Elmi, S. (Eds.), Proceedings Rosso Ammonitico Sympo-sium. Tecnoscienza, Rome, pp. 419–434.

García-Hernández, M., López-Garrido, A.C., Martín-Algarra, A., Molina, J.M., Ruiz-Ortiz,P.A., Vera, J.A., 1989. Las discontinuidades mayores del Jurásico de las Zonas Exter-nas de las Cordilleras Béticas: análisis e interpretación de los ciclos sedimentarios.Cuadernos de Geología Ibérica 13, 35–52.

Gobeil, C., MacDonald, R., Sundby, B., 1997. Diagenetic separation of cadmium andmanganese in suboxic continental margin sediments. Geochimica et Cosmochi-mica Acta 61, 4647–4654.

Goldhammer, R.K., 1998. Second-order accommodation cycles and points of “strati-graphic turnaround”: implications for high-resolution sequence stratigraphy andfacies architecture of the Haynesville and Cotton Valley Lime “Pinnacle Reefs” ofthe East Texas Basin. West Texas Geological Bulletin 37, 5–9.

Gooday, A., 1996. Epifaunal and shallow infaunal foraminiferal communities at threeabyssal NE Atlantic sites subject to differing phytodetritus input regimes. Deep-Sea Research I 9, 1395–1421.

Goy, A., Gómez, J.J., Bacelar, J., 1973. El Bathoniense, Calloviense y Oxfordiense, en fa-cies de oolitos ferruginosos, en el límite SW del Rincón de Ademuz (Valencia-Cuenca). Estudios Geológicos 29, 355–366.

Gradstein, F.M., Ogg, J., Smith, A.G., Agterberg, F.P., Bleeker, W., Cooper, R.A., Davydov,V., Gibbard, P., Hinnov, L.A., House, M.R., Lourens, L., Luterbacher, H.P., McArthur, J.,Melchin, M.J., Robb, L.J., Shergold, J., Villeneuve, M., Wardlaw, B.R., Ali, J., Brinkhuis,H., Hilgen, F.J., Hooker, J., Howarth, R.J., Knoll, A.H., Laskar, J., Monechi, S., Plumb,K.A., Powell, J., Raffi, I., Röhl, U., Sadler, P., Sanfilippo, A., Schmitz, B., Shackleton,N.J., Shields, G.A., Strauss, H., Van Dam, J., van Kolfschoten, T., Veizer, J., Wilson,D., 2004. A geologic time scale 2004. Cambridge University Press, Cambridge.589 pp.

Gygi, R.A., 1981. Oolitic iron formations: marine or not marine? Eclogae GeologicaeHelvetiae 74, 233–254.

Gygi, R.A., 1986. Eustatic sea level changes of the Oxfordian (Late Jurassic) and their ef-fect documented in sediments and fossil assemblages of an epicontinental sea.Eclogae Geologicae Helvetiae 79, 455–491.

Gygi, R.A., Persoz, F., 1987. The epicontinental sea of Swabia (southern Germany) in theLate Jurassic: factors controlling sedimentation. Neues Jahrbuch für Geologie undPaläontologie Abhandlungen 176, 49–65.

Haig, D.W., Lynch, D.A., 1993. A late early Albian marine transgressive pulse overnortheastern Australia, precursor to epeiric basin anoxia: foraminiferal evidence.Marine Micropaleontology 22, 311–362.

Hallam, A., 1986. The origin of minor limestone-shale cycles: climatically induced ordiagenetic? Geology 14, 609–612.

Hantzpergue, P., 1991. Biogéographie des ammonites et variations du niveau marin:apport de la stratigraphie séquentielle dans l'analyse des peuplements du Kimmér-idgien Nord-Aquitaine. Geobios 24, 59–64.

Haq, B.U., Hardenbol, J., Vail, P.R., 1987. Chronology of fluctuating sea level since theTriassic. Science 235, 1156–1167.

Haq, B.U., Hardenbol, J., Vail, P.R., 1988. Mesozoic and Cenozoic chronostratigraphyand cycles of sea-level change. In: Wilgus, C.K., Hastings, B.S., Kendall, C.G., Posa-mentier, H.W., Ross, C.A., van Wagoner, J.C. (Eds.), Sea-Level Changes — An inte-grated approach: Society for Sedimentary Geology (SEPM) Special Publication,42, pp. 71–108.

Hardenbol, J., Thierry, J., Farley, M.B., Jacquin, T., De Gracianski, P.C., Vail, P.R., 1998. Me-sozoic and Cenozoic sequence chronostratigraphic framework of European basins.In: De Gracianski, P.C., Hardenbol, J., Jacquin, T., Vail, P.R. (Eds.), Mesozoic and Ce-nozoic Sequence Stratigraphy of European Basins: Society for Sedimentary Geolo-gy (SEPM) Special Publication, 60. charts 1–8, Tulsa, OK.

Helland-Hansen, W., Gjelberg, J.G., 1994. Conceptual basis and variability in sequencestratigraphy: a different perspective. Sedimentary Geology 92, 31–52.

Henderson, A.S., Hart, M.B., 2000. The distribution of Foraminiferida in the Oxfordiansequences of North Dorset, UK. GeoResearch Forum 6, 311–320.

Holland, S.M., Patzkowsky, M.E., 2004. Ecosystem structure and stability: middle UpperOrdovician of central Kentucky, USA. Palaios 19, 316–331.

Holland, S.M., Patzkowsky, M.E., 2009. Stratigraphic architecture of a tropical carbonateplatform and its effect on the distribution of fossils: Ordovician Bighorn Dolomite,Wyoming, USA. Palaios 24, 303–317.

Holland, S.M., Meyer, D.L., Miller, A.I., 2000. High-resolution correlation in apparentlymonotonous rocks: Upper Ordovician Kope Formation, Cincinnati Arch. Palaios15, 73–80.

Huber, B., Müller, B., Luterbacher, H., 1987. Mikropaläontologische Untersuchungen ander Callovien/Oxfordien-Grenze im Schweizer Jura und auf der Schwäbischen Alb(vorläufige Mitteilung). Eclogae Geologicae Helvetiae 80, 449–459.

Hunt, D., Tucker, M.E., 1992. Stranded parasequences and the forced regressivewedge systems tract: deposition during base-level fall. Sedimentary Geology81, 1–9.

Hunt, D., Tucker, M.E., 1993. Sequence stratigraphy of carbonate shelves with an exam-ple from the mid-Cretaceous (Urgonian) of Southeast France. In: Posamentier,H.W., Summerhayes, C.P., Haq, B.U., Allen, G.P. (Eds.), Sequence Stratigraphy andFacies Associations: International Association of Sedimentologists (IAS) SpecialPublication, 18, pp. 307–341.

Hylton, M.D., Hart, M.B., 2000. Benthic foraminiferal response to Pliensbachian-Toarcian (Lower Jurassic) sea level change and oceanic anoxia in NW Europe.GeoResearch Forum 6, 455–462.

Insalaco, E., 1999. Facies and Palaeoecology of Upper Jurassic (Middle Oxfordian) coralreefs in England. Facies 40, 81–100.

Jerez-Mir, L., 1973. Geología de la Zona Prebética, en la transversal de Elche de la Sierray sectores adyacentes (provincias de Albacete y Murcia). Ph.D. Thesis, Univ. Grana-da, 749 pp.

Jervey, M.T., 1988. Quantitative geological modeling of siliciclastic rock sequencesand their seismic expression. In: Wilgus, C.K., Hastings, B.S., Kendall, C.G.St.C.,Posamentier, H.W., Ross, C.A., VanWagoner, J.C. (Eds.), Sea-Level Changes: An In-tegrated Approach: Society for Sedimentary Geology (SEPM) Special Publication,42, pp. 47–69.

Jørgensen, B.B., Glud, R., Holby, O., 2005. Oxygen distribution and bioirrigation in Arcticfjord sediments (Svalbard, Barents Sea). Marine Ecology Progress Series 292, 85–95.

Jorissen, F., 1999. Benthic foraminiferal microhabitats below the sediment–water inter-face. In: Sen Gupta, B. (Ed.), Modern Foraminifera. Kluwer Academic Publishers,Dordrecht, pp. 161–180.

Jorissen, F.J., De Stigter, H.C., Widmark, J.G.V., 1995. A conceptual model explainingbenthic foraminiferal habitats. Marine Micropaleontology 26, 3–15.

Kauffman, E.G., 1986. High resolution event stratigraphy: regional and global Creta-ceous bio-events. In: Walliser, O.H. (Ed.), Global Bio-Events. Lecture Notes inEarth Sciences, 8. Springer, Berlin, pp. 279–335.

Kauffman, E.G., 1988. Concepts and methods of high-resolution event stratigraphy. An-nual Review Earth Planetary Science Letters 16, 605–654.

Kendall, C.G., Levine, P.A., Ehrich, R., 1995. The enigma of third-order sea level cycles: acosmic connection? In: Haq, B.Q. (Ed.), Sequence Stratigraphy and Depositional


Response to Eustatic, Tectonic and Climatic Forcing. Kluwer Academic, Dordrecht,pp. 367–376.

Kerans, C., Loucks, R.G., 2002. Stratigraphic setting and controls on occurrence of highenergy carbonate beach deposits: Lower Cretaceous of the Gulf of Mexico. Transac-tions, Gulf Coast Association of Geological Societies 52, 517–526.

Kidwell, S.M., 2001. Preservation of species abundance in marine death assemblages.Science 294, 1091–1094.

Kidwell, S.M., 2002. Mesh-size effects on the ecological fidelity of death assemblages: ameta-analysis of molluscan live–dead studies. Geobios 35 (Supplement 1),107–119.

Kidwell, S.M., Bosence, D.W.J., 1991. Taphonomy and time-averaging of marine shellyfaunas. In: Allison, P.A., Briggs, D.E.G. (Eds.), Taphonomy. : Releasing the DataLocked in the Fossil Record. Plenum, New York, pp. 115–209.

Kuhnt, W., Moullade, M., Kaminski, M.A., 1996. Ecological structuring and evolution ofdeep sea agglutinated foraminifera — a review. Revue de Micropaleontologie 39,271–281.

Leinfelder, R.R., Wilson, R.C., 1998. Third-order sequences in an upper Jurassic rift-related second-order sequence, Central Lusitanian Basin, Portugal. In: De Gracianski,P.C., Hardenbol, J., Jacquin, T., Vail, P.R. (Eds.), Mesozoic and Cenozoic Sequence Stra-tigraphy of European Basins: Society for Sedimentary Geology (SEPM) Special Publi-cation, 60, pp. 507–525.

Leinfelder, R.R., Nose, M., Schmid, D., Werner, W., 1993. Microbial crusts of the late Ju-rassic: composition, palaeoecological significance and importance in reef construc-tion. Facies 29, 195–230.

Leinfelder, R.R., Krautter, M., Laternser, R., Nose, M., Schmid, D.U., Schweigert, G., Werner,W., Keupp, H., Brugger, H., Herrmann, R., Rehfeld-Kiefer, U., Schroeder, J.H., Reinhold,C., Koch, R., Zeiss, A., Schweizer, V., Christmann, H., Menges, G., Luterbacher, H., 1994.The origin of Jurassic reefs: current research developments and results. Facies 31,1–56.

Lorin, S., Courville, P., Collin, P.Y., Thierry, J., Tort, A., 2004. Modalités de réinstallationd'une plate-forme carbonatée après une crise sédimentaire: exemple de la limiteOxfordien moyen – Oxfordien supérieur dans le Sud-Est du Bassin de Paris. Bulletinde la Société Géologique de France 175, 289–302.

Magniez-Jannin, F., 1992. Foraminifers tolerance to dysaerobic conditions within EarlyCretaceous depositional sequences of the Vocontian Basin (SE France). SequenceStratigraphy of European Basins, CNRS-IFP, Abstract Volume, pp. 284–285.

Marques, B., Olóriz, F., Rodríguez-Tovar, F.J., 1991. Interactions between tectonics andeustasy during the Upper Jurassic and Lowermost Cretaceous. Examples from thesouth of Iberia. Bulletin de la Société Géologique de France 162, 1109–1124.

Marques, B., Olóriz, F., Caetano, P.S., Rodríguez-Tovar, F.J., 1993. Relative sea-level fluc-tuations and ecostratigraphy; applications to Middle/Upper Oxfordian fossil as-semblages of east-central Algarve, Portugal. Acta Geologica Polonica 43, 289–298.

Marques, B., Olóriz, F., Rodríguez-Tovar, F.J., 1996. Preliminary ecostratigraphic re-searches in the Middle-Upper Oxfordian from the Lusitanian basin (west Portugal).Comparison with the Eastern Algarve basin (south Portugal). Comptes RendusAcademie Sciences Paris (II) 322, 757–764.

Martin, R.E., 1991. Ecostratigraphy datums and sequence stratigraphy: application tothe marine quaternary. American Association of Petroleum Geologists (AAPG) Bul-letin 75, 630.

Martinsson, A., 1973. Ecostratigraphy. Lethaia 6, 441–443.McRoberts, C.A., Aberhan, M., 1997. Marine diversity and sea-level changes: numerical

test for associations using Early Jurassic bivalves. Geologische Rundschau 86,160–167.

Meléndez, G., Ramajo, J., Martínez-Cotanda, S., 2005. El desarrollo de la Capa de Arroyo-frío (límite Calloviense-Oxfordiense) al Sur de Zaragoza, entre Ricla y Aguilón:bioestratigrafía y facies. Geogaceta 38, 3–6.

Miall, A.D., 1997. The geology of stratigraphic sequences. Springer-Verlag, BerlinHeidelberg New York. 433 pp.

Miall, A.D., 2004. Empiricism and model building in stratigraphy: the historical roots ofpresent-day practices. Stratigraphy 1, 3–25.

Miall, A.D., Miall, C.E., 2001. Sequence stratigraphy as a scientific enterprise: theevolution and persistence of conflicting paradigms. Earth-Science Reviews 54,321–348.

Mitchum, R.M., Vail, P.R., 1977. Seismic stratigraphy and global changes of sea level,Part. 7: seismic stratigraphy interpretation. In: Payton, C.E. (Ed.), Seismic Stratigra-phy — Applications to Hydrocarbon Exploration: American Association of Petro-leum Geologists (AAPG) Memoir, 26, pp. 135–163. Tulsa, OK.

Mitchum, R.M., Van Wagoner, J.C., 1991. High-frequency sequences and their stackingpatterns: sequence stratigraphic evidence of high-frequency eustatic cycles. Sedi-mentary Geology 70, 131–160.

Mitchum, R.M., Vail, P.R., Thompson, S., 1977. Seismic stratigraphy and global changesof sea level, Part 2: the depositional sequence as a basic unit for stratigraphy anal-ysis. In: Payton, C.E. (Ed.), Seismic Stratigraphy — Applications to HydrocarbonExploration: American Association of Petroleum Geologists (AAPG) Memoir, 26,pp. 202–212.

Muto, T., Steel, R.J., 2000. The accommodation concept in sequence stratigraphy:some dimensional problems and possible redefinition. Sedimentary Geology 130,1–10.

Nagy, J., 1992. Environmental significance of foraminiferal morphogroups in JurassicNorth Sea deltas. Palaeogeography, Palaeoclimatology, Palaeoecology 95, 111–134.

Nagy, J., Gradstein, F.M., Kaminski, M.A., Holbourn, A.E., 1995. Foraminiferal mor-phogroups, paleoenvironments and new taxa from Jurassic to Cretaceous strataof Thakkhola, Nepal. Grzybowski Foundation Special Publication 3, 181–209.

Nagy, J., Finstand, E.K., Dypvik, H., Bremer, M.G.H., 2001. Response of foraminiferal fa-cies to transgressive-regressive cycles in the Callovian of Northeast Scotland. Jour-nal of Foraminiferal Research 31, 324–349.

Nagy, J., Reolid, M., Rodríguez-Tovar, F.J., 2009. Foraminiferal morphogroups in dysoxicshelf deposits from the Jurassic of Spitsbergen. Polar Research 28, 214–221.

Neagu, T., Dragastan, O., Csiki, Z., 1997. Early Cretaceous shelf paleocommunities ofCernavoda (south Dobrogea, SE Romania). Acta Paleontologica Romaniae 1, 28–36.

Nummedal, D., 2004. Tectonic and eustatic controls on Upper Cretaceous stratigraphyof Northern New Mexico. In: Mack, G.H., Giles, K.A. (Eds.), The Geology of NewMexico— A Geologic History, vol. 11. NewMexico Geological Society, pp. 169–182.

Nummedal, D., Swift, D.J.P., 1987. Transgressive stratigraphy at sequence-bounding un-conformities: some principles derived from Holocene and Cretaceous examples. In:Nummedal, D., Pilkey, O.H., Howard, J.D. (Eds.), Sea-level Fluctuation and Coastal Evo-lution: Society for Sedimentary Geology (SEPM) Special Publication, 41, pp. 241–260.

Ogg, J., Przybylski, P., 2006. Jurassic chronostratigraphic charts and a display interface.Abstract for 7th International Congress on the Jurassic System, Krakow, Poland[poster session]. Volumina Jurassica 4, 199.

Olivier, N., Pittet, B., Mattioli, E., 2004. Palaeoenvironmental control on sponge-microbialite reefs and contemporaneous deep-shelf marl-limestone deposition(Late Oxfordian, southern Germany). Palaeogeography, Palaeoclimatology,Palaeoecology 212, 233–263.

Olóriz, F., 1985. Paleogeography and ammonites in the Upper Jurassic. Outlines for apattern. In: Pallini, G. (Ed.), Fossili, Evoluzione, Ambiente. Atti I Convengo Interna-zionale, Pergola, Belli, Pesaro, pp. 1–9.

Olóriz, F., 1990. Ammonite phenotypes and ammonite distributions. Notes and com-ments. In: Pallini, G., Cecca, F., Cresta, S., Santantonio, M. (Eds.), Fossili, Evoluzione,Ambiente. Atti II Convengo Internazionale, Pergola, Tecnostampa, pp. 417–426.

Olóriz, F., 2000. Time-averaging and long-term palaeoecology in macroinvertebratefossil assemblages with ammonites (Upper Jurassic). Revue de Paléobiologie Vol.Spéc. 8, 123–140.

Olóriz, F., Rodríguez-Tovar, F.J., 1999. Biostratigraphic review and ecostratigraphicanalysis of the Fuente Alamo profile (External Prebetic, Betic Cordillera, Spain).Profil 16, 73–81.

Olóriz, F., Marques, B., Rodríguez-Tovar, F.J., 1991. Eustatism and faunal associations.Examples from the South Iberian Margin during the Late Jurassic (Oxfordian-Kim-meridgian). Eclogae Geologicae Helvetiae 84, 83–106.

Olóriz, F., Rodríguez-Tovar, F.J., Marques, B., Caracuel, J.E., 1993. Ecostratigraphy andsequence stratigraphy in high frequency sea level fluctuations: examples from Ju-rassic macroinvertebrate assemblages. Palaeogeography, Palaeoclimatology,Palaeoecology 101, 131–145.

Olóriz, F., Rodríguez-Tovar, F.J., Marques, B., 1994. Macroinvertebrates assemblagesand ecostratigraphic structuration within a highstand system tract. An examplefrom the Lower Kimmeridgian in southern Iberia. Geobios M.S. 17, 605–614.

Olóriz, F., Caracuel, J., Rodríguez-Tovar, F.J., 1995. Using ecostratigraphic trends in se-quence stratigraphy. In: Haq, B.U. (Ed.), Sequence Stratigraphy and DepositionalResponse to Eustatic, Tectonic and Climatic Forcing. Kluwer Academic Publisher,Dordrecht, pp. 59–85.

Olóriz, F., Caracuel, J.E., Ruiz-Heras, J.J., Rodríguez-Tovar, F.J., Marques, B., 1996. Ecos-tratigraphic approaches, sequence stratigraphy proposals and block tectonics: ex-amples from epioceanic swell areas in south and east Iberia. Palaeogeography,Palaeoclimatology, Palaeoecology 121, 273–295.

Olóriz, F., Reolid, M., Rodríguez-Tovar, F.J., 1999. Fine-resolution ammonite biostratig-raphy at the Rio Gazas-Chorro II section in Sierra de Cazorla (Prebetic Zone, Jaénprovince, southern Spain). Profil 16, 83–94.

Olóriz, F., Caracuel, J.E., Rodríguez-Tovar, F.J., Tavera, J.M., 2002a. Upper Jurassic: BeticCordillera. In: Gibbons, W., Moreno, M.T. (Eds.), The Geology of Spain. GeologicalSociety, London, pp. 247–251.

Olóriz, F., Reolid, M., Rodríguez-Tovar, F.J., 2002b. Fossil assemblages, lithofacies andtaphofacies for interpreting depositional dynamics in epicontinental Oxfordian(Prebetic Zone, Betic Cordillera, southern Spain). Palaeogeography, Palaeoclimatol-ogy, Palaeoecology 185, 53–75.

Olóriz, F., Reolid, M., Rodríguez-Tovar, F.J., 2003a. Palaeogeographic and stratigraphicdistribution of Mid-Late Oxfordian foraminiferal assemblages in the PrebeticZone (Betic Cordillera, southern Spain). Geobios 36, 733–747.

Olóriz, F., Reolid, M., Rodríguez-Tovar, F.J., 2003b. A Late Jurassic carbonate ramp colo-nized by sponges and benthic microbial communities (External Prebetic, SouthernSpain). Palaios 18, 528–545.

Olóriz, F., Villaseñor, A.B., González-Arreola, C., 2003c. Major lithostratigraphic units inland-outcrops of north-central Mexico and the subsurface along the northern rimof Gulf of Mexico Basin (Upper Jurassic–lowermost Cretaceous): a proposal for cor-relation of tectono-eustatic sequences. Journal of South American Earth Sciences16, 119–142.

Olóriz, F., Reolid, M., Rodríguez-Tovar, F.J., 2004a. Taphonomy of ammonite assem-blages from the Middle-Upper Oxfordian (Transversarium?-Bifurcatus Zones) inthe Internal Prebetic (Betic Cordillera, Southern Spain): taphonic populationsand taphofacies for ecostratigraphic interpretations. Rivista Italiana PaleontologiaStratigrafia 110, 239–248.

Olóriz, F., Reolid, M., Rodríguez-Tovar, F.J., 2004b. Microboring and taphonomy in MiddleOxfordian to lowermost Kimmeridgian (Upper Jurassic) from the Prebetic Zone(southern Iberia). Palaeogeography, Palaeoclimatology, Palaeoecology 212, 181–197.

Olóriz, F., Reolid, M., Rodríguez-Tovar, F.J., 2006. Approaching trophic structure in Late Ju-rassic neritic shelves: a western Tethys example from southern Iberia. Earth-ScienceReviews 79, 101–139.

Olóriz, F., Reolid, M., Rodríguez-Tovar, F.J., 2008. Taphonomy of fossil macro-invertebrate assemblages as a tool for ecostratigraphic interpretation in Upper Ju-rassic shelf deposits (Prebetic Zone, southern Spain). Geobios 41, 31–42.

Olszewski, T.D., 1999. Taking advantage of time-averaging. Paleobiology 25, 226–238.Olszewski, T.D., 2004. Modeling the influence of taphonomic destruction, reworking,

and burial on time-averaging in fossil accumulations. Palaios 19 (1), 39–50.


Oxford, M.J., Hart, M.B., Watkinson, M.P., 2000. Micropaleontological investigations ofthe Oxford clay-corallian succession of the Dorset Coast. Geosciences SW England10, 9–13.

Oxford, M.J., Gregory, F.J., Hart, M.B., Henderson, A.S., Simmons, M.D., Watkinson, M.P.,2002. Jurassic planktonic foraminifera from the United Kingdom. Terra Nova 14,205–209.

Pittet, B., Gorin, G.E., 1997. Distribution of sedimentary organic matter in a mixed car-bonate–siliciclastic platform environment: Oxfordian of the Swiss Jura Mountains.Sedimentology 44, 915–937.

Pittet, B., Mattioli, E., 2002. The carbonate signal and calcareous nannofossil distribu-tion in an Upper Jurassic section (Balingen-Tieringen, Late Oxfordian, southernGermany). Palaeogeography, Palaeoclimatology, Palaeoecology 179, 71–96.

Pittet, B., Strasser, A., 1998a. Depositional sequences in deep-shelf environmentsformed through carbonate mud import from the shallow platform (Late Oxfordian,German Swabian Alb and eastern Swiss Jura). Eclogae Geologicae Helvetiae 91,149–169.

Pittet, B., Strasser, A., 1998b. Long-distance correlations by sequence stratigraphy andcyclostratigraphy: examples and implications (Oxfordian from the Swiss Jura,Spain, and Normandy). Geologische Rundschau 86, 852–874.

Pittet, B., Strasser, A., Mattioli, E., 2000. Depositional sequences in deep-shelfenvironments: a response to sea-level changes and shallow-platform carbonateproductivity (Oxfordian, Germany and Spain). Journal of Sedimentary Research70, 392–407.

Plint, A.G., Nummedal, D., 2000. The falling stage systems tract: recognition and impor-tance in sequence stratigraphic analysis. In: Hunt, D., Gawthorpe, R.L. (Eds.), Sedi-mentary Response to Forced Regression: Geological Society of London SpecialPublication, 172, pp. 1–17.

Pomar, L., 1993. High-resolution sequence stratigraphy in prograding carbonates: ap-plication to seismic interpretation. In: Loucks, B., Sarg, J.F. (Eds.), Carbonate Se-quence Stratigraphy: Recent Developments and Applications: AmericanAssociation of Petroleum Geologists (AAPG) Memoir, 57, pp. 389–407.

Pomar, L., 2001a. Ecological control of sedimentary accommodation: evolution of a car-bonate ramp to rimmed shelf, Upper Miocene, Balearic Islands. Palaeogeography,Palaeoclimatology, Palaeoecology 175, 249–272.

Pomar, L., 2001b. Types of carbonate platforms: a genetic approach. Basin Research 13,313–334.

Pomar, L., Kendall, C.G., 2008. Architecture of carbonate platforms: a response to hy-drodynamics and evolving ecology. In: Lukasik, J., Simo, A. (Eds.), Controls on Car-bonate Platform and Reef Development: Society for Sedimentary Geology (SEPM)Special Publication, 89, pp. 187–216.

Pomar, L., Ward, W.C., 1995. Sea-level changes, carbonate production and platform ar-chitecture: the Llucmajor Platform, Mallorca, Spain. In: Haq, B.U. (Ed.), SequenceStratigraphy and Depositional Response to Eustatic. : Tectonic and Climatic Forc-ing. Kluwer Academic Press, Dordrecht, pp. 87–112.

Pomar, L., Gili, E., Obrador, A., Ward, W.C., 2005. Facies architecture and high-resolution sequence stratigraphy of an Upper Cretaceous platform margin succes-sion, southern central Pyrenees, Spain. Sedimentary Geology 175, 339–365.

Posamentier, H.W., Allen, G.P., 1999. Siliciclastic sequence stratigraphy: concepts andapplications. Concepts in Sedimentology and Paleontology, 7. Society for Sedimen-tary Geology (SEPM). 210 pp.

Posamentier, H.W., James, D.P., 1993. An overview of sequence-stratigraphic concepts:uses and abuses. In: Posamentier, H.W., Summerhayes, C.P., Haq, B.U., Allen, G.P.(Eds.), Sequence Stratigraphy and Facies Associations: International Associationof Sedimentologists (IAS) Special Publication, 18, pp. 3–18.

Posamentier, H.W., Kolla, V., 2003. Seismic geomorphology and stratigraphy of deposi-tional elements in deep-water settings. Journal of Sedimentary Research 73,367–388.

Posamentier, H.W., Morris, W.R., 2000. Aspects of the stratal architecture of forced regres-sive deposits. In: Hunt, D., Gawthorpe, R.L. (Eds.), Sedimentary Responses to ForcedRegressions: Geological Society of London Special Publication, 172, pp. 19–46.

Posamentier, H.W., Vail, P.R., 1988. Eustatic controls on clastic deposition 2. Sequenceand system tract models. Sea Level Changes: An Integrated Approach: Society forSedimentary Geology (SEPM) Special Publication, 42, pp. 125–154.

Posamentier, H.W., Walker, R.G., 2006. Deep-water turbidites and submarine fans. In:Posamentier, H.W., Walker, R.G. (Eds.), Facies Models Revisited. Special Publica-tion, 84. Society for Sedimentary Geology (SEPM), pp. 397–520.

Posamentier, H.W., Jervey, M.T., Vail, P.R., 1988. Eustatic controls on clastic deposition1. Conceptual framework. Sea Level Changes: An Integrated Approach: Society forSedimentary Geology (SEPM) Special Publication, 42, pp. 109–124.

Rabe, B.D., Cisne, J.L., 1980. Chronostratigraphic accuracy of Ordovician ecostrati-graphic correlation. Lethaia 13, 109–118.

Raffi, S., Serpagli, E., 1993. Introduzione alla paleontologia. UTET, Torino. 654 pp.Ramajo, J., Aurell, M., 1997. Análisis sedimentológico de las discontinuidades y depós-

itos del Calloviense superior-Oxfordiense medio en la Cordillera Ibérica Norocci-dental. Cuadernos de Geología Ibérica 22, 213–236.

Ramajo, J., Aurell, M., 2008. Long-term Callovian–Oxfordian sea-level changes and sed-imentation in Iberian carbonate platform (Jurassic, Spain): possible eustatic impli-cations. Basin Research 20, 163–184.

Ramajo, J., Aurell, M., Cepría, J., 2002. Análisis de facies de la Capa de oolitos ferrugino-sos de Arroyofrío en la Sierra de Arcos (Jurásico, Cordillera Ibérica septentrional).Journal of Iberian Geology 28, 45–64.

Reitner, J., Neuweiler, F., 1995. Mud mounds: a polygenetic spectrum of fine-grainedcarbonate buildups. Facies 32, 1–70.

Reolid, M., 2003. Dinámica eco-sedimentaria durante el Oxfordiense medio–Kimmer-idgiense temprano en la Zona Prebética: Interpretación ecoestratigráfica y secuen-cial. Ph.D. Thesis, Univ. Granada, 254 pp.

Reolid, M., 2005. Asociaciones de braquiópodos del Oxfordiense medio-Kimmeridgienseinferior en la Zona Prebética (Sureste de España): relación con las lithofacies y el gra-diente proximal-distal de la plataforma. Revista Española de Paleontología 20, 21–36.

Reolid, M., 2007. Taphonomy of the Oxfordian-Lowermost Kimmeridgian siliceoussponges of the Prebetic Zone (Southern Iberia). Journal of Taphonomy 5, 71–90.

Reolid, M., 2008. Taphonomic features of Lenticulina as a tool for paleoenvironmentalinterpretation of mid-shelf deposits of the Upper Jurassic (Prebetic Zone, SouthernSpain). Palaios 23, 482–494.

Reolid, M., 2011. Interactions between microbes and siliceous sponges from Upper Ju-rassic buildups of External Prebetic (SE Spain). In: Reitner, J., Quéric, N.V., Arp, G.(Eds.), Advances in Geobiology of Stromatolite Formation. Lecture Notes in EarthSciences, 131, pp. 319–330.

Reolid, M., Gaillard, C., 2007. Microtaphonomy of bioclasts and paleoecology of micro-encrusters from Upper Jurassic spongiolithic limestones (External Prebetic, South-ern Spain). Facies 53, 97–112.

Reolid, M., Nagy, J., 2008. Jurassic transgressive-regressive cycles in carbonate and sili-ciclastic shelf facies: different response of foraminiferal assemblage trends to sea-level changes. Grzybowski Foundation Special Publication 13, 199–213.

Reolid, M., Gaillard, C., Olóriz, F., Rodríguez-Tovar, F.J., 2005. Microbial encrustationsfrom the Middle Oxfordian–Earliest Kimmeridgian lithofacies in the PrebeticZone (Betic Cordillera, southern Spain): characterization, distribution and control-ling factors. Facies 50, 529–543.

Reolid, M., Abad, I., Martín-García, J.M., 2008a. Palaeoenvironmental implications offerruginous deposits related to a Middle–Upper Jurassic discontinuity (PrebeticZone, Betic Cordillera, Southern Spain). Sedimentary Geology 203, 1–16.

Reolid, M., Nagy, J., Rodríguez-Tovar, F.J., Olóriz, F., 2008b. Foraminiferal assemblages aspaleoenvironmental bioindicators in Late Jurassic epicontinental platforms: rela-tion with trophic conditions. Acta Palaeontologica Polonica 53, 705–722.

Reolid, M., Rodríguez-Tovar, F.J., Nagy, J., Olóriz, F., 2008c. Benthic foraminiferal mor-phogroups of mid to outer shelf environments of the Late Jurassic (Prebetic Zone,Southern Spain): characterization of biofacies and environmental significance.Palaeogeography, Palaeoclimatology, Palaeoecology 261, 280–299.

Rey, J., Cubaynes, R., Fauré, P., Hantzpergue, P., Pélisié, T., 1988. Stratigrapie séquencelleet évolution d'une plate-forme carbonatée: le Jurassique du Quercy (Sud-Ouets dela France). Compte Rendus de l'Academie des Sciences de Paris (II) 306,1009–1015.

Rey, J., Bonnet, L., Cubaynes, R., Qajoun, A., Ruget, C., 1994. Sequence stratigraphy andbiological signals: statistical studies of benthic foraminifera from Liassic series.Palaeogeography, Palaeoclimatology, Palaeoecology 111, 149–171.

Rodríguez-Tovar, F.J., 1993. Evolución sedimentaria y ecoestratigráfica en plataformasepicontinentales del Margen Subibérico durante el Kimmeridgiense inferior.Ph.D. Thesis, Univ. Granada, 377 pp.

Rodríguez-Tovar, F.J., Reolid, M., Pardo-Igúzquiza, E., 2010. Planktonic versus benthicforaminifera response to Milankovitch forcing (Late Jurassic, Betic Cordillera): test-ing methods for cyclostratigraphic analysis. Facies 56, 459–470.

Salas, R., Guimerá, J., Mas, R., Martín-Closas, C., Meléndez, A., Alonso, A., 2001. Evolutionof the Mesozoic Central Iberian Rift System and its Cainozoic Inversion (IberianChain). In: Cavazza, W., Roberston, A.H.F.R., Ziegler, P. (Eds.), Peri-Tethyan Rift/Wrench Basins and Passive Margins: Museum National d'Histoire Naturelle,Paris, Memoires, 186, pp. 145–185.

Sarg, J.F., 1988. Carbonate sequence stratigraphy. In: Wilgus, C.K., Hastings, B.S., Ken-dall, C.G. (Eds.), Sea-level Changes: An Integrated Approach: Society for Sedimen-tary Geology (SEPM) Special Publication, 42, pp. 155–182.

Schlager, W., 1991. Depositional bias and environmental change— important factors insequence stratigraphy. Sedimentary Geology 70, 109–130.

Schlager, W., 1992. Sedimentology and sequence stratigraphy of reefs and carbonateplatforms: American Association of Petroleum Geologists (AAPG). Continuing Edu-cation Course Note Series, 34. 71 pp.

Schlager, W., 2005. Carbonate sedimentology and sequence stratigraphy. Concepts inSedimentology and Paleontology, 8. Society for Sedimentary Geology (SEPM).200 pp.

Schmiedl, G., Pfeilsticker, M., Hembleben, C., Mackensen, A., 2004. Environmental andbiological effects on the stable isotope composition of recent deep sea benthic fo-raminifera from the western Mediterranean sea. Marine Micropaleontology 51,129–152.

Scouflaire, Q., Marchand, D., Bonnot, A., Courville, P., Raffray, M., Huault, V., 1997. Lecontact Callovien-Oxfordien dans les environs de Chaignay: nouvelles donnéesstratigraphiques et paléontologiques. Bulletin Sciences Bourgogne 49, 45–63.

Sokolov, B.K., 1988. Ecostratigraphy, its place and role in modern stratigraphy. Interna-tional Geology Review 30, 3–10.

Strasser, A., 2007. Astronomical time scale for theMiddle Oxfordian to Late Kimmeridgianin the Swiss and French Jura Mountains. Swiss Journal of Geosciences 100, 407–429.

Strasser, A., Hillgärtner, H., Hug, W., Pittet, B., 2000. Third-order depositional sequencesreflecting Milankovitch cyclicity. Terra Nova 12, 303–311.

Tomašových, A., Zuschin, M., 2009. Variation in brachiopod preservation along a car-bonate shelf-basin transect (Red Sea and Gulf of Aden): environmental sensitivityof taphofacies. Palaios 24, 697–716.

Tyszka, J., 1994. Response of Middle Jurassic benthic foraminiferal morphogroups todysoxic/anoxic conditions in the Pieniny Klippen Basin, Polish Carpathians. Palaeo-geography, Palaeoclimatology, Palaeoecology 110, 55–81.

Vail, P.R., Mitchum, R.M., Thompson, S., 1977. Seismic stratigraphy and global changesof sea level, part 3: global cycles of relative changes of sea level. In: Payton, C.E.(Ed.), Seismic stratigraphy – Applications to hydrocarbon Exploration: AmericanAssociation of Petroleum Geologists (AAPG) Memoir, 26, pp. 63–81.

Vail, P.R., Colin, J.P., Chene, R.J., Kuchly, J., Mediavilla, F., Triflieff, V., 1987. La stratigra-phie séquencelle et son application aux corrélations chronostratigraphiques dans


le Jurassique du bassin de Paris. Bulletin de la Société Géologique de France 8,1301–1321.

Vail, P.R., Audemard, F., Bowman, S.A., Eisner, P.N., Pérez-Cruz, C., 1991. The stratigraphicsignatures of tectonics, eustasy and sedimentology: an overview. In: Einsele, G.,Ricken, W., Seilacher, A. (Eds.), Cycles and Events in Stratigraphy. Springer-Verlag,Berlin, pp. 617–659.

Valentine, J.W., 1973. Evolutionary Paleoecology of the Marine Biosphere. Prentice-Hall, Englewood Cliffs, NJ. 510 pp.

Van der Zwaan, G.J., Duijnstee, I.A.P., Den Dulk, M., Ernst, S.R., Jannink, N.T., Kouwenho-ven, T.J., 1999. Benthic foraminifers: proxies or problem? A review of paleoecolog-ical concepts. Earth-Science Reviews 46, 213–236.

Van Wagoner, J.C., Mitchum, R.M., Posamentier, H.W., Vail, P.R., 1987. An overview ofsequence stratigraphy and key definitions. In: Bally, A.W. (Ed.), Atlas of SeismicStratigraphy, Vol. 1: American Association of Petroleum Geologists (AAPG) Studiesin Geology, 27, pp. 11–14.

Van Wagoner, J.C., Posamentier, H.W., Mitchum, R.M., Vail, P.R., Sarg, J.F., Loutit, T.S.,Hardenbol, J., 1988. An overview of the fundamentals of sequence stratigraphy

and key definitions. In: Wilgus, C.K., Hastings, B.S., Kendall, C.G., Posamentier, H.W.,Ross, C.A., Van Wagoner, J.C. (Eds.), Sea-level Changes: An integrated approach: Soci-ety for Sedimentary Geology (SEPM) Special Publication, 42, pp. 39–45.

Walker, K.R., Bambach, R.K., 1971. The significance of fossil assemblages from finegrained sediments: time-averaged communities. Geological Society of America Ab-stracts & Programs 3, 783–784.

Waterhouse, J.B., 1976. The significance of ecostratigraphy and need for biostratigraph-ic hierarchy in stratigraphic nomenclature. Lethaia 9, 317–325.

Yin, H., Tong, J., Zhang, K., Wu, S., 1997. Application of ecostratigraphy to sequence stra-tigraphy. Science in China, Series D, Earth Sciences 40, 137–144.

Ziegler, M.A., 1962. Beiträge zur Kenntnis des unteren Malm im zentralen Jura. Mittei-lungen aus dem Geologischem Institut der Eidgen. Technischen Hochschule undder Universität Zurich 82, 1–51.

Documents

Palaeogeography and relative sea-level history forcing eco-sedimentary contexts in Late Jurassic epicontinental shelves (Prebetic Zone, Betic Cordillera): An ecostratigraphic approach