Disponible en ligne sur www.sciencedirect.

com

Geobios 41 (2008) 3142 http://france.elsevier.com/direct/GEOBIO

Original article

Taphonomy of fossil macro-invertebrate assemblages as a tool for ecostratigraphic interpretation in Upper Jurassic shelf deposits (Prebetic Zone, southern Spain) ` ` Apport de la taphonomie des assemblages fossiles a macro-invertebres a linterpretation ecostratigraphique des depots de plate-forme du Jurassique superieur (zone Prebetique, Sud de lEspagne)
Federico Olriz a,*, Matas Reolid a,b, Francisco J. Rodrguez-Tovar a
a

Departamento Estratigrafa y Paleontologa, Facultad de Ciencias, Universidad de Granada, Fuentenueva s/n, 18002 Granada, Spain b Departamento Geologa, Universidad de Jan, Campus Las Lagunillas 23071 Jan, Spain Received 27 October 2005; accepted 2 March 2006 Available online 26 December 2007

Abstract Composition and taphonomy of macro-invertebrate fossil assemblages, together with facies analysis, have been approached in order to interpret shifting paleoenvironmental conditions in the External Prebetic (S-SE Spain) during the early Late Jurassic (Middle Oxfordian). In oolitic and spongiolitic limestones, the size of fossil remains, mode of preservation, within-bed position, corrasion, fragmentation, epibiont and biogenic encrustation, disarticulation and uncoupling, allow recognition of two taphofacies, respectively. Identied ecostratigraphic events and trends accord with rapid ooding under high-energy conditions related to ecospace enlargement for cephalopods and then the persistence of lower energy, long-lasting exposure of skeletals and higher sedimentary rates. The paleoenvironmental interpretation is consistent with neritic environments shifting from shallow carbonate to hemipelagic sedimentation and enlarging of shelf ecospace for marine invertebrates. # 2007 Elsevier Masson SAS. All rights reserved. Rsum Lanalyse taphonomique et la composition des associations fossiles principalement constitues par des macro-invertbrs, en combinaison avec une analyse de facis, sont utilises pour interprter des changements paloenvironnementaux au dbut du Jurassique suprieur (Oxfordien moyen) dans le Prbtique Externe (S-SE Espagne). Dans les calcaires oolitiques et spongiolitiques, la taille des fossiles, leur prservation, orientation dans les horizons sdimentaires, corrasion, fragmentation, encrotement biognique, dsarticulation et dcouplage de moules internes permettent la reconnaissance de deux taphofacis, respectivement. Lidentication dvnements et tendances costratigraphiques est en accord avec une inondation rapide sous conditions de haute nergie et agrandissement de lespace cologique pour les cphalopodes, suivi de conditions de basse nergie persistantes avec exposition de longue dure pour les coquilles et un taux plus lev de sdimentation. Linterprtation paloenvironnementale est en accord avec le passage dune sdimentation carbonate environnement nritique hmiplagique accouple llargissement de lespace cologique pour les invertbrs marins. # 2007 Elsevier Masson SAS. All rights reserved.
Keywords: Macro-invertebrate assemblages; Taphonomy; Taphofacies; Palaeoenvironment; Ecostratigraphy; Oxfordian; Prebetic Zone (SE Spain) Mots cls : Assemblages de macro-invertbrs ; Taphonomie ; Taphofacis ; Paloenvironnement ; Ecostratigraphie ; Oxfordien ; zone Prbtique (Sud de lEspagne)

* Corresponding author. E-mail address: foloriz@ugr.es (F. Olriz). 0016-6995/$ see front matter # 2007 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.geobios.2006.03.003

32

F. Olriz et al. / Geobios 41 (2008) 3142

1. Introduction Ecostratigraphy, or ecosystem stratigraphy, was rst described as the natural development of biostratigraphy, to give paleoecological support to the temporal subdivision of the fossil record (Martinsson, 1973; Boucot, 1982; Sokolov and Kaljo, 1986, among others), working on stratigraphic resolution at the orders of time intervals as ne as 105 years (Rabe and Cisne, 1980; Martin, 1990). In this context, ecostratigraphic events, determined by relatively short-time ecological changes, are especially informative (Kauffman, 1988). Thus, the potential relationship between changes in ecospace, as multidimensional ecological volume and lithofacies reinforces the usefulness of ecostratigraphic interpretations for analyzing major traits of the geobiological evolution in a given basin. Hence, ecostratigraphic approaches could serve as a key for rening recent innovations in basin analysis (i.e., Olriz et al., 1993, 1994, 1995, 1996, 2002b, 2004a, 2004b; RodrguezTovar, 1993; Reolid, 2005). To complement ecostratigraphic events, the recognition of ecostratigraphic trends (Olriz et al., 1995) informs about lengthy uctuations in faunal compositions, and therefore about ecology at a longer term, which is of special relevance for interpretation of palaeoenvironmental dynamics as an integrated aspect of basin evolution. In the ecostratigraphic procedure, the truly important aspect is the interdisciplinary approach applied to fossils and associated sedimentary rocks, on the basis of a bed-by-bed analysis (Boucot, 1982). In order to complement ecological and depositional interpretations, the integration of taphonomic analysis into the ecostratigraphic approach is crucial. Taphonomic analyses of macro-invertebrate assemblages have revitalize modern interpretation of both depositional and paleoecological conditions, as well as that of ecological dynamics in a hierarchical frame (Speyer and Brett, 1988; Miller, 1990; Allison and Briggs, 1991; Kidwell and Bosence, 1991; Parsons and Brett, 1991; Brett and Baird, 1993; Kidwell, 1993; Kowalewski et al., 1994; Sageman et al., 1997; Smith and Nelson, 2003). All of this is of special relevance for stratigraphic analysis within modern basin research based on outcrops records (i.e., sequence stratigraphy and related approaches, Rollins et al., 1990; Brett, 1995; Pittet et al., 2002). In the Jurassic of Iberia, ammonite taphonomy provides valuable data to interpret the ecological and depositional evolution in shelf environments (e.g., Fernndez-Lpez and Melndez, 1994; Fernndez-Lpez, 1997, 2000; Olriz et al., 2002b, 2004b). The combined analysis of fossil preservation and lithofacies enables the identication of taphonic populations, taphonomic trends, and taphofacies to improve ecostratigraphic and sequence stratigraphic interpretations in Upper Jurassic sections from the south-Iberian paleomargin (Olriz et al., 1996, 2002b, 2004b). During the late-Middle to the earliest-Late Jurassic plate dynamics affected the North Sea Graben, the growing central North Atlantic, and regions related to the Tethys, determining ooding events across European shelves under variable tectonic forcing (e.g., Marques et al., 1991; Norris and Hallam, 1995; Jacquin et al., 1998; Gygi et al., 1998; Leinfelder and Wilson,

1998; Olriz et al., 2003a,b; and references therein). In southern Europe, Middle/Upper Jurassic boundary deposits from Tethyan epicontinental areas are usually related to ferruginous deposits associated with discontinuity surfaces. These deposits have been registered in Iberia (the Prebetic Zone of the Betic Cordillera; Acosta, 1989; Olriz et al., 2004b; Reolid, 2005, the Iberian Range; Aurell et al., 1994, 1999; Ramajo and Aurell, 1997; Ramajo et al., 2002, and the eastern Algarve Basin; Marques et al., 1991; Leinfelder, 1993); in France (the western Subalpine Basin; Dromart, 1989, Cte dOr; Courville and Collin, 1997, Schaignay; Scouaire et al., 1997, Haute-Marne; Collin and Courville, 2000, and the southeastern Paris Basin; Lorin et al., 2004); and in Switzerland (the Jura region; Ziegler, 1962; Gygi, 1969, 1981; Huber et al., 1987). Focused on characterizing Middle-Upper Jurassic boundary deposits in Western Europe, Norris and Hallam (1995) related ferruginous-oolitic deposition resting on shallow carbonates to result from the onset of wide-regional transgression, which has been largely recognized in Spain following a non-sequence stratigraphic interval embracing the Callovian/Oxfordian boundary (e.g., Bulard et al., 1979; Ramon et al., 1992; Aurell et al., 2002, 2003; Barn et al., 2004; Martn-Algarra and Vera, 2004). Across southern Iberia (the Algarve Basin included), the inner Prerift in Morocco, and the Lusitanian Basin, Marques et al. (1991) interpreted a complex unconformity DIII embracing Middle/Upper Jurassic boundary local deposits and widespread non-sequences, which relate to their Callovian Oxfordian Crisis identied in Terthyan and Atlantic areas. Courville and Collin (2002) interpreted lowermost Oxfordian deposition in the Paris Basin to result in complex sequences. Vera et al. (2004) recognized the character of complex discontinuity to the top of the Middle Jurassic, with potential over-imposition of at least three late Bathonian discontinuities, underlying oldest Late Jurassic deposition in pelagic swells of the South-Iberian paleomargin. Within the Mesozoic epicontinental shelf-system developed in the south-Iberian paleomargin, the External Prebetic provides a record of mid-shelf Upper Jurassic deposits (spongiolithic limestones and marl-limestone rhythmites) overlying inner-shelf LowerMiddle Jurassic sediments (oolitic and dolomitic limestones). Occasionally, the local record of ferruginous oolitic limestones overlying the stratigraphic discontinuity on top of the Middle Jurassic carbonates (Olriz et al., 2004a, 2004b) represents deposition during transitions phases to mid-shelf environments and then hemipelagic conditions in the area. The aim of this paper is to interpret the course of ecological and depositional changes related to switching from carbonate to carbonate-siliciclastic shelf occurred during the early Late Jurassic in a mid-shelf setting in the External Prebetic that rst received ferruginous-oolitic and then spongiolithic carbonate deposition. Hence, to propose a local scenario for the initial ooding of the Middle Jurassic carbonate platform based on the comparative analysis of composition and taphonomy of macroinvertebrate fossil assemblages recorded in the ferruginous oolitic limestones and the overlying spongiolithic limestones.

F. Olriz et al. / Geobios 41 (2008) 3142

33

2. Geological setting The studied outcrop (Fig. 1) is located in the External Prebetic that is the outermost, landward fringe of the Betic Cordillera (southeastern Iberia). Paleogeographically, the Prebetic represents part of the epicontinental system developed in the south-Iberian margin during the Jurassic (see Olriz et al., 2002a for extended treatment). The preserved External Prebetic constituted a carbonate inner-shelf environment during the EarlyMiddle Jurassic, and a carbonate-siliciclastic midshelf environment during the Late Jurassic (Middle Oxfordian to EarlyMiddle Kimmeridgian, three-fold division). Jurassic deposits in the External Prebetic correspond to the Chorro Formation and the Lorente Formation (Pendas, 1971).

The Chorro Formation consists of ca. 400 m thick Lower Jurassic dolomitized limestones, and 50 m of Middle Jurassic oolitic limestones with megaripples and oncoliticbioclastic facies. This formation represents a shallow environment undergoing carbonate sedimentation and short episodes of continental deposition; its upper part shows oolitic bars. Upper Jurassic (Middle Oxfordian to Lower Kimmeridgian) deposits correspond to the Lorente Formation, and constitute the rst interval of pelagichemipelagic sedimentation in the southeastern epicontinental system occurring in Iberia during the Mesozoic. The comparatively most proximal, landward preserved sectors of the platform (External Prebetic) are mainly represented in the Lorente Formation by a 70100 m thick succession made of spongiolithic limestones and marl-limestone

Fig. 1. Location and geological sketch of the Betic Cordillera (A) and central sector of External Prebetic (B). Synthetic lithological succession for Jurassic rocks in Sierra de Cazorla according to Garca-Hernndez and Lpez-Garrido (1988) (C). Fig. 1. Situations gographique et gologique de la Chane Btique (A) et de la partie centrale du Prbtique Externe (B). Succession lithologique synthtique des roches jurassiques dans la Sierra de Cazorla, daprs Garca-Hernndez et Lpez-Garrido (1988) (C).

34

F. Olriz et al. / Geobios 41 (2008) 3142

rhythmites (Rodrguez-Tovar, 1993; Olriz et al., 2003a,b; Reolid, 2005). The Riogazas-Chorro section (RGCH) crops out in the Sierra de Cazorla area (central sector of the External Prebetic; Fig. 1). The RGCH section is located in the El Chorro sheet (Foucault, 1971), and includes the contact between the Chorro Formation and the Lorente Formation. The studied section (Fig. 1) is 1.1 m thick and shows, from bottom to top, one or locally two limestone beds containing ferruginous ooids and pisoids (Middle Oxfordian Plicatilis Zone), three limestone beds (% 0.7 m total) characterised by a high content of sponge remains (Middle Oxfordian Transversarium Zone), and then several meters of alternating marls and limestones (Upper Oxfordian Bifurcatus Zone to Lower Kimmeridgian Platynota Zone). 3. Methodology

ammonite, bivalve, brachiopod and gastropod carcasses. The size of skeletal remains has been controlled to recognize its potential relationship with their variable completeness. Size quantication was made taking into account maximum lengths to allow identication of size intervals. For fragmentation and corrasion, the fragmentation index (F i) and the corrasion index (Ci) sensu Olriz et al. (2002b, 2002c, 2004b) were used. The Ci informs about the degree of corrasion of a group of samples, calculated as the mean value obtained by summing the products of the number of samples (n) presenting different degrees of corrasion [high corrasion degree (HCD, > 60% worn) 100; medium corrasion degree (MCD, 1060% worn) 50; low corrasion degree (LCD, < 10% worn) 1], 1], and dividing the resulting value by the total number of samples considered (N), including those presenting no signs of corrasion. Ci nHCD 100 nMCD 50 nLCD 1 N

Sedimentological and paleontological analyses were performed separately for each of the two studied facies: ferruginous oolitic limestones and spongiolithic limestones. Outcrop observations were complemented by microscopic analysis on polished slabs and thin sections, under transmittedlight microscope and binocular magnifying glass, for characterizing microfacies. Macro-invertebrate assemblages were studied following the ecostratigraphic procedure used previously (Olriz et al., 1993, 1994, 1995). Remains of fossil macro-invertebrates (including fragments and complete specimens) were collected bed-by-bed together with preliminary taphonomic observations in the RGCH section. To avoid counting problems, sponge remains were excluded from the analysis (because of the difculty in their extraction and subsequent evaluation). Other remains that are normally abundant and disarticulated (e.g., crinoids) were excluded because of the impossibility of determining the number of individuals. Thus, sponges and crinoids were analysed only qualitatively. Laboratory processing of samples yielded a total of 775 specimens of macro-invertebrates, which were characterized as follows. Three compositional spectra were considered and represented in pie-diagrams: spectrum A as a general picture representing the relative abundance in ammonoids, belemnoids and benthos; spectrum B reecting the internal composition of ammonoids assemblages (perisphinctoids, Sowerbyceras, Phylloceratina and Lytoceratina, haploceratids and others Ammonitina; Olriz et al., 1995); and spectrum C applied to show the internal composition of benthic assemblages (brachiopods, bivalves, regular echinoids, irregular echinoids and others; mainly gastropods and ahermatypic corals). In addition, taphonomic features were characterized in detail, including: mode of preservation, size of skeletal remains, within-bed position, corrasion (sensu Brett and Baird, 1986), fragmentation, epibiont and biogenic encrustation, disarticulation and uncoupling (sensu Olriz et al., 2002b, 2002c, 2004b). The preservation mode refers to both, the relative occurrence of inner moulds with and/or without neomorphic shell and the type of sediments that lled up

The F i indicates the degree of fragmentation among a set of individuals, and is calculated as the mean value obtained by summing the products of the number of individuals (n) presenting different degrees of fragmentation [high fragmentation degree (HFD, remains are little representative of original dimensions and shape) 100; medium fragmentation degree (MFD, fragmentation affect appreciably the shape and size) 50; low fragmentation degree (LFD, fragmentation that does not affect signicantly the shape and size) 1], and dividing the resulting value by the total number of samples considered (N), including those with no fragmentation. Fi nHFD 100 nMFD 50 nLFD 1 N

Uncoupling refers to disaggregation in internal moulds of embedded ammonoids, in relation to location of septa and whorl overlapping (sensu Olriz et al., 2002b, 2002c, 2004b; phragmocone disarticulation sensu Fernndez-Lpez and Melndez, 2004). The combined analysis of lithofacies, composition and taphonomic features in macro-invertebrate assemblages will allow to identify taphofacies to be used as a standard tool in reconstructing ancient depositional environments (e.g., Brett and Baird, 1986; Speyer and Brett, 1986, 1988; Kowalewski et al., 1994; Olriz et al., 2002b, 2002c; Zuschin et al., 2003). 4. Material and database 4.1. Ferruginous oolitic limestone Ferruginous oolitic deposits in the Prebetic Zone show thin, local deposition related to the Middle/Upper Jurassic boundary. A planar, ferruginous surface, or a thin ferruginous crust, separates Upper Jurassic ferruginous oolitic limestones from Middle Jurassic carbonates. Generally, ferruginous deposits pinch-out and their thicknesses appear to be related to the number of beds (usually one or two). In the RGCH section, this lithofacies consists of two beds showing thickness variation

F. Olriz et al. / Geobios 41 (2008) 3142

35

Fig. 2. Stratigraphic column for the Riogazas-Chorro section showing composition and mean values of macro-invertebrate fossil assemblages and taphonomic features, both of these gathered from the ferruginous oolitic limestone (lower pie-diagrams) and the spongiolithic limestone (upper pie-diagrams). Fig. 2. Colonne stratigraphique de la section de Riogazas-Chorro indiquant la composition et les valeurs moyennes des assemblages fossiles macro-invertbrs, ainsi que les caractristiques taphonomiques.

between 10 and 40 cm and a decreasing upwards abundance in ferruginous ooids (Fig. 2). The lower boundary is irregular and locally related to the record of a ferruginous crust. The ferruginous oolitic limestone shows a grain-supported bioclastic fabric (packstone) (Olriz et al., 2004b; Reolid, 2005) with abundant peloids (30%), ferruginous ooids and pisoids (26%), and bioclasts (26%). Other grains, including quartz, are secondary. Among bioclasts, echinoderm remains, mollusc fragments (mainly ammonoids) and foraminifera dominate. Planktic foraminifera are dominant (82%), while benthic foraminifera are scarcer (mainly spirillinids, nodosariids, agglutinated forms, and Epistomina). Notable is the combined record of Epistomina and Trocholina, as well as the high abundance of thick-shelled, great-size, fragmented specimens of Lenticulina (72% of the record of this genus; Olriz et al., 2004b; Reolid, 2005).

4.1.1. Fossil assemblages of macro-invertebrates A total of 600 specimens, including fragments, were analyzed from the ferruginous oolitic limestone lithofacies. The macro-invertebrate fossil assemblage (Fig. 2) is composed of ammonoids (70%), belemnoids (21%), and benthics (9%). Among ammonoids, phylloceratids are dominant (28%, with genus Sowerbyceras representing 11% of ammonoids), followed by perisphinctoids (14%). Dominant benthics are gastropods, ahermatypic corals and bivalves. Belemnoids are especially frequent in the lower part of the lithofacies overlying the ferruginous crust, but their record decreases upward. 4.1.2. Taphonomic analysis Taphonomic analysis (Olriz et al., 2004b) revealed a high abundance of specimens per rock volume (Fig. 3), the frequent preservation of neomorphized carbonate shells,

36

F. Olriz et al. / Geobios 41 (2008) 3142

Fig. 3. [upper left] Lower surface of ferruginous oolitic limestone bed showing abundant belemnoids (B) and ammonoids (A). [upper right] Spongiolithic limestone with common sponge remains (Sp) and crinoid stem (Cr). Lower views showing ferruginous oolitic limestone with abundant fossil remains such as belemnite rostra (B), ammonite phragmocones with neomorphized shell-walls and septa (A), and indeterminate shells. Scale bar = 1 cm. Fig. 3. [En haut gauche] Surface infrieure dun banc calcaire oolitique ferrugineux riche en blemnites (B) et en ammonites (A). [En haut droite] Calcaire ponges (Sp) et tige de crinode (Cr). (Photos du bas) Calcaire oolitique ferrugineux avec dabondants restes de fossiles comme des rostres de blemnites (B), des phragmocnes dammonites coquilles nomorphoses (A), ainsi que des coquilles indtermines. chelle = 1 cm.

which originally were aragonitic carcasses (ammonoids and gastropods), as well as macrocrystalline calcite lling in the innermost chambers (e.g., gastropods, ammonoids). Fossil remains size smaller than 30 mm dominates (Fig. 2), especially in the case of ahermatypic corals (9.0 mm mean-size) and gastropods (11.3 mm mean-size). Macrofossils showing larger mean-size are belemnoids (37.7 mm) and Phylloceratina (28.1 mm). The largest belemnoids are located at the base of the lowermost bed (Fig. 3A). The most frequent within-bed orientation of fossil remains is oblique to quasi-vertical (61%), while quasi-horizontal orientation is most common among larger specimens. The orientation of shell fragments is mainly concave up (61%). Features revealing long-lasting exposure of skeletals on the sea-oor are rarely observed, as indicated by low frequencies of corrasion (Ci < 2%), microbial encrustation (absent), and macroscopic epibionts (rare; < 3% serpulids). In contrast, the mean value of fragmentation is high (F i = 55%), especially in ammonoids (higher in perisphinctoids 67% and lower in Sowerbyceras 29%). Evidences of uncoupling were recorded only rarely. In the

case of benthic organisms, disarticulation is variable; common in bivalves (100%) and echinoderms (100%) and less frequent in brachiopods (25%). 4.2. Spongiolithic limestone Spongiolithic limestones crop out in the External Prebetic and are developed as well-stratiphied limestones, with bedthickness ranging from 12 to 50 cm, and a high abundance of siliceous sponge remains (Olriz et al., 2003a,b; Reolid, 2005; Fig. 3B). In the RGCH section, this lithofacies overlies the ferruginous oolitic limestones and consists of three beds 24, 20 and 23 cm thick (Fig. 2). The fabric under microscope is grainto matrix-supported, microfacies being characterised as wackestone (locally packstone) mainly composed of bioclasts (25%), oncoids (23%), peloids (18%), ooids (13%), lumps (10%) and tuberoids (5.5%), showing secondary aggregate grains and quartz. The most abundant bioclasts are echinoderm remains, indeterminate mollusc fragments, bivalves and foraminifera. Benthic foraminifera (agglutinated forms, spir-

F. Olriz et al. / Geobios 41 (2008) 3142

37

illinids, nubeculariids, ofthalmidiids and nodosariids) are dominant (62%) while planktics are scarce. 4.2.1. Fossil assemblages of macro-invertebrates A total of 175 fossil macro-invertebrates were analysed (Fig. 2), showing the clear dominance of ammonoids (69%), especially Sowerbyceras (44% of ammonoids). Benthic organisms represent 19% of the macro-invertebrate assemblages, among which infaunal forms are dominated by irregular echinoids (51% of benthics), while brachiopods (21% of benthics) dominates suspension-feeding epifauna, aside from the occurrence of abundant sponge remains and crinoids (only qualitatively analyzed; see above). Siliceous sponges correspond to Dictyida, Lychniskida and Lithistida. 4.2.2. Taphonomic analysis The taphonomic analysis (Fig. 2) reveals a high proportion of small size macrofossils (<30 mm) per rock volume, apart from sponges. The smallest remains belong to brachiopods and irregular echinoids (mean size of 17 mm and 23 mm, respectively). Perisphinctids are the largest macro-invertebrates, except for sponges. The frequency of specimens with a quasi-horizontal within-bed orientation is notable (51%) and dish-shaped sponges are frequently overturned. Aragonitic shells (e.g., ammonoids, gastropods and some bivalves) are absent, or preserved as neomorphic calcite. Original preservation of shell material is frequent in belemnoids (rostra), brachiopods and echinoids, and less common in bivalves. Siliceous sponges are preserved as sponge mummies (Schwamm-Mumien in Fritz, 1958), in which neomorphic calcite spicules maintain their original arrangement. Recrystallization of spicules occurred early during diagenesis, with initial transformation from opaline to crystalline silica (Hartman et al., 1980) and subsequent replacement by calcite. There are no marked differences in sediment composition between moulds of fossil remains and that of the matrix surrounding them. Nevertheless, microcrystalline calcitic llings are sometimes observed within small brachiopods (rhynchonellids). In spongiolithic limestones, the corrasion index (24%) and the colonization by macroscopic epibionts (32% serpulids and encrusting foraminifera) are comparatively high, with corrasion being more pervasive on upward-oriented surfaces of the largest ammonoid shells. The fragmentation index is 34% (Fig. 2), being lower for echinoids and brachiopods. Fragmentation in sponges could be related to size; the size of dish-shaped sponges is usually around 15 cm, and their thickness does not usually exceed 1 cm. These size ratios indicate that sponges were delicate structures, liable to fragmentation and then disintegration before burial of their spicules. Nevertheless, despite their apparent fragility, many virtually complete sponge specimens have been found (Olriz et al., 2003a,b). There is not disarticulation registered in brachiopods. In contrast, disarticulation is common in echinoderms, although stems of crinoids preserved with many ossicles (eight to 15)

exist. There is not occurrence of specimens of ammonoids showing uncoupling. The biogenic encrustation on sponges by annelids (serpulids and terebellids) and encrusting foraminifera, as well as by microbialite growths, is very frequent (see Olriz et al., 2003a,b; Reolid, 2005). Benthic microbial communities and nubeculariids dominate in upward-oriented surfaces of skeletal remains and sponges while annelids colonized downwardoriented ones. The taphonomic analysis reveals no difference in biogenic encrustation polarity between sponges recorded in a live position or overturned; thus, colonisation after death is assumed (Gaillard, 1983; Olriz et al., 2003a,b). 5. Interpretation Composition and taphonomy of Middle Oxfordian macroinvertebrate assemblages in the RGCH section allow to interpret ecologic and depositional parameters related to the environmental dynamics (e.g., ecospace, proximal-distal gradient or relative distance from the shore, exposure time, water energy and both velocity i.e. accumulation rate and rate of sedimentation) occurring at the beginning of pelagichemipelagic sedimentation in the area, early during the Late Jurassic (Figs. 4 and 5). Among the taphonomic features analyzed, corrasion, encrustment and the presence of epibionts relate to the exposure time at the seaoor, which is directly proportional to the rate of sedimentation. Fragmentation, disarticulation and uncoupling mainly point to physical controls of deposition. The combination of physical and biogenic controls determined mean-size of skeletals and withinbed position, while corrasion and the mode of preservation would be mainly forced by physical-chemical factors. The relative signicance of taphonomic features and their relation to lithofacies allow recognition of two taphofacies:  Taphofacies I, recognized in the ferruginous oolitic limestones. The macro-invertebrate assemblages are characterized by:  High occurrence of pelagic-to-demersal, carnivores-tonecrophagous organisms (ammonoids, mainly phylloceratids) and scarce benthos made of suspension-feeders, microcarnivorous (ahermatypic corals) and carnivorous (gastropods),  Frequent macrocrystalline calcite lling of carcasses,  High fragmentation and dominance of bioclasts in an oblique to quasi-vertical within-bed orientation,  Near absence of evidence for prolonged exposure (low corrasion, absence of microbial encrustation, and rare occurrence of macroscopic epibionts);  Taphofacies II, recognized in the overlying spongiolithic limestones. The macro-inverbrate assemblages are characterized by:  Higher occurrence of benthos (epibenthos, mainly sponges, and endobenthos) and poorer record of phylloceratids,  Lower abundance of macrocrystalline calcite lling of carcasses and fragmentation,  Over-dominance of bioclasts with a quasi-horizontal position,

38

F. Olriz et al. / Geobios 41 (2008) 3142

Fig. 4. Environmental reconstruction for Middle Jurassic oolitic limestones and the overlying deposits analyzed. (A) Oolitic limestone corresponding to Middle Jurassic oolitic bars containing terebratulids and bivalves, and local colonization by corals. (B) Middle Oxfordian ferruginous oolitic limestone showing abundant ammonoids and belemnoids, episodes of high-energy conditions and the resulting preservation context. (C) Middle Oxfordian spongiolithic limestone showing common benthic organisms (sponges, crinoids, echinoids, brachiopods, bivalves and serpulids), a more stable low-energy environment, lower accumulation rate, and the resulting preservation context.

F. Olriz et al. / Geobios 41 (2008) 3142

39

Fig. 5. Traits selected from both litho- and bioclasts and the interpreted environment, relative sea level and ecostratigraphy, all these according to ammonite biostratigraphy. Ecostratigraphic events (EE), ecostratigraphic trend (ET), high energy (HE), low energy (LE). Fig. 5. Paramtres slectionns partir des bioclastes et lithoclastes, environnements interprts, niveau marin relatif et signals costratigraphiques. Information cale sur lchelle biostratigraphique ammonites. Abrviations : EE : vnements costratigraphiques ; ET : tendances costratigraphiques ; HE : haute nergie ; LE : basse nergie.

 Longer bottom exposure of skeletal remains and inner moulds (higher corrasion and colonization by epibionts). The taphonomic traits of the spongiolithic limestone are congruent with Taphofacies II described by Olriz et al. (2002b). These features reveal that the beginning of pelagic sedimentation over the Middle Jurassic carbonate-shelf in the studied area, which resulted in ferruginous oolitic limestones, occurred in a context of ecosedimentary, environmental turnover (Figs. 4 and 5) related to:  Rapid ooding on the shelf;  Relative high water-energy;  Ecospace enlargement favouring shelled cephalopods;

 Short exposure time of skeletal remains (Olriz et al., 2004b);  High accumulation but low sedimentation rates. The scarcity in benthic macro-invertebrates compared with spongiolithic limestones was related to foreseeable, unfavourable ecological conditions at the seabed, agreeing with proliferation of opportunistic organisms such as some gastropods and ahermatypic corals. The occurrence of microfossils and lithoclasts of variable provenance (ferruginous ooids, angular quartz grains, and Trocholina from emerged and near-shore environments, and Globuligerina and Epistomina from outer marine environments; Olriz et al., 2004b; Reolid, 2005), is consistent with rapid ooding of the shelf under highenergy conditions. This interpretation is also supported by abundance of fragmented nodosariids (mainly thick-shelled

Fig. 4. Reconstitution environnementale des calcaires oolitiques du Jurassique moyen et dpts analyss. (A) Calcaire oolitique correspondant des barrires oolitiques trbratules et bivalves, localement colonises par des coraux. (B) Calcaire oolitique ferrugineux de lOxfordien moyen form par dabondantes ammonites et blemnites, caractris par des pisodes de haute nergie et montrant son contexte de prservation. (C) Calcaire spongiolitique de lOxfordien moyen organismes benthiques communs (ponges, crinodes, chinodes, brachiopodes, bivalves et serpules), traduisant un milieu nergie basse, un taux daccumulation infrieur et son contexte de prservation.

40

F. Olriz et al. / Geobios 41 (2008) 3142

Lenticulina). Storm-dominated deposition offers an appropriate scenario for the studied ferruginous oolitic limestones, which probably related to wetter climate forced by the wide-regional transgression. Later, environmental conditions changed during deposition that resulted in the overlying spongiolithic limestones. The new environmental context was mainly related to and/or supported by:  Ecospace readjustment;  Diminishing rate of accumulation in a context of higher rate of sedimentation;  Lower environmental energy;  A longer exposure of skeletal remains before nal burial. The lower content of quartz grains, the absence of ferruginous ooids, foraminiferal assemblages showing higher values of sessile forms and lower fragmentation of Lenticulina, absence of Trocholina and scarcer presence of planktic forms, are consistent with more stable, relatively quiet, marine conditions. Two main scenarios are interpreted for idealizing the environmental change occurring in the mid-shelf External Prebetic during the beginning of the Late Jurassic (Middle Oxfordian):  Sudden environmental restructuring favourable for reception of exported materials (ferruginous oolitic limestone), including postmortem transport of macro-invertebrate carcasses in a high-energy context related to ooding events;  Later and comparatively gradual onset of a lower-energy environment, in a more open marine carbonate platform, favouring less disturbed ecologic relationship between the macro-invertebrate community and its fossil record (spongiolithic limestones). In ecostratigraphic terms, the beginning of Late Jurassic pelagichemipelagic deposition overlying inner-shelf, Lower Middle Jurassic carbonates, was related to a short-time ecological change, recorded as an ecostratigraphic event, reected by the FAD of pelagic macro-invertebrates on the carbonate shelf (macro- and micro-organisms). Afterwards, an analogous turnover, in terms of rapid change in environmental energy, favoured the return to lower-energy conditions and their persistence throughout the rest of the Middle and the Late Oxfordian. Hence, the possibility for recording and identication of the ecostratigraphic trend connected to lower-energy environments, more stable sea-bottoms, and slower burial. 6. Conclusions In the Riogazas-Chorro section (Sierra de Cazorla, External Prebetic), the analysis of Middle Oxfordian lithofacies and fossil assemblages, mainly composed by macro-invertebrates, is of value for understanding the ecological and depositional dynamics related to a major environmental change. Stratigraphic evolution of lithofacies (from ferruginous oolitic limestones to spongiolithic limestones), composition of

macro-invertebrate assemblages and taphonomy (including taphofacies), relates to changing ecosedimentary conditions after initial ooding of a Middle Jurassic carbonate shelf early during the Middle Oxfordian. This ooding forced both ecospace enlargement and the taphonomic process for skeletals on the seabed. Taphofacies I identied in ferruginous oolitic limestone is characterized by scarce benthic forms (9% of the total macroinvertebrate assemblage vs. 70% of ammonoids and 21% of belemnoids) with common macrocrystalline calcite lling of carcasses; high fragmentation (F i = 55%) and dominance of bioclasts in an oblique to quasi-vertical within-bed orientation (61% of the fossil remains); and near absence of corrasion (Ci < 2%) and encrustations. This taphofacies indicates relative high water-energy, short exposure time of skeletons and high accumulation but low sedimentation rates. Taphofacies II identied in spongiolithic limestone is characterized by macro-invertebrate assemblages with higher content of benthos (19% of the macro-invertebrate assemblages vs. 69% of ammonoids), scarcity macrocrystalline calcite lling of carcasses and fragmentation (F i = 34%); overdominance of bioclasts with a quasi-horizontal position (51% of the fossil remains); and higher corrasion (Ci = 24%) and colonization by epibionts and microbialites. This taphofacies is the result of lower environmental energy, diminishing rate of accumulation in a context of higher rate of sedimentation as well as a longer exposure of skeletals on the sea-bottom. The ecostratigraphic signature of the stratigraphic changes observed imply ecostratigraphic events related to pulses of high-energy conditions and their shift to low-energy conditions determining the beginning of an ecostratigraphic trend. Highenergy conditions involved storm dominated deposition characterizing the analysed depocentre as a setting for the reception of both bio- and lithoclasts. Subsequent low-energy conditions started suddenly and progressively determining a persistent, calm environment throughout the analyzed Oxfordian, in which fossil assemblages better inform about the real ecology of the seabed and the overlying water column. Acknowledgements This research was supported by Projects BTE3029, 1316 MCyT and the EMMI Group (RNM178 Junta de Andaluca). We are grateful to Dr. M. Kowalewski (Virginia Tech) and an anonymous reviewer for valuable suggestions and comments. References
Acosta, P., 1989. Estudio del Jursico de un sector de la Sierra de Cazorla (Zona Prebtica). Tesis Lic., Universidad de Granada. Allison, P.A., Briggs, D.E.G., 1991. Taphonomy: releasing the Data Locked in the Fossil Record. Topics in Geobiology 4. Plenum Press, New York. Aurell, M., Bdenas, B., Bello, J., Delvene, G., Melndez, G., Prez-Urresti, I., Ramajo, J., 1999. El Calloviense y el Jursico Superior en la Cordillera Ibrica Nororiental y la zona de enlace con la Cordillera Costero Catalana, en los sectores de Sierra de Arcos, Calanda y Xerta-Pals. Cuadernos de Geologa Ibrica 25, 73110.

F. Olriz et al. / Geobios 41 (2008) 3142 Aurell, M., Fernndez-Lpez, S., Melndez, G., 1994. The Middle-Upper Jurassic oolitic ironstone level in the Iberian Range (Spain). Eustatic implications. In: Cariou, E., Hantzpergue, P. (Eds.), 3e Symposium International de Stratigraphie du Jurassique. Poitiers, France, 2229 septembre 1991, 2e dition. Geobios MS 17, pp. 549561. Aurell, M., Melndez, G., Olriz, F. (coords), Bdenas, B., Caracuel, J., GarcaRamos, J.C., Goy, A., Linares, A., Quesada, S., Robles, S., RodrguezTovar, F.J., Rosales, I., Sandoval, J., Suarez de Centi, C., Tavera, J.M., Valenzuela, M., 2002. Jurassic. In: Gibbons, W., Moreno, M.T. (Eds.), The Geology of Spain. The Geological Society London, pp. 213253. Aurell, M., Robles, S., Rosales, I., Quesada, S., Melndez, A., Bdenas, B., Garca-Ramos, J.C., 2003. Transgressive/regressive cycles and Jurassic palaeogeography of northeast Iberia. Sedimentary Geology 162, 239271. Barn, A., Forns, J.J., Pomar, L., 2004. Eivissa: Sntesis estratigrca. In: Vera, J.A. (Ed.), Geologa de Espaa. Sociedad Geolgica de Espaa; Instituto Geolgico y Minero de Espaa, pp. 459460. Boucot, A.J., 1982. Ecostratigraphic framework for the Lower Devonian of the North American Appohimchi Subprovince. Neues Jahrbuch fr Geologie und Palontologie. Abhandlingen 163, 2181. Brett, C.E., 1995. Sequence stratigraphy, biostratigraphy, and taphonomy in shallow marine environments. Palaios 10, 597616. Brett, C.E., Baird, G.C., 1986. Comparative taphonomy: a key to paleoenvironmental interpretation based on fossil preservation. Palaios 1, 207227. Brett, C.E., Baird, G.C., 1993. Taphonomic approaches to temporal resolution in stratigraphy: examples from Paleozoic marine mudrocks. In: Kidwell, S.M., Behrensmeyer, A.K. (Eds.), Taphonomic approaches to time resolution in fossil assemblages. Short Courses in Paleontology, Knoxville, Tennessee 6, pp. 250274. Bulard, P.F., Feuill, P., Floquet, M., 1979. La limite Jurassique moyenJurassique suprieur dans la Sierra dAralar (Pyrnes basques espagnoles). Cuadernos de Geologa Ibrica 10, 179196. Collin, P.Y., Courville, P., 2000. Paloenvironnements et biostratigraphie dune srie oxfordienne non condense de rfrence (Saint-Blin-Smilly, HauteMarne). Gologie de la France 2000 (1), 5963. Courville, P., Collin, P.Y., 1997. La srie du Callovien et de lOxfordien de Veuxhaulles (Chtillonnais, Cte dOr) : problmes de datation, de gomtrie et de paloenvironnements dans une srie condense . Bulletin des Sciences de Bourgogne 49, 2943. Courville, P., Collin, P.Y., 2002. Taphonomic sequences - A new tool for sequence stratigraphy. Geology 30, 511514. Dromart, G., 1989. Deposition of Upper Jurassic ne-grained limestones in the western Subalpine Basin, France. Palaeogeography, Palaeoclimatology, Palaeoecology 69, 2343. Fernndez-Lpez, S., 1997. Ammonites, ciclos tafonmicos y ciclos estratigrcos en plataformas epicontinentales carbonticas. Revista Espaola de Paleontologa 12, 151174. Fernndez-Lpez, S., 2000. Ammonite taphocycles in carbonate epicontinental platforms. Georesearch Forum 6, 293300. Fernndez-Lpez, S., Melndez, G., 1994. Abrasion surfaces on internal moulds of ammonites as palaeobathymetric indicators. Palaeogeography, Palaeoclimatology, Palaeoecology 110, 2942. Fernndez-Lpez, S., Melndez, G., 2004. Fossilization of ammonites and sedimentary events in deep environments of carbonate platform (highest Middle to lowest Upper Oxfordian, Iberian Range, Spain). Rivista Italiana di Paleontologia e Stratigraa 110, 219230. Foucault, A., 1971. tude gologique des environs des sources du Guadalquivir (Provinces de Jan et de Grenade, Espagne mridionale). Thse de lUniversit de Paris. Fritz, G., 1958. Schammstotzen, Tuberolithe und Schuttbreccien im Weissen Jura der Schwabischen Alb. Arbeiten aus dem Geologisch-Palontologischen Institut der Technischen Hochschule Stuttgart. Neue Folge 13, 1118. Gaillard, C., 1983. Les biohermes spongiaires et leur environnement dans lOxfordien du Jura meridional. Documents des Laboratoires de Gologie de Lyon 90, 1515. Garca-Hernndez, M., Lpez-Garrido, A.C., 1988. The Prebetic platform during the Jurassic: a sedimentary evolution upon a distensive margin. In: Rocha, R.B., Soares, A.F. (Eds.), 2nd International Symposium on Jurassic Stratigraphy, Lisboa, pp. 10171030.

41

Gygi, R.A., 1969. Zur Stratigraphie der Oxford-Stufe (Oberes Jura-System) der Nordschweizund des sddeutschen Grenzgebietes. Beitrge zur geologischen Karte Schweiz 136, 1123. Gygi, R.A., 1981. Oolitic iron formations: marine or not marine? Eclogae geologicae Helvetiae 74, 233254. Gygi, R.A., Coe, A.L., Vail, P.R., 1998. Sequence stratigraphy of the Oxfordian and Kimmeridgian stages (Late Jurassic) in northern Switzerland. Mesozoic and Cenozoic Sequence Stratigraphy of European Basins. Society of Sedimentary Geology, Tulsa, Special Publication 60, 527544. Hartman, W.D., Wendt, J.W., Wiedenmayer, F., 1980. Living and fossil sponges, notes for a short course. Sedimenta 8, 1274. Huber, B., Mller, B., Luterbacher, H., 1987. Mikropalontologische Untersuchungen an der Callovien/Oxfordien-Grenze im Schweizer Jura und auf der Schwbischen Alb (vorluge Mitteilung). Eclogae geologicae Helvetiae 80, 449459. Jacquin, T., Dardeau, G., Durlet, C., de Graciansky, P.C., Hantzpergue, P., 1998. The North Sea Cycle: an overview of 2nd-Order Transgressive-Regressive Facies Cycles in Western Europe. Mesozoic and Cenozoic Sequence Stratigraphy of European Basins. Society of Sedimentary Geology, Tulsa, Special Publication 60, 445466. Kauffman, E.G., 1988. Concepts and methods of high resolution event stratigraphy. Annual Review of Earth and Planetary Science Letters 16, 605654. Kidwell, S.M., 1993. Pattens of time-averaging in the shallow marine fossil record. In: Kidwell, S.M., Behrensmeyer, A.K. (Eds.), Taphonomic approaches to time resolution in fossil assemblages. Short Courses in Paleontology 6, pp. 275300. Kidwell, S.M., Bosence, D.W.J., 1991. Taphonomy and time-averaging of marine shelly faunas. In: Allison, P.A., Briggs, D.E.G. (Eds.), Taphonomy. Releasing the Data Locked in the Fossil Record. Plenum, New York, pp. 115209. Kowalewski, M., Flessa, K.W., Aggen, J.A., 1994. Taphofacies analysis of recent shelly cheniers (Beach Ridges), northeastern Baja California, Mexico. Facies 31, 209242. Leinfelder, R.R., 1993. A sequence stratigraphic approach to the Upper Jurassic mixed carbonate-siliciclastic succession of the central Lusitanian Basin, Portugal. Prol 5, 1191240. Leinfelder, R.R., Wilson, R.C., 1998. Third-Order sequences in an Upper Jurassic rift-related Second-Order sequence, Central Lusitanian Basin, Portugal. Mesozoic and Cenozoic Sequence Stratigraphy of European Basins. Society of Sedimentary Geology, Tulsa, Special Publication 60, 507525. Lorin, S., Courville, P., Collin, P.Y., Thierry, J., Tort, A., 2004. Modalits de rinstallation dune plate-forme carbonate aprs une crise sdimentaire : exemple de la limite Oxfordien moyen Oxfordien suprieur dans le SudEst du Bassin de Paris. Bulletin de la Socit gologique de France 175, 289302. Marques, B., Olriz, F., Rodrguez-Tovar, F.J., 1991. Interactions between tectonics and Eustasy during the Upper Jurassic and lowermost Cretaceous. Examples from the south of Iberia. Bulletin de la Socit gologique de France 162, 11091124. Martin, R.E., 1990. Ecostratigraphy datums and sequence stratigraphy: Application to the marine Quaternary. AAPG Bulletin-American Association of Petroleum Geologists 75, 630. Martn-Algarra, A., Vera, J.A., 2004. Evolucin de la Cordillera Btica. In: Vera, J.A. (Ed.), Geologa de Espaa. Sociedad Geolgica de Espaa, Instituto Geolgico y Minero de Espaa, pp. 437444. Martinsson, A., 1973. Ecostratigraphy. Lethaia 6, 441443. Miller, III, W. (Ed.), 1990. Paleocommunity Temporal dynamics: The longterm development of multispecies assemblies. The Paleontological Society. Special Publication 5. Norris, M.S., Hallam, A., 1995. Facies variation across the Middle-Upper Jurassic boundary in Western Europe and the relationship to sea level changes. Palaeogeography, Palaeoclimatology, Palaeoecology 116, 189245. Olriz, F., Caracuel, J., Rodrguez-Tovar, F.J., 1995. Using ecostratigraphic trends in sequence stratigraphy. In: Haq, B.U. (Ed.), Sequence stratigraphy and depositional response to eustatic, tectonic and climatic forcing. Kluwer Academic Publisher, Netherlands, pp. 5985. Olriz, F., Caracuel, J., Ruiz-Heras, J.J., Rodrguez-Tovar, F.J., Marques, B., 1996. Ecostratigraphic approaches, sequence stratigraphy proposals and

42

F. Olriz et al. / Geobios 41 (2008) 3142 Rabe, B.D., Cisne, J.L., 1980. Chronostratigraphy accuracy of Ordovician ecostratigraphic correlation. Lethaia 13, 109118. Ramajo, J., Aurell, M., 1997. Anlisis sedimentolgico de las discontinuidades y depsitos del Calloviense superior-Oxfordiense medio en la Cordillera Ibrica Noroccidental. Cuadernos de Geologa Ibrica 22, 213236. Ramajo, J., Aurell, M., Cepra, J., 2002. Anlisis de facies de la Capa de Oolitos ferruginosos de Arroyofro en la Sierra de Arcos (Jursico, Cordillera Ibrica septentrional). Journal of Iberian Geology 28, 4564. Ramon, X., Aurell, M., Melndez, G., 1992. Stratigraphy and associated unconformities in the Middle to Upper Jurassic on the south central Pyrenees, Spain. II Congreso Geolgico de Espaa, Simposios 2, 161167. Reolid, M., 2005. Dinmica eco-sedimentaria durante el Oxfordiense medioKimmeridgiense temprano en la Zona Prebtica: Interpretacin ecoestratigrca y secuencial. PhD Thesis Universidad de Granada. Rodrguez-Tovar, F.J., 1993. Evolucin sedimentaria y ecoestratigrca en plataformas epicontinentales del margen Sudibrico durante el Kimmeridgiense inferior. PhD Thesis Universidad de Granada. Rollins, H.B., West, R.R., Busch, R.M., 1990. Hierarchical Genetic Stratigraphy and Marine Paleoecology. In: Miller, III, W. (Ed.), Paleocommunity Temporal dynamics: The long-term development of multispecies assemblies The Paleontological Society. Special Publication 5, pp. 273300. Sageman, B.B., Kauffman, E.G., Harries, P.J., Elder, W.P., 1997. Cenomanian/ Turonian Bioevents and Ecostratigraphy in the Western Interior Basin. In: Contrasting Scales of Local, Regional, and Global Events, Regional, Global, Events, In: Brett, C.E., Baird, G.C. (Eds.), Paleontological Events - Stratigraphic, Ecological, and Evolutionary Implications. Columbia University Press, New York, pp. 520570. Scouaire, Q., Marchand, D., Bonnot, A., Courville, P., Raffray, M., Huault, V., 1997. Le contact Callovien-Oxfordien dans les environs de Chaignay : nouvelles donnes stratigraphiques et palontologiques. Bulletin des Sciences de Bourgogne 49, 4563. Smith, A.M., Nelson, C.S., 2003. Effects of early sea-oor processes on the taphonomy of temperate shelf skeletal carbonate deposits. Earth-Science Reviews 63, 131. Sokolov, B., Kaljo, D., 1986. Theory and practice of ecostratigraphy (excerpts). International Geology Review 30, 12. Speyer, S.E., Brett, C.E., 1986. Trilobite taphonomy and Middle Devonian taphofacies. Palaios 1, 312327. Speyer, S.E., Brett, C.E., 1988. Taphofacies models for epeiric sea environments: middle Paleozoic examples. Palaeogeography, Palaeoclimatology, Palaeoecology 63, 225262. Vera, J.A. (coord.), Arias, C., Garca-Hernndez, M., Lpez-Garrido, A.C., Martn-Algarra, A., Martn-Chivelet, J., Molina, J.M., Rivas, P., Ruz-Ortiz, P., Sanz de Galdeano, C., Vilas, L., 2004. Las Zonas Externas Bticas y el Paleomargen Sudibrico. In: Vera, J.A. (Ed.), Geologa de Espaa. Sociedad Geolgica de Espaa, Instituto Geolgico y Minero de Espaa, pp. 354361. Ziegler, M.A., 1962. Beitrge zur Kenntnis des unteren Malm im zentralen Jura. Mitteilungen aus dem Geologischen Institut der Eidgen. Technischen Hochschule und der Universitat Zurich 82, 151. Zuschin, M., Stachowisch, M., Stanton, R.J., 2003. Patterns and processes of shell fragmentation in modern and ancient marine environments. EarthScience Reviews 63, 3382.

block tectonics: examples from epioceanic swell areas in south and east Iberia. Palaeogeography, Palaeoclimatology, Palaeoecology 121, 273295. Olriz, F., Linares-Rodrguez, A., Goy, A., Sandoval, J., Caracuel, J., Rodrguez-Tovar, F.J., Tavera, J.M., 2002a. The Betic Cordillera and Balearic islands. In: Gibbons, W., Moreno, T. (Eds.), The Geology of Spain, 11 Jurassic. The Geological Society, London, pp. 235253. Olriz, F., Reolid, M., Rodrguez-Tovar, F.J., 2002b. Fossil assemblages, lithofacies and taphofacies for interpreting depositional dynamics in epicontinental Oxfordian (Prebetic Zone, Betic Cordillera, southern Spain). Palaeogeography, Palaeoclimatology, Palaeoecology 185, 5375. Olriz, F., Reolid, M., Rodrguez-Tovar, F.J., 2002c. Taphonomic features in Upper Oxfordian ammonite assemblages (Bifurcatus Zone) from the Navalperal section (Internal Prebetic, Betic Cordillera). In: de Renzi, M., Pardo, M.V., Belinchn, M., Pealver, E., Montoya, P., Mrquez-Aliaga, A. (Eds.), Currents topics on taphonomy and fossilization. Col-lecci Encontres 5, Valencia, pp. 215222. Olriz, F., Reolid, M., Rodrguez-Tovar, F.J., 2003a. A Late Jurassic carbonate ramp colonized by sponges and benthic microbial communities (External Prebetic, Southern Spain). Palaios 18, 528545. Olriz, F., Reolid, M., Rodrguez-Tovar, F.J., 2004a. Ferruginous oolitic limestone from the Prebetic Zone (South Iberian Margin): pulse of pelagic sedimentation on epicontinental platform during the Late Jurassic. Recueil des Rsums du 2me Colloque sur le Jurassique Marocain, 126127. Olriz, F., Reolid, M., Rodrguez-Tovar, F.J., 2004b. Taphonomy of ammonite assemblages from the Middle-Upper Oxfordian (Transversarium?-Bifurcatus Zones) in the Internal Prebetic (Betic Cordillera, southern Spain): Taphonic populations and taphofacies to support ecostratigraphic interpretations. Rivista Italiana di Paleontologia e Stratigraa 110, 239 248. Olriz, F., Rodrguez-Tovar, F.J., Caracuel, J.E., 1994. Faunal assemblages, ecostratigraphy and high resolution sequence stratigraphy. In: Johnson, S.D. (Ed.), High Resolution Sequence Stratigraphy: Innovation and Application. University of Liverpool, Oxford, pp. 198203. Olriz, F., Rodrguez-Tovar, F.J., Marques, B., Caracuel, J.E., 1993. Ecostratigraphy and sequence stratigraphy in high frequency sea level uctuations: examples from Jurassic macro-invertebrate assemblages. Palaeogeography, Palaeoclimatology, Palaeoecology 101, 131145. Olriz, F., Villaseor, A.B., Gonzlez-Arreola, C., 2003b. Major lithostratigraphic units in land-outcrops of north-central Mexico and the subsurface along the northern rim of the Gulf of Mexico Basin (Upper Jurassiclowermost Cretaceous): a proposal for correlation of tectono-eustatic sequences. Journal of South American Earth Sciences 16, 119142. Parsons, K.M., Brett, C.E., 1991. Taphonomic processes and biases in modern marine environments: an actualistic perspective on fossil assemblage preservation. In: Donovan, S.K. (Ed.), The Process of Fossilization. Columbia University Press, New York, pp. 2265. Pendas, F., 1971. Denicin morfolgica de los embalses subterrneos del alto sureste espaol. I Congreso Hispano-Luso-Americano de Geologa Econmica Seccin Hidrogeologa II, 529550. Pittet, B., Van Buchem, F.S.P., Hillgrtner, H., Razin, P., Grtsch, J., Droste, H., 2002. Ecological succession, palaeoenvironmental change, and depositional sequences of the Barremian-Aptian shallow-water carbonates in northern Oman. Sedimentology 49, 555581.