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Cretaceous Research (1999) 20, 663–683Article No. cres.1999.0176, available online at http://www.idealibrary.com on

Aptian bio-events—an integratedbiostratigraphic analysis of the AlmadichFormation, Inner Prebetic Domain, SE Spain

*Roque Aguado, †Jose Manuel Castro, ‡Miguel Company and†Gines Alfonso de Gea

*Departamento de Geologıa, Universidad de Jaen, Escuela Universitaria Politecnica de Linares,Alfonso X El Sabio 28, 23700 Linares, Spain†Departamento de Geologıa, Universidad de Jaen, Facultad de Ciencias Experimentales, Campus Universitario LasLagunillas, 23071 Jaen, Spain‡Departamento de Estratigrafıa y Paleontologıa, Universidad de Granada, Facultad de Ciencias, 18002 Granada,Spain

Revised manuscript accepted 12 July 1999

The Upper Barremian to Aptian Almadich Formation (Inner Prebetic Domain of the Betic Cordillera) is composed ofhemipelagic sediments deposited on a distal carbonate ramp in the southern Iberian Palaeomargin. Within this facies we havefound a thick interval of blue to black shales and marls that is interpreted as deposited under oxygen-depleted conditions. Wethink that this interval, dated as early Aptian, represents the local record of Ocean Anoxic Event 1a. The integratedbiostratigraphic analysis of a section in the Almadich Formation, by means of planktonic foraminifera, calcareousnannofossils and ammonites, enables us to recognize most of the biostratigraphic units based on these three fossil groups andto correlate between them. The Sartousiana, Sarasini, Weissi, Deshayesi and Furcata (ammonite) Zones were identified forthe Upper Barremian–Lower Aptian interval. By means of calcareous nannofossil biostratigraphy the Micrantholithushoschulzii, Hayesites irregularis and Rhagodiscus angustus Zones, plus several additional biohorizons, were identified. Aquantitative study performed on a set of 27 Lower Aptian samples has enabled the precise identification of the ‘nannoconidcrisis’, the lower limit of which clearly precedes the main anoxic event, and its correlation with other bioevents. Planktonicforaminifera occur consistently throughout the Lower to Upper Aptian of the Cau section and are moderately well preserved.This fact allows us to use the most recent taxonomic framework, based on wall texture, to identify the Blowiella blowi,Schackoina cabri, Globigerinelloides ferreolensis, Globigerinelloides algerianus, Hedbergella trocoidea and Ticinella bejaouaensis Zones.Coincident with the anoxic episode, the planktonic foraminiferal assemblages are composed of a significant number of formswith elongated chambers and/or tubulospines assigned to the genera Claviblowiella, Lilliputianella, Leupoldina and Schackoina.Most of the planktonic foraminiferal and nannofossil taxa are illustrated. � 1999 Academic Press

K W: Barremian; Aptian; biostratigraphic events; calcareous nannofossils; planktonic foraminifera; ammonites;anoxic events; Tethys; Betic Cordillera; southeastern Spain.

1. Introduction

The Lower Aptian marine sediments globally record amajor turnover in marine fauna and flora. This ismainly reflected in changes in the ammonite,belemnite, dinoflagellate, radiolarian and benthicforaminiferal taxa, radiation and diversification ofplanktonic foraminifera and silicoflagellates, anddiversification and later crisis within the calcareousnannofossils (Caron, 1985; Williams & Bujak,1985; Coccioni et al., 1992; McCartney, 1993; Erba,1994; O’Dogherty, 1994; Boudagher-Fadel, 1996;Boudagher-Fadel et al., 1996, 1997a, 1998; Aguado

0195–6671/99/020663+21 $30.00/0

et al., 1997; Bischoff & Mutterlose, 1998; Bown et al.,1998; Cecca, 1998; Mutterlose, 1998, among others).These changes coincided with other major palaeogeo-graphic and palaeoceanographic events, such as thedevelopment of an oceanic anoxic event (Schlanger &Jenkyns, 1976; Arthur et al., 1990), widespreaddrowning of shallow carbonate platforms (Schlager,1989), oceanic volcanic eruptions in the Pacific (the‘superplume episode’ of Larson, 1991a), a global risein sea level (Haq et al., 1988; Hallam, 1992) andthe onset of the Cretaceous greenhouse conditions(Larson, 1991b).

� 1999 Academic Press

664 R. Aguado et al.

The complexity of the temporal and causal relation-ships between biotic and geologic events during theEarly Aptian makes the establishment of an accuratebiostratigraphic framework for this time interval(Erba, 1994; Erba et al., 1996; Aguado et al., 1997;Cobianchi et al., 1997; Bischoff & Mutterlose, 1998)necessary in order to improve dating and correlationof the various recorded events.

The main goal of this paper is to present an inte-grated biostratigraphic analysis of the uppermostBarremian–Aptian interval, based on ammonites,planktonic foraminifera and calcareous nannofossils.With this aim, we have selected a section rich inplanktonic fauna and flora and well known in terms ofboth palaeogeographic setting and sequence strati-graphic analysis (Castro, 1998; Ruiz-Ortiz & Castro,1998).

2. Geologic and palaeogeographic setting

The studied section, located on the western slope ofCau hill, near Calpe (Alicante province, SE Spain)(Figure 1), geologically belongs to the PrebeticDomain of the Betic Cordillera. The Prebetic was avast epeiric carbonate platform, formed on a passivemargin attached to a Hercynian craton to the north(the Iberian Massif). This platform settled during theCretaceous on the northern margin of the Tethys,within the Tethys–Atlantic seaway.

During this time, carbonate ramps developed onthe Prebetic Platform, which evolved from shallowUrgonian environments in the north to distal hemi-pelagic environments to the south, with a very gentleslope. The sediments of the Cau section weredeposited on this epeiric platform under hemipelagicconditions, but only a few tens of metres deep, andpalaeogeographically close to the depositional site ofshallow platform carbonates (Company, 1987;Castro, 1998; Ruiz-Ortiz & Castro, 1998). In thisparticular palaeogeographic setting, hemipelagic sedi-mentation was more sensitive to relative sea-levelfluctuations. The sedimentary evolution was alsoclosely related to that of more proximal environments,well documented in the Alicante and adjacent areas(Vilas et al., 1993; Castro & Ruiz-Ortiz, 1995; Castro,1998; Ruiz-Ortiz & Castro, 1998). Tectonics played akey role on this platform, leading to substantial lateralchanges in the subsidence rate. In this context, thearea of deposition of the section studied correspondsto a strongly subsiding zone, bordered towards moredistal positions by highs, namely the Sierra Heladaand Cabezon de Oro (Granier, 1987; Ruiz-Ortiz &Castro, 1998).

Several third-order sequences have been deducedfrom the record of Aptian sediments in the Prebetic(Vilas et al., 1993; Ruiz-Ortiz & Castro, 1998), withits drastic vertical and lateral facies changes which

Figure 1. Geographical and geological location of the section studied.

An analysis of the Almadich Formation 665

represent rapid lateral shifts in depositional environ-ments. The Lower Aptian beds studied weredeposited in a general transgressive context and rep-resent the lower part of a transgressive-regressivesecond order stratigraphic sequence that comprisesthe entire Aptian (Castro, 1998; Castro & Ruiz-Ortiz,1998). This transgressive trend has been registeredworldwide (Haq et al., 1988; Hallam, 1992). It led tothe submergence of broad continental areas and tosignificant processes of backstepping and drowning ofcarbonate platforms, probably in relation to tectonicpulses (Vilas et al., 1993; Funk et al., 1993; Castro &Ruiz-Ortiz, 1995).

The Cau section was studied by Gignoux & Fallot(1926, in Darder, 1945), and later by Darder (1945)and Rıos et al. (1961), all of whom focused mainly onthe presence of ammonites. More recent studies in theregion have concerned the lithostratigraphy, bio-stratigraphy, sedimentology and sequence stratigraphy(Company et al., 1982; Castro & Ruiz-Ortiz, 1994,1995; Castro, 1996, 1998; Ruiz-Ortiz & Castro,1998).

3. Lithostratigraphy

In the Cau area, a relatively complete sequence of lateBarremian–Aptian age crops out, with a thickness ofroughly 197 m (Figure 2). It consists predominantlyof rhythmic alternations of marls and marly lime-stones with ammonites. These facies belong to theAlmadich Formation (Castro, 1998) which, in thisoutcrop, overlies the marls and marly limestonesof the Los Villares Formation (Ruiz-Ortiz, 1980;Aguado et al., 1996) and underlies the ochre bioclasticcalcarenites of the Seguilı Formation (Castro, 1998).Towards the north, the lower and upper parts of theAlmadich Fm shift, by lateral changes of facies, toUrgonian limestones of the Llopis Formation and theSeguilı Formation, respectively (Castro, 1998). Inlithostratigraphic terms, the Almadich Fm can bedivided into three members.

Lower member

With a visible thickness of about 55 m (the base of theunit does not crop out owing to faulting), this memberis characterized by a rhythmic alternation of marlylimestones and light grey marls containing ammonitesand nannofossils in abundance. Rare to commonplanktonic foraminifera are present only in the upperpart. In general, 1-m-thick cycles are evident withmarly limestones predominating over the marls exceptthrough an 18 m interval in the upper part, in whichmarls predominate. In places, slightly ferruginous

surfaces are visible at the top of these cycles. The bedin which sample 11c is located (Figure 2) shows aferruginous and bioturbated surface (hardground)towards the top, and we have detected, in associationwith this surface, a stratigraphic discontinuity whichcoincides with the Barremian/Aptian boundary (seebelow). This surface corresponds to the basal bound-ary of the Aptian second-order stratigraphic sequence(Ruiz-Ortiz & Castro, 1998). Notable also in theupper part of this member is a bed (14.6 in Figure 2)comprising 35 cm of black marls with pyrite nodules.

Middle member

The middle member is approximately 27 m thick andmainly composed of black to dark blue shales andmarly sediments. In detail, it has a cyclic organization,with decimetre-thick cycles comprising a lower blacklutitic division that evolves gradually upwards to grey,more marly and silty sediments. The top of each cycleis marked by a bed of light grey marlstones and ochreto yellow marls 1–2 cm thick which are sometimescapped by a slightly reddish surface. The dark col-oured sediments are interpreted as having been de-posited under oxygen-depleted conditions and shouldbe characterized by their high organic carbon content(Coccioni et al., 1989; Breheret, 1994; Baudin et al.,1998). Owing to both sedimentologic (very high sedi-mentation rates) and biostratigraphic considerations(see below), we conclude that this middle memberas a whole should be correlated with other lowerAptian black shale deposits in central and northernItaly (‘Livello Selli’), southeastern France (‘NiveauGoguel’) and, probably, in northwestern Germanyand the North Sea area (‘Fischschiefer’). If this iscorrect, then the member, hereafter referred to in thetext as reflecting the main anoxic event, should repre-sent the local record of the Ocean Anoxic Event(OAE) 1a (Arthur et al., 1990; Larson et al., 1993;Bralower et al., 1994). The differences in thicknessbetween it and the ‘Livello Selli’ and the ‘NiveauGoguel’ can be explained by a different palaeogeo-graphic context, the Cau section corresponding to adistal ramp with a very high subsidence rate andterrigenous input from the continent, whereas the‘Livello Selli’ and ‘Niveau Goguel’ accumulated inpelagic conditions.

Upper member

This member is 115 m thick and composed of alter-nating marly limestones and grey marls (white onweathered surfaces) in 1.5-m-thick cycles. These beds


Figure 2. Lithology and integrated biostratigraphy of the Cau section. The figure shows lithologic log and scale, location ofsamples, lithostratigraphic units and stages. The most important calcareous nannofossil and planktonic foraminiferalevents are indicated with respect to sample numbers. Nannofossil events are marked with an asterisk (*). Separatecolumns list calcareous nannofossil, planktonic foraminiferal and ammonite zones. The graph, lower right-hand side,shows the results of a semiquantitative analysis of 27 samples. The proportions of Watznaueria barnesae, Nannoconus spp.and narrow canal nannoconids are shown as percentages. For all values in the graph, the average percentage error for a99.9% confidence index (Dennison & Hay, 1967) is 2.0% (maximum=7.2%, minimum=0.5%).


contain very few ammonites and common to abun-dant planktonic foraminifera and nannofossils. Theinterval begins with beds of marly limestones havingthin intercalated layers of marls. Some beds of marlylimestones have weakly ferruginous surfaces at theirtops. In the uppermost part of the member, the marlylimestone layers become slightly sandy. The lithologyof this member, by comparison to the middle mem-ber, indicates a change in sedimentary environment,in particular a return to more oxygenated conditions.

4. Integrated biostratigraphy

We have analysed the stratigraphic distribution ofthree different fossil groups (calcareous nannofossils,planktonic foraminifera and ammonites) in the Causection. For the study of the nannofossils and plank-tonic foraminifera, the use of the same samples allowsa direct correlation between both groups. As for theammonites, most of the samples were also those usedto study the foraminifera and nannofossils; whendifferent, they were precisely located in the section. Inthis way, we were able to establish a direct correlationbetween the biostratigraphic scales based on the threefossil groups for part of the Upper Barremian andLower Aptian. The stratigraphic distribution of themost significant taxa of these groups is shown in arange chart (Figure 3).

4.1. Nannofossil biostratigraphy

Some 45 samples from the Almadich Fm werestudied. These cover the entire formation at irregularintervals, as shown in Figure 2. Only the lowest fivebelong to the late Barremian; the rest are of Aptianage. Smear slides were prepared from untreated rawmaterial in order to preserve as faithfully as possiblethe original composition of the nannofossil assem-blages. A quantitative study was performed on a groupof 27 samples of late Barremian–early Aptian age inorder to establish the relative proportions of the dif-ferent taxa and to locate the ‘nannoconid crisis’ eventas reliably as possible. A minimum of 200 specimensalong random traverses were counted in each smearslide, although this number was frequently surpassedto reach much higher values (occasionally up to 1500specimens). For each sample, the proportions of thedifferent taxa were expressed as percentages. Figure 2shows partial results from this study, with respect tothe total percentages of Nannoconus spp., the narrowcanal nannoconids and Watznaueria barnesae. For therest of the samples, we performed a routine biostrati-graphic study on smear slides, investigating at least200 fields at 1250�.

In general, the diversity of the assemblages provedto be high, with a total of 55 identified taxa (seeAppendix). The assemblages (Figure 4) are domi-nated by the genera Watznaueria and Nannoconus,with other species, such as Biscutum ellipticum,Zeugrhabdotus noeliae, Lithraphidites carniolensis andRhagodiscus asper, being common. Other less abun-dant but characteristic forms include Assipetra infra-cretacea, A. terebrodentaria, Cyclagelosphaera margerelii,Diazomatolithus lehmanii, Flabellites oblongus, Rhagodis-cus gallagheri and, for the Aptian, Braarudosphaeraafricana, Eprolithus floralis, Hayesites irregularis andRhagodiscus angustus. According to their general com-position, these assemblages were clearly Tethyanfor the Barremian, with a progressive increase incosmopolitan species in the Aptian (Mutterlose,1991, 1992a, b, 1996; Aguado, 1993a, b; Erba, 1994;Aguado et al., 1997; Bischoff & Mutterlose, 1998;Bown et al., 1998; Mutterlose & Bockel, 1998). Sometaxa, such as Crucibiscutum hayi and C. salebrosum,considered typically boreal (Mutterlose, 1992a, b),were also recorded.

This biostratigraphic analysis enabled the identifi-cation of the Micrantholithus hoschulzii, Hayesitesirregularis and Rhagodiscus angustus Zones (Figure 2)according to the zonation of Applegate & Bergen(1988).

Calcareous nannofossil events. The lowermost samplesin the section (10–11c, Figure 2) should be assignedto the M. hoschulzii Zone (Thierstein, 1971, 1973),defined as the interval between the last occurrence(LO) of Calcicalathina oblongata and the first occur-rence (FO) of Hayesites irregularis. Samples 10 and10b (Figure 2) contain Crucibiscutum salebrosum andsamples 11, 11b and 11c contain Rhagodiscus gallagh-eri (Figure 4.28) and Flabellites oblongus (Figure 4.19).The FO of this latter species has been shown to occurin the uppermost part of the M. hoschulzii Zone. Itcorrelates with the lower part of the Sarasini ammo-nite Zone (Aguado et al., 1995; Erba et al., 1996;Aguado et al., 1997), indicating a latest Barremianage. In some sections from the pelagic SubbeticDomain, the FO of R. gallagheri (=Rhagodiscus sp. cf.R. angustus in Aguado et al., 1997) is registered belowthe FO of F. oblongus. In the Cau section, the FOs ofboth taxa probably fall within the small covered inter-val of about 8 m near the base (Figure 2). Nanno-conids are abundant within the interval of the sectioncorresponding to this zone, with abundances of about35–44%, the plexus being dominated by the narrowcanal form Nannoconus steinmannii (18–43%).

Originally, Thierstein (1971, 1973) used the LO ofNannoconus colomii (=Nannoconus steinmannii in this

F


igure 3. Range chart showing the stratigraphic distribution of selected (most age-diagnostic) taxa for ammonites,calcareous nannofossils and planktonic foraminifera in the Cau Section. For calcareous nannofossils, total abundance iscoded as follows: A=abundant (>40 specimens in 10 fields of view); C=common (10–40 specimens in 10 fields of view);R=rare (<10 specimens in 10 fields of view). Species abundance is coded as follows: V=very abundant (>30%);A=abundant (15–30%); C=common (7–15%); F=few (2–7%); R=rare (<2%). For samples 23a–24e, speciesabundance is only an estimation, as no quantitative analysis was performed for this interval. The shaded boxes inAssipetra terebrodentaria indicate samples in which specimens >10 �m were found. Total abundance for planktonicforaminifera is an estimation. For all taxa, preservation is coded as P=poor; M=moderate or G=good.


study) and/or the FOs of Chiastozygus litterarius andRucinolithus irregularis to define the lower limit of hisC. litterarius Zone. It has been shown however, that

these three events are not coeval (Noel, 1980;Covington & Wise, 1987; Aguado et al., 1988, 1992a,1997; Applegate & Bergen, 1988; Erba, 1988, 1989,

Figure 4. 1–3, Assipetra terebrodentaria (large specimens: 1, sample 15.4; 2, sample 15.3; 3, sample 14.5). 4, Manivitellapemmatoidea, sample 15.7 (0� to crossed nicols). 5–7, Nannoconus steinmannii, sample 10b. 8, Zeugrhabdotus embergeri,sample 10b (10� to crossed nicols). 9, Assipetra terebrodentaria, sample 15.5 (normal-sized specimen). 10, Zeugrhabdotusnoeliae, sample 14.5 (30� to crossed nicols). 11, 12, Diazomatolithus lehmanii (28, sample 14.5; 30, sample 15.7). 13,Lithraphidites carniolensis, sample 14.5 (45� to crossed nicols. 14, Nannoconus vocontiensis, sample 13b. 15, Nannoconustruittii, sample 15.7. 16, Percivalia fenestrata, sample 10 (0� to crossed nicols). 17, Rhagodiscus asper, sample 14.9 (0� tocrossed nicols). 18, Retecapsa surirella, sample 14.5 (0� to crossed nicols). 19, Flabellites oblongus, sample 14.5 (30� tocrossed nicols). 20, Assipetra infracretacea, sample 15.7. 21–24, Hayesites irregularis (19, 23, sample 15.7; 20, 21, sample14.5; 22, sample 14.9). 25, Biscutum ellipticum, sample 15.6 (45� to crossed nicols). 26, Braarudosphaera africana, sample15.5. 27, Haqius circumradiatus, sample 14.9. 28, Rhagodiscus gallagheri, sample 15.6 (45� to crossed nicols). 29,Discorhabdus ignotus, sample 15.7. 30, Palaeomicula maltica, sample 10b. 31, Cyclagelosphaera margerelii, sample 15.7. 32,Helenea chiastia, sample 10b (0� to crossed nicols). 33, Chiastozygus sp. cf. C. litterarius, sample 12a (45� to crossednicols). 34, Staurolithites mitcheneri, sample 13b (45� to crossed nicols). 35, Eprolithus floralis, sample 23f. 36, Rhagodiscusangustus, sample 23e (0� to crossed nicols). All figures c. �3500.


1994; Aguado 1993b; Cobianchi et al., 1997). Forthis reason, we prefer to use the Hayesites irregularisnannofossil Zone, which is defined as the intervalbetween the FO of this species and the FO of Epro-lithus floralis (Thierstein, 1971; Manivit et al., 1977;Applegate & Bergen, 1988; Aguado, 1993b; Aguadoet al., 1997). H. irregularis (Figure 4.21–24) is firstregistered in sample 12a and is consistently presentthroughout the rest of the section. Samples 12a–21bare assigned to this zone. Just below sample 12a thereis a 2 m covered interval and, therefore, no samples.There is some evidence of a stratigraphic discontinuityat the base of this interval: sample 12a contains readilyidentifiable specimens of Nannoconus truittii (Figure4.15) and Braarudosphaera africana (Figure 4.26).According to Aguado et al. (1997), typical specimensof N. truittii are not present in the Tuarkyricus andlower part of Weissi ammonite Zones, and the FO ofB. africana is recorded in the upper part of theTuarkyricus ammonite Zone. On the other hand, thissample contains a significantly lower proportion ofnannoconids (about 28%) and very scarce (about0.5%) narrow canal nannoconids (i.e., N. steinmannii)in comparison with the underlying Upper Barremiansamples (Figure 2). This fact may indicate, accordingto Erba (1994), a relatively high position within the H.irregularis Zone. For the above reasons, we concludethat sample 12a should be correlated with the upperpart of the Weissi ammonite Zone. If this is correct,the covered interval should correspond to the com-plete Tuarkyricus and lowermost Weissi ammoniteZones, at least. The ‘nannoconid crisis’ (Erba, 1994)is recorded within the H. irregularis Zone, the begin-ning coinciding with sample 13.2 (Figure 2), and maybe correlated with the upper part of the Weissi or thelower part of the Deshayesi ammonite Zones. Nonarrow canal nannoconids have been found within orabove the interval corresponding to the ‘nannoconidcrisis’ (Figure 2). This interval is also characterized bythe presence of large (about 10 �m in diameter)specimens of Assipetra terebrodentaria (Figure 4.1–3)together with abundant (35–60%) Watznaueria bar-nesae (Figure 2). Figure 2 shows that the beginning ofthe ‘nannoconid crisis’ clearly preceded the mainanoxic event, which is marked by a net change in thelithologic record.

Thierstein (1971, 1973), defined the Rhagodiscusangustus Zone as the interval between the FOs of R.angustus and/or E. floralis and the FO of Predisco-sphaera columnata. It has been demonstrated that theFOs of R. angustus and E. floralis are not coeval in theTethyan realm (Applegate & Bergen, 1988; Aguadoet al., 1991, 1992a, b; Aguado, 1993b; Erba, 1994;Cobianchi et al., 1997). In the Cau section, the FO of

R. angustus is unclear, owing to the presence of rare tofew specimens of Rhagodiscus gallagheri and intermedi-ate forms between the two taxa. These very rareintermediate forms were found in samples 19.1 and19.5, slightly below the FO of E. floralis. Commontrue specimens of R. angustus (Figure 4.36) first occurin sample 23a, within the Globigerinelloides ferreolensisplanktonic foraminiferal Zone, postdating the FOof E. floralis. To avoid taxonomic misinterpretationsrelated to the R. gallagheri-R. angustus plexus, weprefer to use the FO of E. floralis (Figure 4.35) todefine the base of R. angustus Zone. This event isrecorded in sample 22 of the Cau section, enabling usto assign samples 22–24e to the R. angustus nanno-fossil Zone. As can be seen in Figure 2, the base of theR. angustus Zone falls within the Furcata ammoniteZone and correlates with the upper part of the Schack-oina cabri planktonic foraminiferal Zone, at the top ofthe Lower Aptian. Diazomatolithus lehmanii (Figure4.11, 12) is scarce to rare throughout the UpperBarremian and Lower Aptian of the Cau section. ItsLO was recorded from sample 23f, within the lowerpart of the R. angustus Zone. This correlates with theGlobigerinelloides algerianus planktonic foraminiferalZone, which is late Aptian in age according toCobianchi et al. (1997). The species Conusphaerarothii, commonly found throughout the UpperBarremian–Lower Aptian of the pelagic SubbeticDomain (Aguado et al., 1992a, 1997; Aguado, 1993b)was not found in the more neritic sediments of theCau section.

4.2. Planktonic foraminiferal biostratigraphy

About 60 samples from micritic, marly, marly-limestone and shale layers from the Almadich Fmwere collected for foraminiferal analysis. These areirregularly spaced throughout the Lower–UpperAptian reflecting outcrop conditions and lithology.Sediments near and within the dysoxic/anoxic facieswere most closely sampled. The majority of thesamples were also investigated for their calcareousnannofossil content. They were disaggregated andwashed through sieves, the residue being separatedinto three fractions (>200 �m, 100–200 �m and 50–100 �m). For each sample, the three residues wereinvestigated, but the richest planktonic foraminiferalassemblages were found in the 100–200 �m residues.In samples containing rare planktonic foraminifera,the complete residue was investigated, and at leastone quarter of residues comprising abundant well-preserved assemblages were examined.

Planktonic foraminiferal abundances in productivesamples vary from scarce with only rare specimens to


very abundant and diverse. Overall, abundanceincreases up to the mid part of the section (dysoxic/anoxic facies) and remains high for much of the rest,but decreases near the top. By contrast, in most LowerAptian sections of northern and central Italy andsoutheastern France, the presumed equivalent anoxicfacies (‘Livello Selli’ and ‘Niveau Goguel’), are barrenor contain no age-diagnostic planktonic foraminiferalassemblages (Breheret & Delamette, 1989; Tornaghiet al., 1989; Coccioni et al., 1992). Preservation ishighly variable depending on lithology. The best pres-ervation normally correlates with marly lithotypeswhich are common in the mid to upper part of thesection and the poorest is associated with the limy andslightly silty lithotypes, which are more abundant inthe lower part (Figure 2). Although the walls of thespecimens have been modified by recrystallization, theoriginal shell texture can be identified in most taxa(Figures 5–10). This encouraged us to use the mostrecent taxonomic classification (Banner & Desai,1988; BouDagher-Fadel, 1995, 1996; BouDagher-Fadel et al., 1995, 1996, 1997a, 1998) mainly basedon shell texture and morphology. Owing to theuneven record of planktonic foraminifera, because ofthe control of the depositional environment on pres-ervation, only a qualitative analysis of the foramin-iferal associations was performed. A complete list ofthe taxa identified is provided in the Appendix.

Planktonic foraminiferal events. The lowermostsamples studied (13b–14.5 in Figure 2) are within theHayesites irregularis nannofossil Zone, immediatelyabove the horizon reflecting the ‘nannoconid crisis’event, and may correspond to the uppermost Weissior lowermost Deshayesi ammonite Zones. Theirplanktonic foraminiferal assemblages are poorly pre-served and characterized by forms with smooth micro-perforated walls having four or five (exceptionally six)rounded chambers showing no radial elongation. Allthese forms were assigned to the genera Blefuscuiana(B. aptiana, Figure 5.13–17; B. daminiae, Figure5.21–23; B. excelsa, Figure 6.20; B. infracretacea,Figure 5.18–20; B. laculata, Figure 5.24, 25; and B.occulta, Figure 6.3–6), Blowiella (B. blowi, Figure8.1–6), Gorbachikella (G. kugleri, Figure 5.1–4) andPraehedbergella (P. sigali, Figure 5.5–8; and P. tuschep-sensis, Figure 5.9–12). No forms with smooth, micro-perforated walls and radially elongated chambers (i.e.,Lilliputianella) were found in this interval. Scarcespecimens of Lilliputianella with minor chamber elon-gation (L. globulifera, Figure 7.3–10; and L. longorii,Figure 7.1–2) together with Blefuscuiana gorbachikae(Figure 6.1–2) and Blowiella duboisi (Figure 8.7–13)were found in samples 14.6–15. In the entire set

of these samples (13b–15), the assemblages aredominated by trochospiral non-elongated forms.

From sample 15.2 to 17.3, the assemblages containa high proportion of trochospiral Lilliputianella withgreat radial chamber elongation (L. bizonae, Figure7.22–24; L. kuhryi; and L. roblesae, Figure 7.13–21).Within this interval (sample 15.3), the FO of plani-spiral forms with smooth microperforated walls andgreat chamber elongation assigned to Claviblowiella(C. saundersi, Figure 9.1–8) was also recorded. Thischamber elongation has recently been regarded as anadaptive response of early planktonic foraminifera tooxygen-depleted sea water (BouDagher-Fadel et al.,1997c; Magniez-Jannin, 1998). The FO of Schackoina(S. cabri, Figure 9.16–20) is registered in sample 15.5,together with the first Leupoldina (L. protuberans, Fig-ure 9.9–15). Other species, which were first recordedwithin this interval, include Blefuscuiana convexa(Figure 6.19), Blowiella gottisi, B. maridalensis (Figure8.21–24), and B. moulladei (Figure 8.16–20). Figure 2shows that the FO of S. cabri, together with themaximum in the abundance of forms with elongatedchambers, appears to coincide with the lower part ofthe main anoxic event and lies within the upper Weissior lower Deshayesi ammonite Zone and H. irregularisnannofossil Zone. This latter fact is important becausein most of the published literature, the FO of S. cabriis located, or assumed to be located, immediatelyabove the top of the OAE 1a (Tornaghi et al., 1989;Coccioni et al., 1992; Erba & Quadrio, 1987; Erba,1988, 1994; Baudin et al., 1998; Weissert et al., 1998,among others). In the Cau section, no forms thatcould be assigned to Leupoldina or Schackoina werefound below the main anoxic event, althoughMagniez-Jannin et al. (1997) reported rare specimensof S. cabri below the ‘Niveau Goguel’ within theuppermost beds assigned to the Weissi Zone in sec-tions in the Vocontian Basin. An increase in thenumber of chambers in B. blowi may be observed fromthe top of this interval upwards. At higher strati-graphic levels, this leads to the FO of Globigerinelloidesferreolensis (BouDagher-Fadel et al., 1997b).

A third interval (samples 18.4–23d) is characterizedby a progressive decrease in forms with radially elon-gated chambers (Claviblowiella and Lilliputianella) anda significantly high proportion of planispiral formsbelonging first to Blowiella and then to Globigerinel-loides. Some specimens assigned to Claviblowiella sigali(Figure 8.25–30), six-to-seven-chambered planispiralforms reported as Blowiella sp. cf. B. blowi (Figure10.1–3), and six-to-seven-chambered trochospiralones reported as Blefuscuiana sp. cf. B. occulta (Figure6.7–12), were registered within this interval. The LO


Figure 5. 1–4, Gorbachikella kugleri, sample 16. 5–8, Praehedbergella sigali (5, sample 20b; 6, sample 17.3; 7, sampleCAU19.3; 8, sample 19.5). 9–12, Praehedbergella tuschepsensis (9, sample 18.4; 10, sample 14.1; 11, sample 19.5; 12,sample 17.3). 13–17, Blefuscuiana aptiana s. s. (13, 17, sample 19.5; 14, sample 21; 15, sample 17.1; 16, sample 18.4).18–20, Blefuscuiana infracretacea occidentalis (18, sample 17.3; 19, sample 19.3; 20, sample 19.5). 21–23, Blefuscuianadaminiae (21, 22, sample 14.5; 23, sample 17.1). 24, 25, Blefuscuiana laculata (24, sample 20.5; 25, sample 14.1). 26,27, Blefuscuiana speetonensis (26, sample 17.3; 27, sample 20.1). 28, Blefuscuiana sp. (last two chambers tend to beelongated), sample 19.5. 29, 30, Blefuscuiana sp. cf. B. rudis (29, sample 19.5; 30, sample 14.1). All figures c. �124.


Figure 6. 1, 2, Blefuscuiana gorbachikae (1, sample 19; 2, sample 21). 3–6, Blefuscuiana occulta, sample 21. 7–12, Blefuscuianasp. cf. B. occulta, sample 21 (more evolute, six to seven chambered morphologies). 13, 16–18, Hedbergella trocoidea (13,sample 23f; 16–18, sample 23e). 14, 15, Blefuscuiana sp. cf. B. praetrocoidea, sample 21. 19, Blefuscuiana convexa, sample14.9. 20, Blefuscuiana excelsa, sample 17.1. All figures c. �124.


Figure 7. 1, 2, Lilliputianella longorii (1, sample 16; 2, sample 21). 3–10, Lilliputianella globulifera (3, 6, sample 17.3; 4,sample 21; 5, sample 17.1; 7, 10, sample 19.3; 8, sample 20.1; 9, sample 20.3). 11, 12, Lilliputianella sp. cf. L. globulifera(forms intermediate between L. globulifera and L. roblesae: 11, sample 20.3; 12, sample 17.3). 13–21, Lilliputianellaroblesae (13, 14, 16, 17, 19, sample 16; 15, sample 17.3; 18, sample 17.1; 20, 21, sample 17.3). 22–24, Lilliputianellabizonae, sample 16. All figures c. �124.


Figure 8. 1–6, Blowiella blowi (1, sample 16; 2–5, sample 19.3; 6, sample 19.5 ). 7–13, Blowiella duboisi (7, sample 20.3;8–12, sample 17.3; 13, sample 19.3). 14, 15, Blowiella sp. cf. B. gottisi (14, sample 17.3; 15, sample 18b). 16–20,Blowiella moulladei (16, 18, 19, sample 17.3; 17, sample 18b; 20, sample 19.5). 21–24, Blowiella maridalensis (21, 22,sample 19.3; 23, sample 15.3; 24, sample 20b). 25–30, Claviblowiella sigali (25, 30, sample 18b; 26, sample 18.4; 27,sample 20.5; 28, sample 19.5; 29, sample 20.3). All figures c. �124.


Figure 9. 1–8, Claviblowiella saundersi (1–4, sample 16; 5, sample 17.3; 6, sample 19.5; 7, 8, sample 20.1). 9–15, Leupoldinaprotuberans (9, 10, 12, 13, sample 21; 11, sample 19b; 14, sample 17.1; 15, sample 19.5). 16–20, Schackoina cabri (16,18, sample 21; 17, sample 17.1; 19, sample 17.3; 20, sample 19.5). All figures c. �124.


Figure 10. 1–3, Blowiella sp. cf. B. blowi (six to seven chambered forms: 1, 2, sample 18b; 3, sample 23e). 4–12,Globigerinelloides ferreolensis (4, sample 23a; 5–11, sample 23e; 12, sample 23f). 13–15, ‘Biglobigerinella’ barri (13, sample23e; 14, sample 23c2; 15, sample 24). 16–19, Globigerinelloides algerianus (16, 17, sample 23f; 18, sample 24; 19, sample23d). 20–23, Hedbergella trocoidea (20, 21, 23, sample 23e; 22, sample 24). 24–27, Ticinella bejaouaensis, sample 24d. Allfigures c. �62.

of S. cabri, together with the FO of true Globigerinel-loides ferreolensis (Figure 10.4–12), with perforationcones in early chambers, and ‘Biglobigerinella’ barri(Figure 10.13–15) occured in sample 22a. Thesethree ‘events’ fall within the Furcata ammonite Zoneand near the base of the R. angustus nannofossil Zone(Figure 2). Finally, the FO of Globigerinelloides algeri-anus (Figure 10.16–19) was recorded from sample23d. From sample 22a upwards the average size of theplanktonic foraminiferal assemblages increases, owingto the appearance and great abundance of trueGlobigerinelloides and, at higher stratigraphic levels,Hedbergella and Ticinella. With the exception of rarespecimens of Blowiella sp. cf. B. blowi, no otherPraehedbergellidae were recorded from sample 23c2upwards (Figure 3).

From sample 23e to the top of the section (sample24e), the assemblages are dominated by trochospiralforms belonging to Hedbergella and finally Ticinella.Hedbergella trocoidea (Figures 6.16–18, 10.20–23) isfirst registered in sample 23e, being very abundantfrom this sample upwards. The LO of Globigerinel-loides algerianus was found in sample 24 and the FO ofTicinella bejaouaensis (Figure 10.24–27) in sample24b1. All of these ‘events’ occur within the R. angustusnannofossil Zone. No identifiable ammonites werefound in the Upper Aptian sediments of the Causection, thus preventing the establishment of corre-lations with respect to the planktonic foraminiferalbiostratigraphy. Planomalina cheniourensis, a character-istic Upper Aptian species (Caron, 1985; Aguadoet al., 1988, 1992a; Cobianchi et al., 1997),was not found. In the uppermost part of the Causection, the planktonic foraminiferal assemblages areimpoverished and poorly preserved.

The biostratigraphical framework adopted is basedon previous work by several authors (Moullade, 1966;Longoria, 1974, 1984; Sigal, 1977; Caron, 1985;Aguado et al., 1988, 1992a; Aguado, 1993b; Coccioni& Premoli Silva, 1994; Cobianchi et al., 1997) and isbriefly described below.

The Blowiella blowi Zone was defined by Moullade(1974) as the interval from the FO of B. blowi to theFO of Schackoina cabri. The lower limit of this zonehas not been identified in the Cau section. Samples13b–15.4 are included within this zone and the‘nannoconid crisis’ is recorded from the uppermostpart (see also Erba, 1994; Cobianchi et al., 1997).

The Schackoina cabri Zone was defined by Bolli(1959) as encompassing the total range of S. cabri. Inthe Cau section, S. cabri ranges from sample 15.5 tosample 22a, the latter also containing the FO ofGlobigerinelloides ferreolensis. Both the LO of S. cabriand FO of G. ferreolensis occur within the Furcata

ammonite Zone of early Aptian age. The FO of thenannofossil Eprolithus floralis occurs in the uppermostpart of the S. cabri Zone. The lower part of the zonecoincides with the main anoxic event which, as men-tioned previously, is marked by a net change in thelithologic record.

The Globigerinelloides ferreolensis Zone was definedby Moullade (1966) as the interval from the LO ofL. cabri to the FO of G. algerianus. The FO of G.algerianus is registered in sample 23d. The lowermostpart of this zone should be early Aptian in age, andcorrelatable with the Furcata ammonite Zone. Noconsistent ammonite data are known for the rest of theG. ferreolensis Zone.

The Globigerinelloides algerianus Zone was definedby Moullade (1966) as encompassing the total rangeof G. algerianus. The LO of G. algerianus is recordedin sample 24 in the Cau section. Samples 23d–24 are,therefore, assigned to this zone. True hedbergellids(Hedbergella trocoidea) first occur within it.

The Hedbergella trocoidea Zone is used here in thesense of Sigal (1977; non sensu Longoria, 1974), whodefined it as the interval from the LO of G. algerianusto the FO of Ticinella bejaouaensis. T. bejaouaensis firstappears in sample 24b1 in the Cau section; hence,samples 24a–24.2 are assigned to this zone.

The Ticinella bejaouaensis Zone, according to Caron(1985), is defined as the interval from the FO of T.bejaouaensis to the FO of Ticinella primula. This latter‘event’ has not been recognized in the Cau sectionowing to unfavourable outcrop conditions. As a result,only part of the zone can be identified.

4.3. Ammonite biostratigraphy

The lower part of the Cau section has yielded rela-tively abundant ammonites. Although their state ofpreservation is mediocre, they have enabled us torecognize most of the zones commonly used forthe Upper Barremian–Lower Aptian interval in theMediterranean area (Hoedemaeker & Company,1993).

The lowest beds of the section can be assignedwithout difficulty to the Sartousiana Zone, given thepresence (in samples 10 and 10b) of numerous speci-mens of the index species, Heinzia sartousiana,accompanied by Camereiceras sp. and Barremites sp.

Slightly higher in the succession (sample 11), smallnuclei of heteroceratids appear, possibly indicatingeither the Giraudi Zone or the Sarasini Zone. We leantowards the latter because, as indicated above(Section 3.1), we have detected the FO of Flabellitesoblongus, in the same levels an ‘event’ that in othersections of the Betic Cordillera is recorded from the



lower half of the Sarasini Zone (Aguado et al., 1997).This implies that sediments corresponding to theFeraudianus and Giraudi Zones, if they exist in thearea studied, are hidden in the covered intervalbetween beds 10b and 11.

Beds 12a and 12 have furnished some specimens ofDeshayesites cf. euglyphus and Deshayesites cf. luppovi,together with a less significant fauna includingBarremites strettostoma. Both of these species have beenrecorded from the Tuarkyricus Zone and the WeissiZone (Bogdanova, 1979; Bogdanova & Tovbina,1995). The nannoplankton assemblage at these samelevels (Section 3.1) leads us to attribute these ammo-nites to the Weissi Zone. Sample 13 contains not onlyDeshayesites cf. euglyphus also a species characteristicof this zone, Deshayesites forbesi. The sediments corre-sponding to the lowermost Aptian (Tuarkyricus Zone)are probably not, therefore, represented in the Causection; this stratigraphic interval should be includedin the discontinuity at the top of level 11c.

From level 13b upwards, coinciding with the ‘nanno-conid crisis’, and throughout the predominantly marlyinterval corresponding to the anoxic event, the ammo-nite assemblages are composed exclusively of forms ofminor biostratigraphic significance. They are juveniledesmoceratids (mainly Pseudohaploceras sp. indet.)which can be very abundant at some levels, andoccasionally accompanied by indeterminate fragmentsof heteromorphs and cheloniceratids.

Up to level 20.4, which coincides with the top of thedark marls, no new characteristic ammonites appear.In this bed, the index species of the Deshayesi Zone,Deshayesites deshayesi, appears together with somespecimens of Australiceras? sp. and Pseudohaplocerassp. As a result, we cannot place the boundary betweenthe Weissi and the Deshayesi Zones in the Cau sectionat the moment. In the Vocontian Basin (SE France),this boundary has been claimed to coincide with thebase of the ‘Niveau Goguel’ (Delanoy, 1995;Magniez-Janin et al., 1997), but the arguments useddo not appear to be definitive.

The youngest identifiable ammonites are Dufrenoyiadufrenoyi, Costidiscus gr. recticostatus and Pseudohap-loceras sp. They occur at levels 21b and 22a andindicate the Furcata Zone of the uppermost LowerAptian. Above these levels, it has not been possible toidentify the scarce ammonites collected.

5. Conclusions

The Upper Barremian–Aptian deposits of the Causection in the Inner Prebetic Domain of the BeticCordillera are characterized by hemipelagic facies thatindicate deposition on a distal carbonate ramp. They

are assigned to the Almadich Formation. Within thesefacies a thick interval of blue to black shales and marlsis interpreted to be the result of an anoxic episode.Dated as early Aptian, this may represent the localrecord of Ocean Anoxic Event 1a.

By means of an integrated biostratigraphic analysisof the section, we have pursued two main objectives; arefined biostratigraphic framework, and a detaileddating of events as important as the OAE 1a and thebase of a second order Aptian stratigraphic sequence.

The nannofossil assemblages recorded from theCau section have a marked Tethyan character. TheMicrantholithus hoschulzii, Hayesites irregularis andRhagodiscus angustus Zones have been identified. Inaddition to the biohorizons containing the zonalmarkers, others have been identified, such as the FOof R. angustus and the LOs of Nannoconus steinmannii,N. bucheri and Diazomatolithus lehmanii. Owingto a stratigraphic discontinuity encompassing theBarreman/Aptian boundary and to the outcrop con-ditions, the lower part of the H. irregularis Zone is notrepresented. This fact is indicated by the presence ofboth Nannoconus truitti and Baarudosphaera africanawithin the lowermost beds assigned to this zone in theCau section. It is known from other sections in theSubbetic Domain (Aguado et al., 1997) that the FOsof these species are recorded above the lower limit ofthe zone. A quantitative study has enabled the preciseidentification of the ‘nannoconid crisis’, the beginningof which clearly precedes the local record of the OAE1a, and its correlation with other bio-events.

The state of preservation of the planktonicforaminifera in the Aptian sediments of the Causection has made it possible for us to use the mostrecent taxonomy based fundamentally on wall textureand chamber morphology. Coinciding approximatelywith the Lower Aptian/Upper Aptian boundary, weencountered a major change in the composition of theassemblages, from being dominated by representativesof the Family Praehedbergellidae to being mainlycomposed of Hedbergellidae. We have identified theBlowiella blowi, Schackoina cabri, Globigerinelloidesferreolensis, G. algerianus, Hedbergella trocoidea andTicinella bejaouaensis Zones. Within the interval corre-sponding to the local record of OAE 1a, we have alsoidentified other biohorizons such as those containingthe FOs of Blowiella duboisi and the genera Lilli-putianella and Claviblowiella and observed an increasein the number of chambers in the plexus of Blowiellablowi. Coinciding with the main anoxic episode, theplanktonic foraminiferal assemblages are composedof a significant number of forms with elongatedchambers and/or tubulospines assigned to the genera


Claviblowiella, Leupoldina, Lilliputianella and Schack-oina. The FO of S. cabri was recorded near the baseof the anoxic interval that represents the local recordof the OAE 1a.

Most of the Upper Barremian and Lower Aptianammonite zones have been identified. We detected adiscontinuity affecting the Tuarkyricus Zone and thelower part of the Weissi Zone which could not beidentified. It was not possible to establish a clearboundary between the Weissi and Dehayesi Zonesowing to the absence of deshayesitids in levels near,and coincident with, the anoxic event.

Our integrated biostratigraphic study has enabledus to establish direct correlations, on many occasionsfrom the same samples, between the zonal scales andbioevents of the three fossil groups analysed. Accord-ing to all the biostratigraphic data, the base of theabove-mentioned second order Aptian stratigraphicsequence is placed within the discontinuity affectingthe lowermost Aptian deposits (Tuarkyrikus andlower part of the Weissi Zones).

Acknowledgements

This study was co-financed by Projects PB97-0826,PB-960429 and PB-93-1150-C02-02 (DGICYT) andResearch Groups 4064 and RNM200 (Junta deAndalucıa). Sample preparation (smear slides andwashing) was performed by A. Carrillo at the GeologyDepartment, EU Politecnica de Linares. We thank MrDavid Nesbitt for his help in translating theoriginal version of this paper into English, and Dr J.Mutterlose (Bochum), Dr M. Simmons (Aberdeen)and Prof. D. Batten (Aberystwyth) for critical reviewsand constructive suggestions.

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Appendix

List of identified taxa, with author attributions and dates.

Calcareous nannofossilsAssipetra infracretacea (Thierstein, 1973) Roth, 1973Assipetra terebrodentaria (Applegate et al. in Covington & Wise,

1987) Rutledge & Bergen, 1994Axopodorhabdus dietzmannii (Reinhardt, 1965) Wind & Wise in

Wise & Wind, 1977Biscutum ellipticum (Gorka, 1957) Grun in Grun & Allemann, 1975Braarudosphaera africana Stradner, 1961Calciosolenia murrayi Gran, 1912Chiastozygus litterarius (Gorka, 1957) Manivit, 1971Conusphaera rothii (Thierstein, 1971) Jakubowski, 1986Cretarhabdus conicus Bramlette & Martini, 1964Crucibiscutum hayi (Black, 1971) Jakubowski, 1986Cyclagelosphaera margerelii Noel, 1965Diazomatolithus lehmanii Noel, 1965Discorhabdus ignotus (Gorka, 1957) Perch-Nielsen, 1968Ellipsagelosphaera britannica (Stradner, 1963) Perch-Nielsen, 1968Ellipsagelosphaera fossacincta Black, 1971Ellipsagelosphaera ovata (Bukry, 1969) Black, 1973Flabellites oblongus (Bukry, 1969) Crux, 1982Haqius circumradiatus (Stover, 1966) Roth, 1978Hayesites irregularis (Thierstein in Roth & Thierstein, 1972)

Covington & Wise, 1987Helenea chiastia Worsley, 1971Lithraphidites carniolensis Deflandre, 1963Manivitella pemmatoidea (Deflandre ex Manivit, 1961) Thierstein,

1971Micrantholithus hoschulzii (Reinhardt, 1966) Thierstein, 1971Micrantholithus obtusus Stradner, 1963Micrantholithus stellatus Aguado, 1997Mitosia infinita Worsley, 1971Nannoconus bucheri Bronnimann, 1955Nannoconus circularis Deres & Acheriteguy, 1980Nannoconus elongatus Bronnimann, 1955Nannoconus minutus Bronnimann, 1955Nannoconus steinmannii Kamptner, 1931 ssp. steinmanniiNannoconus truittii Bronnimann, 1955Nannoconus vocontiensis Deres & Acheriteguy, 1980Nannoconus wassallii Bronnimann, 1955Palaeomicula maltica (Worsley, 1971) Varol & Jakubowski, 1989Percivalia fenestrata (Worsley, 1971) Wise, 1983Podorhabdus gorkae Reinhardt, 1969


Retecapsa surirella (Deflandre in Deflandre & Fert, 1954) Grun inGrun & Allemann, 1975

Rhagodiscus asper (Stradner, 1963) Reinhardt, 1967Rhagodiscus gallagheri Rutledge & Bown, 1996Rhagodiscus splendens (Deflandre, 1953) Verbeek, 1977Rotelapillus laffittei (Noel, 1957) Noel, 1973Staurolithites crux (Deflandre in Deflandre & Fert, 1954) Caratini,

1963Staurolithites matalosus (Stover, 1966) Cepek & Hay, 1969Staurolithites mitcheneri (Applegate & Bergen, 1988) Rutledge &

Bown, 1998Stoverius achylosus (Stover, 1966) Perch-Nielsen, 1986Tegumentum stradneri Thierstein in Roth & Thierstein, 1972Tetrapodorhabdus decorus (Deflandre in Deflandre & Fert, 1954)

Wind & Wise in Wise & Wind, 1977Tranolithus gabalus Stover, 1966Watznaueria barnesae (Black in Black & Barnes, 1959) Perch-

Nielsen, 1968Watznaueria biporta Bukry, 1969Zeugrhabdotus diplogrammus (Deflandre in Deflandre & Fert, 1954)

Burnett in Gale et al., 1996Zeugrhabdotus embergeri (Noel, 1959) Perch-Nielsen, 1984Zeugrhabdotus noeliae Rood, Hay & Barnard, 1971Zeugrhabdotus scutula (Bergen, 1994) Rutledge & Bown, 1996

Planktonic foraminifera‘Biglobigerinella’ barri (Bolli, Loeblich & Tappan, 1957)Blefuscuiana aptiana (Bartenstein, 1965) s.s.Blefuscuiana convexa (Longoria, 1974)Blefuscuiana daminiae Banner, Copestake & White, 1993Blefuscuiana excelsa (Longoria, 1974) s.s.Blefuscuiana gorbachikae (Longoria, 1974)Blefuscuiana hispaniae (Longoria, 1974)Blefuscuiana infracretacea (Glaessner, 1936) ssp. occidentalis

Boudagher-Fadel et al., 1995Blefuscuiana laculata Banner, Copestake & White, 1993Blefuscuiana mitra Banner & Desai, 1988Blefuscuiana occulta (Longoria, 1974)Blefuscuiana sp. cf. B. occulta (Longoria, 1974)Blefuscuiana sp. cf. B. praetrocoidea (Krechmar & Gorbachik, 1986)Blefuscuiana rudis Banner, Copestake & White, 1993Blefuscuiana speetonensis Banner & Desai, 1988 s.s.Blowiella blowi (Bolli, 1959)Blowiella sp. cf. B. blowi (Bolli, 1959)Blowiella duboisi (Chevalier, 1961)Blowiella gottisi (Chevalier, 1961)

Blowiella maridalensis (Bolli, 1959)Blowiella moulladei Boudagher-Fadel, 1995Blowiella solida Krechmar & Gorbachik, 1986Claviblowiella saundersi (Bolli, 1959)Claviblowiella sigali (Longoria, 1974)Globigerinelloides algerianus Cushman & Ten Dam, 1948Globigerinelloides ferreolensis (Moullade, 1961)Gorbachikella anteroapertura Boudagher-Fadel et al., 1995Gorbachikella kugleri (Bolli, 1959)Hedbergella trocoidea (Gandolfi, 1942)Leupoldina protuberans Bolli, 1957Lilliputianella bizonae (Chevalier, 1961)Lilliputianella globulifera (Kretchmar & Gorbachik, 1971)Lilliputianella sp. cf. L. globulifera (Kretchmar & Gorbachik, 1971)Lilliputianella kuhryi (Longoria, 1974)Lilliputianella labocaensis (Longoria, 1974)Lilliputianella longorii Banner & Desai, 1988Lilliputianella roblesae (Longoria, 1974)Praehedbergella ruka Banner, Copestake & White, 1993Praehedbergella sigali (Moullade, 1966)Praehedbergella tuschepsensis (Antonova, 1964)Schackoina cabri Sigal, 1952Ticinella bejaouaensis Sigal, 1966

AmmonitesAustraliceras sp.Barremites sp.Barremites strettostoma (Uhlig, 1883)Camereiceras sp.Cheloniceratidae indet.Costidiscus gr. recticostatus (d’Orbigny, 1841)Deshayesites cf. euglyphus Casey, 1964Deshayesites cf. luppovi Bogdanova, 1983Deshayesites deshayesi (d’Orbigny, 1841)Deshayesites forbesi Casey, 1961Deshayesites sp.Desmoceratidae indet.Dufrenoyia dufrenoyi (d’Orbigny, 1841)Dufrenoyia sp.Heinzia sartousiana (d’Orbigny, 1841)Heteroceratidae indet.Pseudohaploceras sp.Pseudosaynella sp.Toxoceratoides sp.Zuercherella sp.