GeolCarp_Vol67_No3_223

www.geologicacarpathica.com

GEOLOGICA CARPATHICA

, JUNE 2016, 67, 3, 223–238

doi: 10.1515/geoca-2016-0015

Calcareous nannofossils of the Jurassic/Cretaceous boundary

strata in the Puerto Escaño section (southern Spain)

— biostratigraphy and palaeoecology

ANDREA SVOBODOVÁ

1,2

and MARTIN KOŠŤÁK

Institute of Geology, The Czech Academy of Sciences, v.v.i., Rozvojová 269, 165 00 Prague 6 - Lysolaje; asvobodova@gli.cas.cz

Institute of Geology and Palaeontology, Faculty of Science, Charles University in Prague, Albertov 6, 128 43 Prague 2;

andrea.svobodova@natur.cuni.cz, martin.kostak@natur.cuni.cz

(Manuscript received July 14, 2015; accepted in revised form March 10, 2016)

Abstract: We obtained material from the Puerto Escaño section (southern Spain) to study the Jurassic/Cretaceous

(J/K) boundary interval. The same samples had already been processed for magnetostratigraphic studies and biostrati-

graphic zonation based on calpionellids and ammonites (Pruner et al. 2010), but not for calcareous nannofossils. The aim

of this study was to process the samples using micropalaeontological analysis and to compare and calibrate results for

calcareous nannofossils with existing magnetostratigraphic and other biostratigraphic data. The calcareous nannofossil

assemblage was dominated by the genera Watznaueria, Cyclagelosphaera, Nannoconus, Conusphaera and Polycostella.

Several nannofossil bioevents were recorded on the basis of the distribution of stratigraphically important taxa, including

zonal and subzonal markers. Based on the lowest occurrences (LO) of M. chiastius, N. globulus minor, N. wintereri,

N. steinmanii minor, N. steinmannii steinmannii, N. kamptneri minor and N. kampteri kamptneri, two nannofossil

subzones (NJT 15b, NJT 17a) and two nannofossil zones (NJT 16, NK-1) were recognized. The paper introduces new

palaeoecological data based on geochemical analysis and macrofauna occurrences.

Key words: Jurassic/Cretaceous boundary, southern Spain, Tethys, biostratigraphy, calcareous nannofossils,

palaeoecology.

Introduction

The existence of two different temperate Realms in the

Northern hemisphere (the Tethyan and the Boreal) during the

J/K boundary interval was associated with the occurrence of

different biotic elements in the two Realms due to their spe-

cific climatic and palaeoceanographic conditions (Zakharov

et al. 2014). Also from this point of view, the J/K boundary

is the last System boundary remaining to be defined. For the

clarification and precise determination of the J/K boundary

strata, several markers have been suggested as keys for the

correlation — namely calpionellids, calcareous nannofossils,

magnetostratigraphy (base M18r, M19.1n, M19n.1r), ammo-

nites, palynomorphs, geochemistry (Wimbledon et al. 2011).

One of the possible markers of the J/K boundary interval is

based on the Calpionella alpina ‘acme zone’. This marker

was identified in the Puerto Escaño section approximately in

the middle part of the M19n magnetozone (Pruner et al.

2010) and it is partly re-interpreted here. The Jurassic/Creta-

ceous (J/K) boundary interval is currently one of the most

studied, because no appropriate stratotype for the base of the

Cretaceous has yet been defined despite an extensive re-

search effort (e.g., Hoedemaeker et al. 1998; Houša et al.

1999, 2004; Lakova et al. 1999; Oszczypko et al. 2004; Mi-

chalík et al. 2009; Reháková et al. 2009; Channell et al.

2010; Grabowski et al. 2010; Lukeneder et al. 2010; Mi-

chalík & Reháková 2011; Wimbledon et al. 2011, 2013;

Wimbledon 2014).

Globally, plankton assemblages with rock-forming poten-

tial developed during this time interval. The remarkable

radiation of the some groups, especially nannoconids and

calpionellids, significantly increases their biostratigraphical

value. Moreover, the richness of different fossil groups (am-

monites, calcareous nannofossils, calpionellids) offers the

opportunity to compare and calibrate different biozonations

and bioevents, improving the knowledge of Jurassic and Cre-

taceous biochronology (Marino et al. 2004). This study is fo -

cused on calcareous nannofossil biostratigraphy, an essential

component of standard multidisciplinary J/K boundary re-

search.

According to Pruner et al. (2010), the lithological succes-

sion of the Puerto Escaño section in southern Spain shows

relatively continuous sedimentation with minor incidence of

hiatuses, conditions favourable for study of palaeomagnetic

polarity. They also reported rich fossil assemblages of

calpionellids and ammonites; calpionellids, in particular,

were very well preserved, highly diversified, and with a full

record of their evolution.

The ammonite fauna of this section has been evaluated by

Olóriz (1978); Olóriz & Tavera (1989, 1990); Tavera et al.

(1994); Olóriz et al. (1995, 2004). Lithology, development

and associated ichnofabric assemblages were investigated by

Caracuel (1996); Caracuel et al. (2000). The first relevant

geochemical data (stable isotopes) from this section, cali-

brated by bio- and magnetostratigraphy were given by Žák

et al. (2011).

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The aim of this paper is to describe the Late Jurassic– Early

Cretaceous calcareous nannofossil assemblage from Puerto

Escaño, and to discuss its biostratigraphy and possible

palaeoecological affinities. We compare these calcareous

nannofossil results with the magnetostratigraphic and bio-

stratigraphic (ammonites, calpionellids) data of Pruner et al.

(2010) to define the J/K boundary interval.

Geological setting

The section at Puerto Escaño, in the province of Córdoba,

south-eastern Spain (Fig. 1) is located in the External

Subbetic, which was positioned during the Late Jurassic and

Early Cretaceous in a more distal, rather epioceanic environ-

ment, located in the S-E Iberian subplate palaeomargin (Co-

imbra et al. 2014a). Generally, the Subbetic Zone is a

complex tectonostratigraphic unit which, palaeogeographi-

cally, was part of the pelagic domain of the southern passive

margin of the Iberian Plate. Palaeoenvironmental characteris-

tics, based on geochemistry of carbonates, were recently

provided by Coimbra et al. (2014a,b) and Coimbra et al.

(2015).

According to Pruner et al. (2010), the Upper Jurassic to

lowermost Cretaceous deposits in the studied section (spe-

cifically GA-7, UTM 30SUG44859) consist of upper Am-

monitico Rosso and related facies, ranging from well-bedded

limestones to clayey nodular limestone horizons reflecting

deposition on a distal, epioceanic swell. The section studied is

dominated by wackestone showing microfacies with

variable contents of radiolarians, calcareous dinoflagellates,

planktonic crinoids, planktonic and benthic foraminifers,

calpionellids, ostracods, cephalopods, echinoderms (plates

and spines), sponge spicules and pelagic bivalves, among

others. Macroscopic fossil remains are mostly ammonites,

and less abundant components are belemnites, brachiopods

and echinoids. Their occurrence is concentrated into several

limited horizons (beds — see below). Tavera et al. (1994)

also reported relatively abundant calcareous nannofossils

from this section.

Fig. 1. Location of the studied section (A). The position of section GA-7 at Puerto Escaño with roads in the surroundings (B). Black area

represents the Betic Cordillera. Maps modified from Pruner et al. 2010.

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Material and methods

Calcareous nannofossils

We used the same rock material as in the previous study by

Pruner et al. (2010) for calcareous nannofossil analysis.

These were unused, very small fragments from samples in

the same stratigraphic interval (an 8.1m-thick part of the sec-

tion straddling the upper Tithonian and lower Berriasian),

and numbering. These remnants provided enough material

for detailed micropalaeontological analysis of the nannofos-

sils. Samples numbered from No. 5 to No. 42 (Fig. 2) were

chosen from throughout the profile (Fig. 2).

Calcareous nannofossils were analysed in 51 smear slides

prepared using the standard techniques described by Švábe-

nická (2001), under an Olympus BX51 light microscope

using an immersion objective with a magnification of 100×.

Digital images of nannofossil specimens were taken with an

Olympus DP70 digital camera. At least 200 specimens were

counted on each slide, to obtain relative abundances and

semiquantitative information about nannofossil species di-

versity. Each slide was then scanned for identification of

scarce, but biostratigraphically important, species.

The set of smear slides is stored at the Institute of Geology

and Palaeontology (Chlupáč´s museum of Earth history),

Faculty of Sciences, Charles University in Prague as item

(IGP/2015/PE001).

Preservation of calcareous nannofossils was characterized

using the abbreviations described by Bown (1992):

VP (very poor)=extreme etching

P (poor)=etching and overgrowth; obscured, damaged, or

destroyed central area structures

M (moderate)=moderate etching or overgrowth.

The scale for the estimate of nannofossil total abundance

was modified after Casellato (2010):

A (abundant): >11 specimens per field of view

C (common): 1–10 specimens per field of view

F (few): 1 specimen every 1–5 fields of view

R (rare): 1 specimen every 6 or more fields of view.

The individual abundances for each species per sample

were counted according to the classification proposed by

Bown (1992):

R (rare)=1–2 specimens

F (few)=3–10 specimens

C (common)=11–100 specimens

A (abundant)=more than 100 specimens (out of 200 specimens).

A full list of the calcareous nannofossil taxa found in this

study is given in alphabetical order in Appendix A. The listed

calcareous nannofossils are indexed according to Perch-

Nielsen (1985) and Bown & Cooper (1998). Biostratigraphic

data were interpreted with reference to the nannofossil zona-

tion of Casellato (2010), commonly used for the Upper Ju-

rassic and the Lower Cretaceous in the Tethyan/

Mediterranean area.

Macrofossil record and geochemistry

The abundance of the macrofossils, also including the

benthic assemblage is based on a simple quantitative analy-

sis, calculating the number of specimens recorded in a

1m-wide unit of the bed. An abundance exceeding 5–10

specimens or fragments per unit is considered herein to be

significantly high (in relation to other beds within the pro-

file, where almost no benthic fauna has been recorded).

Higher belemnite abundance (≥3–4 per unit) should be con-

sidered as a bio-event and it is reported in the “Palaeoecolo-

gy” chapter.

Stable isotope curves have been published by Žák et al.

(2011) and they are reduced herein into a single column

(δ

O — Fig. 3) as δ

C values (varying from 1.14 to 1.53 ‰

V-PDB): they show no significant expressions suitable for

palaeoecological interpretations (see below).

We have used additional AAS geochemical methods as

a tool for recognizing possible terrigeneous (siliciclastic

SiO

and Al

) input within the Puerto Escaño section. All

analysis (concentrated to oxide and element detections) was

carried out using a VARIAN (type SpectrAA 280 FS) instru-

ment at the Faculty of Science, Charles University in Prague

(Laboratories of the Geological Institutes).

Results

In the samples studied, calcareous nannofossils were rare

to abundant and very poorly to moderately well preserved. In

total, 35 calcareous nannofossil taxa were identified (Fig. 2).

The most common component of the assemblage is the genus

Watznaueria, forming nearly 58 % of the nannofossils, in-

cluding W. barnesiae (Fig. 4/1), W. manivitiae (Fig. 4/2), W.

communis (Fig. 4/3), W. fossacincta (Fig. 4/4) and W. britan

nica (Fig. 4/5). W. barnesiae was present in all samples and

ranged from 38 % to 59 % of the total assemblage. The

highest abundance of this species was recorded in bed 19.

The percentages of W. manivitiae ranged from 3 % to 17 %

of the total assemblage and the highest abundances were re-

corded from beds 11 to 22. W. britannica formed approxi-

mately 2 % of the total assemblage, while W. fossacincta and

W. communis occurred only sporadically, representing ~0.5

% of the genus Watznaueria.

The second most common genus is Cyclagelosphaera spp.

(~27 % of all identified nannofossils), particularly C. margerelii

(Fig. 4/6), C. deflandrei (Fig. 4/7) and C. argoensis (Fig. 4/8).

Other abundant genera are: Conusphaera spp. (~7 %) repre-

sented by C. mexicana mexicana (Fig. 4/19, 20) and C. mexi

cana minor, Nannoconus spp. (~4 %) represented by

Nannoconus sp. (Fig. 5/1), N. infans, N. erbae (Fig. 5/2, 3),

N. puer (Fig. 5/4, 5), N. globulus minor (Fig. 5/6–9),

N. globulus globulus (Fig. 5/10–13), N. wintereri (Fig. 5/14–17),

N. steinmannii minor, N. steinmannii steinmannii (Fig. 5/18–21),

N. kamptneri minor, and N. kamptneri kamptneri (Fig. 5/22,

23), and finally Polycostella beckmannii (Fig. 4/22, 23), rep-

resenting more than 2 % of the total nannofossil assemblage.

Three nannoliths - Conusphaera spp., Nannoconus spp. and

Polycostella sp. showed the most significant fluctuations in

the nannofossil assemblage, as shown in previous studies

(e.g., Tremolada et al. 2006a). The percentage representation

and peaks in abundance of these most abundant genera

across the studied profile are presented in Fig. 6.

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Fig. 4. Calcareous nannofossils from the Puerto Escaño section. Cross polarized light; scale bar represents 5 µm. 1 — Watznaueria bar

nesiae; sample 25-1 t. 2 — W. manivitiae; sample 25-2 b. 3 — Watznaueria communis; sample 23-3. 4 — W. fossacincta; sample 41.

5 — W. britannica; sample 24 b. 6 — Cyclagelosphaera margerelii; sample 25-1 t. 7 — C. deflandrei; sample 24 b. 8 — C. argoensis;

sample 30. 9, 10 — Cruciellipsis cuvillieri; 9 — sample 25-2 t, 10 — sample 28. 11–13 — Diazomatolithus lehmanii; 11 — sample

26-2, 12 — sample 41, 13 — sample 39. 14, 15 — Hexalithus noeliae; sample 13. 16 — Microstaurus chiastius; sample 25-2 t.

17 — Zeugrhabdotus embergeri; sample 40. 18 — Z. cooperi; sample 41. 19, 20 — Conusphaera mexicana mexicana; 19 — sample 22 b,

20 — sample 26-3 b. 21 — Pentalith; sample 25-2 t. 22, 23 — Polycostella beckmannii; sample 11. 24 — Lithraphidites carniolensis;

sample 26-3 t.

Discussion

Biostratigraphy

Generally, calcareous nannoflora represent an important

source of biostratigraphic data for the Mesozoic and Cenozoic

ages. Indeed, in some multiproxy studies, the most signifi-

cant stratigraphic data were obtained by the analysis of cal-

careous nannofossils (e.g. Halásová et al. 2012).

Despite the relatively poor preservation of the calcareous

nannofossils in our material, several biostratigraphic events

have been defined. The lowest occurrence (LO) of M. chias

Other nannoliths represented by Faviconus multicolumna

tus, Assipetra infracretacea, Hexalithus noeliae (Fig. 4/14,

15), Lithraphidites carniolensis (Fig. 4/24) and an unidenti-

fied pentalith (one specimen, Fig. 4/21) occurred less fre-

quently, as did other members of the coccolithophorids such as

Zeugrhabdotus embergeri (Fig. 4/17), Z. cooperi (Fig. 4/18),

Z. erectus, Rhagodiscus asper, Diazomatolithus lehmanii

(Fig. 4/11–13), Cruciellipsis cuvillieri (Fig. 4/9, 10), Dis

corhabdus ignotus, Microstaurus chiastius (Fig. 4/16) and

Biscutum constans. The preservation and total abundance of

calcareous nannofossils and the individual abundances for

each species are summarized in Fig. 7.

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Fig. 5. Calcareous nannofossils from the Puerto Escaño section. XPL — cross polarized light, PPL — plane polarized light. Scale bar repre-

sents 5 µm. 1 — Nannoconus sp., cross section; sample 25-2 b, XPL. 2, 3 — N. erbae; 2 — sample 23-3, XPL; 3 — sample 23-3, PPL

(the same specimen). 4, 5 — N. puer; 4 — sample 25-2 b, XPL; 5 — sample 25-2 b, PPL (the same specimen). 6–9 — N. globulus minor;

6 — sample 25-1 b, XPL; 7 — sample 25-1 b, PPL (the same specimen); 8 — sample 25-1 t, XPL; 9 — sample 25-1 t, PPL (the same spec-

imen); 10–13 — N. globulus globulus; 10 — sample 25-1 t, XPL; 11 — sample 25-1 t, PPL (the same specimen); 12 — sample 30, XPL;

13 — sample 30, PPL (the same specimen). 14 –17 — N. wintereri; 14 — sample 35, XPL; 15 — sample 35, PPL (the same specimen);

16 — sample 40, XPL; 17 — sample 40, PPL (the same specimen). 18–21 — N. steinmannii steinmannii; 18 — sample 35, XPL; 19 — sam-

ple 35, PPL (the same specimen); 20 — sample 35, XPL; 21 — sample 35, PPL (the same specimen). 22, 23 — Nannoconus kamptneri

kamptneri; 22 — sample 35, XPL; 23 — sample 35, PPL (the same specimen). 24 — Faviconus multicolumnatus; sample 13.

the NJT 17b Subzone, that continues to the LO of N. stein

mannii minor sensu Casellato (2010). Unfortunately, this

bioevent has not been identified in the studied section and

the LO of N. steinmannii minor has been observed in the

middle parts of the Calpionella Zone and the Jacobi Zone

and corresponds to the LOs of N. steinmannii steinmannii,

N. kamptneri minor and N. kamptneri kamptneri (Figs. 6, 7),

that indicates the beginning of the NK-1 Zone (sensu Bra-

lower et al. 1989). In the interval between the LO of N. win

tereri and the LOs of N. steinmannii minor, N. steinmannii

steinmannii, N. kamptneri minor and N. kamptneri kampt

neri, the Calpionella alpina ‘acme zone’ and the beginning

tius indicating the beginning of the NJT 16 Zone sensu

Casellato (2010) was recorded in the middle part of the

M20n magnetozone, at the end of the Chitinoidella Zone and

in the middle part of the Transitorius Zone (Tavera et al.

1994). The LO of N. globulus minor is found in the middle

part of the M19r magnetozone and indicates the beginning of

the NJT 17 Zone sensu Casellato (2010). The LOs of

N. globulus globulus and C. cuvillieri were recorded in the

lower part of the M19n magnetozone, in the middle part of

the Crassicolaria Zone and at the beginning of the Jacobi

Zone. In the upper part of the Crassicolaria Zone, the LO of

N. wintereri was observed. This indicates the beginning of

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of the Calpionella Zone were observed (Pruner et al. 2010).

It should be noted, that the LO of N. kamptneri minor usually

appears a little above the LO of N. steinmannii minor, but in

this paper it occurs together with the LOs of N. steinmannii

steinmannii and N. kamptneri kampteri in bed 35. This anoma-

ly can be explained by the very poor preservation and ex-

treme etching of calcareous nannofossils between beds 32

and 34 (Fig. 7). Moreover, the appearance of these four spe-

cies together suggests the presence of a hiatus.

In the previous study by Tavera et al. (1994), the zonal

scheme proposed by Bralower et al. (1989) was used. How-

ever, the zonation of Bralower et al. (1989) and specifically

the definition of subzones NJK-A, NJK-B and NJK-C is

based on the lowest occurrences of species with relatively

delicate structures (Umbria granulosa granulosa and Rote

lapillus laffittei) which may disappear from the association

as a result of preservational effects (Tavera et al. 1994). The

material studied in Tavera et al. (1994) was also not well pre-

served and U. granulosa and R. laffittei taxa have not been

recorded. Indeed, only the NJK-D subzone based on the LO

of N. steinmannii minor was determined. This subzone is cor-

related with the lowermost Berriasian (Tavera et al. 1994),

corresponding to the NKT Zone sensu Casellato (2010). In

comparison, two additional nannofossil subzones (NJT 15b,

NJT 17a) and two nannofossil zones (NJT 16, NK-1) were

recognized in this study. Moreover, the time interval of the

NJT 17 and NKT Zones sensu Casellato (2010) has been ob-

served.

Correlation of the main bioevents with magnetostrati-

graphic data (Pruner et al. 2010) is given in Fig. 8, where the

comparison with other studied sections of the J/K boundary

is shown. The LO of N. wintereri has been recorded in the

M19n magnetozone in Puerto Escaño, as well as in Le Chouet,

Brodno, Foza and Torre de’ Bussi sections. In Fiume Bosso,

this bioevent has been observed in M18r and in Barlya sec-

tion in M17r magnetozone. Similarly, the LO of N. steinman

nii minor within the M19n magnetozone has been recorded in

Puerto Escaño, as well as in Le Chouet, Lokut and Torre

de’ Bussi sections. This bioevent has been recorded in M17n

zone in Foza and Fiume Bosso and in M18r in Barlya sec-

tion. The LO of N. steinmannii steinmannii in M19n magne-

tozone was observed only in Puerto Escaño section, then in

M17r zone in Foza and Fiume Bosso sections.

Palaeoecological interpretations

This chapter is divided into four major parts: the calcareous

nannofossils palaeoecology, macrofossil record, new addi-

tional geochemical data and remarks on the key Bed 28 (J/K

boundary sensu Pruner et al. 2010).

Remarks on selected Late Jurassic calcareous nannofossils

In some cases, calcareous nannofossils provide only spo-

radic information about the palaeoecological conditions of

a depositional area. But there is also evidence reported from

the literature that calcareous nannoplankton may give impor-

tant information regarding the trophic levels of the superfi-

cial oceanic water, about the water temperature or salinity

(e.g., Erba 1992; Tremolada et al. 2006b; Aguado et al.

2008, Mattioli et al. 2008). Some palaeoecological aspects

can also be presented from our study.

Generally, the most dominant genera in studied material

are Watznaueria, Cyclagelosphaera, Nannoconus, Conus

phaera and Polycostella (see chapter Results). This compo-

sition of calcareous nannofossil assemblages across J/K

boundary interval is typical for “low latitudes“ sections and

has been already observed in previous studies (e.g. Tavera et

al. 1994; Bornemann et al. 2003; Tremolada et al. 2006a;

Michalík et al. 2009; Lukeneder et al. 2010).

The most common taxon recorded is Watznaueria

barnesiae, representing 48 % of the entire taphocoenosis. This

species is not susceptible to dissolution and is resistant to dia-

genetic alteration (Hill 1975; Thierstein 1980; Roth 1981;

Roth & Bowdler 1981; Roth & Krumbach 1986). Roth &

Krumbach (1986) described assemblages containing more

than 40 % of W. barnesiae as heavily altered by diagenesis.

However, the genus Watznaueria is generally considered to

be the robust and most successful Mesozoic coccolitho-

phore, in terms of abundance, across the widest range of en-

vironments. By some authors Watznaueria is considered to

be an opportunistic, r-strategist taxa (e.g., Tremolada et al.

2006a; Lees et al. 2006; Tantawy et al. 2009; Colombié et al.

2014; Suchéras-Marx et al. 2015). It is ubiquitous and domi-

nant through most of the Mesozoic, further illustrating its

wide palaeoecological tolerance. In the fossil record, W. bar

nesiae displays a eurytopic, ecologically robust form, and

was one of the first species to settle new habitats. In an eco-

logical sense, it is similar to a recent species, Emiliania hux

leyi. Together with Cyclagelosphaera, these two genera are

stratigraphically long-ranging (Jurassic–earliest Palaeocene)

and morphologically conservative, characteristics of gene-

ralist rather than specialist taxa (e.g., Mutterlose & Wise

1990; Street & Bown 2000; Melinte & Mutterlose 2001;

Bown & Concheyro 2004; Lees et al. 2004). Both are con-

sidered to be cosmopolitan taxa and Lees et al. (2006) sug-

gest, that C. margerelii lived in a higher trophic position

than W. barnesiae and was more r-selected, possibly with

more extreme nutrification affinities.

Species such as Biscutum constans, Discorhabdus ignotus,

Diazomatolithus lehmanii and Zeugrhabdotus erectus oc-

curred only sporadically in our samples. These species indi cate

eutrophic environments and preferred higher nutrient

levels in the oceanic surface water (Roth 1981; Roth &

Bowdler 1981; Roth & Krumbach 1986; Bornemann et al.

2003; Tremolada et al. 2006a,b). Dominance of B. constans

and Zeugrhabdotus spp. is usually considered indicative of

upwelling of cold water rich in nutrients (e.g., Mutterlose &

Kessels 2000; Lees et al. 2005; Hardas & Mutterlose 2007;

Lowery et al. 2014). Higher abundances of the taxa B. con

stans (~11 % of the total assemblage) and Z. erectus (~33 %

of the total assemblage) have been associated with the oc-

currence of black shales and phosphorite deposition (Kessels

et al. 2003). However, these species show very low abun-

dances in our samples — only one specimen of each species

(Fig. 7). It can be explained by high susceptibility to dissolu-

tion of these small coccoliths such as Z. erectus, B. constans

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Fig. 7. Age, calcareous nannofossil zonation and calcareous nannofossil range chart of the Puerto Escaño section, including information on

preservation, total abundance and relative abundance of each taxon. Question marks indicate uncertain species identification of heavily dis-

solved specimens or of nannofossils that occutred only as fragments. Uncertain occurrence was often associated with beds where preserva-

tion was very poor (marked by grey colour) (b=bottom, m=middle, t=top of the bed).



Both situations are probably connected with controlling

factors, namely shallowing but rather the presence of higher

nutrient flux. Shallowing should be excluded, maybe excep-

ting that in Beds 10–17 (i.e. lower and middle part of the

Transitorius ammonite Zone, Fig. 3), where siliciclastic and

influx (with some signs of cyclicity) has been detected

(Fig. 3), but this phenomenon could also be connected with

periods with higher run-off. Probably, the sequences with

abundant benthos represent the result of larger nutrient flux

and subsequent biological productivity. In this context, the

co-occurrence of Biscutum constans, Discorhabdus ignotus,

Diazomatolithus lehmanii and Zeugrhabdotus erectus with

abundant benthic organisms (Fig. 3) — namely eutrophic

communities, should indicate some correlation. This should

also support the results of Bornemann et al. (2003) and

Tremolada et al. (2006a,b), who suggested these nannofossil

taxa were dependent on a eutrophic environment. However,

we assume that more relevant data are needed. We could not

confirm the hypothesis of Mutterlose & Kessels (2000), Lees

et al. (2005), Hardas & Mutterlose (2007), Lowery et al.

(2014) and others relative to the oxygen stable isotope curve,

as no marked cooling (related to upwelling of cold waters) is

recorded in the δ

values (Fig. 3). Nevertheless, we agree

with the conclusion that these taxa represent rather eutrophic

organisms.

Benthic assemblages with skeleton remains occur in Bed

complex 10–17 (in the middle part of the Transitorius

Zone), 22–25 (corresponding approximately to the Duran-

gites ammonite Zone), 28 (“Saccocoma Bed”, suggested

J/K boundary), 33 (lower part of the standard Calpionella

Zone) and 36 (i.e. the middle part of the Calpionella Zone)

(Fig. 3).

Higher abundances of belemnites (active nektonic ani-

mals) have been recorded in beds 14A, 21, 23, 33B–C, and

42. Their occurrences partly correlate with the benthos

presence — namely with more eutrophic conditions (see

above). It has been suggested that higher belemnite abun-

dances (predominantly hibolithids and pseudobelids) are de-

pendent on the transgressive/regressive tracts (Mitchell

2005; Wiese et al. 2009), however, the sea-level oscillations

in the epioceanic development of the Puerto Escaño se-

quence (Olóriz et al. 2004) seems to be under the analytical

limits and further investigations are needed. In this respect,

the winnowing effect, concentrating macrofossil records,

cannot be excluded. On the other hand, the presence of these

remains only in several horizons suggests that their original

habitat, in terms of the environment, lithology and geochemi-

cal record in other beds, in which they are missing, is fully

comparable.

Additional geochemical data

New geochemical data are based on the AAS silicate analy-

sis (see above), which detects especially oxides and element

and D. ignotus (Hill 1975; Thierstein 1980; Roth 1981,

1983; Roth & Krumbach 1986), because a diagenetic over-

print may imply a decrease in their relative abundance

(Giraud 2009; Giraud et al. 2013).

Irregular occurrence of Rhagodiscus asper and Lithraphi

dites carniolensis began in bed 26, which corresponds to the

NJT 17a nannofossil subzone, close below the J/K boundary

interval. Erba (1987) and Erba et al. (1992) interpreted these

species as thermophilous warm-water taxa indicating warm

surface water that was poor in nutrients. In this context, there

is also increased abundance of Nannoconus spp. in this part

of the section (Fig. 6). Nannoconids have been characterized

as typical Tethyan taxa of warm, low-latitude, carbonate-

shelf environments. Similar to Watznaueria are considered

to be r-selected taxa, moreover robust and not susceptible to

dissolution (Street & Bown 2000; Melinte & Mutterlose

2001; Bown & Concheyro 2004; Tremolada et al. 2006b).

These extinct ‘incertae sedis’ nannofossils are often inter-

preted as living in the lower photic zone (e.g., Erba 1994;

Bornemann et al. 2003; Herrle 2003; Barbarin et al. 2012).

The ecological affinities of other abundantly represented

species, Conusphaera and Polycostella, are unknown, except

considering Conusphaera to be a warm water taxon (e.g.,

Melinte & Mutterlose 2001; Mutterlose et al. 2005). However,

Bornemann et al. (2003) proposed for them an ecological

setting similar to that of Nannoconus, due to some similari-

ties in the skeletons of these taxa.

Macrofossil abundance

The ammonite fauna has been studied in great details by

numerous authors (Olóriz & Tavera 1989; Tavera et al.

1994; Pruner et al. 2010; etc.) and the belemnite record (sta-

ble isotopes) has partly been investigated by Žák et al.

(2011). However, the presence and abundance of the macro-

fauna, predominantly benthic filter feeders and substrate

feeders (i.e. brachiopods, echinoids, less abundant bivalves,

echinoderms, sponges) is of great interest. Their presence/

absence is shown in Fig. 3. The constant sedimentation rate

(i.e. 1–5 mm/ky; 2.87 mm/ky with standard deviation

1.17 mm/ky for the whole section; Pruner et al. 2010) as well

as palaeocological conditions — namely a very low variation

of geochemical content in carbonates (also supported by the

stable isotope data; Žák et al. 2011, Fig. 3) within the lime-

stone beds through the whole section do not suggest any rapid

changes either in sedimentology (and bathymetry, not ex-

ceeding the CCD — supported by almost continuous ammo-

nite record) or in taphonomy. Skeletons of all the above

mentioned benthic groups as well as belemnites are com-

posed predominantly by calcite. Thus, we assume very low

fluctuations within the preservation potential of these fossils.

So, the basic scheme for benthic assemblage distribution

should be introduced based on simple fact — namely on rela-

tively abundant/absent.

233

CALCAREOUS NANNOFOSSILS OF THE J/K PUERTO ESCAÑO SECTION (S SPAIN)

GEOLOGICA CARPATHICA

, 2016, 67, 3, 223–238

SAMPLE

PRESER

TION

TOT

ABUNDANCE

Assipetra infracr

etacea

Biscutum constans

Conusphaera m. mexicana

Conusphaera mexicana minor

Cruciellipsis cuvillieri

Cyclagelosphaera ar

goensis

Cyclagelosphaera deflandr

Cyclagelosphaera mar

ger

elii

Diazomatolithus lehmanii

Discor

habdus ignotus

Faviconus multicolumnatus

Hexalithus noeliae

Lithraphidites carniolensis

Micr

ostaurus chiastius

Nannoconus sp.

Nannoconus erbae

Nannoconus globulus globulus

Nannoconus globulus minor

Nannoconus infans

Nannoconus k. kamptneri

Nannoconus kamptneri minor

Nannoconus puer

Nannoconus steinmannii minor

Nannoconus s. steinmannii

Nannoconus winter

Pentalith

Polycostella beckmannii

Rhagodiscus asper

atznaueria barnesiae

atznaueria britannica

atznaueria communis

atznaueria fossacincta

atznaueria manivitae

Zeugr

habdotus cooperi

Zeugr

habdotus ember

geri

Zeugr

habdotus er

ectus

CALCAREOUS NANNO

FOSSILS ZONA

TION

AGE

P C - - R - - - C C R - - - - - F - - R - - R - R - - - - - C F - R C - F -

NK-1

LOWER BERRIASIAN

BERRIASIAN

P C - - R - - - C C R - - - R - F - R F - - - - F - R - - - C F - R C R - -

P C - - F - - - C C - - - - R - C - F F - - - - F - F - - R C F - F C - R -

P C - - F - - - C C R - - - R - F - R F - - R - - - F - - - C F - - C - R -

38 t

VP F - F - - - C C - - - - - - C - - R - - - - - - R - - - A R - - C - - -

38 b

VP R - - F - - - C C - - - - - - C - R - - - R - - - R - - - A - - - C - R -

VP C - - C R - - C C - - - - R - C - R R - - - - - - R - - - A R - - C - R -

P C - - C R - - C C - - - - R - C - R R - - F - F R - - - - C F - - F - R -

M A - - R R - - C C F - - - - - C - F R - R F - C F R - - R C F - - F R R -

VP C - - F - - - C C - - - - - - F ?R - ?R - - - - - - R - - - A R - R C - - -

NJT

17 / NKT

J/K

33B

P C R - C F - - C F - - - - - - F - R - - - - - - - - - - - C R - R C - R -

33A

VP C - - C F - - C C - - - - - - C - - R - - - - - - - - - - C R - - C - R -

32 t

P C R - C R - - C C - - - R R - F - R R - - - - - - - - - - C F - R C - - -

32 m VP R - - R R - - C C R - - - - - F - ?R - - - - - - - - - R - A R - R C - - -
32 b

VP C - - F F - - C C - - - - - - C ?R R R - - - R ?R - F - - - A R - - F - - -

P C - - F F - - C C - - - - R - F - R R - - - R - - R - - - C R - R C R - -

M C - - F F - R C C - - - - - - C R R F - - - R - - R - - R C R - R C R - -

VP C R - C C - - C C R - - - - - F - - R - - - - - - R - - - C R - F C - - -

P F - - C C R - C C - - - - - - F - - R - - - - - - - - - - C F - - C - - -

P C - - C C - - C C R - - - - - F - R R - - - R - - R - - - C F R - C - - -

26-3 t P C - - F F - - C C

- - - R - F - - R - - - - - - - - - R C F - - C - R -

NJT

17a

UPPER TITHONIAN

T I

T H O N I

26-3 b P C - - C C - - C C - - - - - - F - R R - - - - - - - - - - C F - - C - R -
26-2

P F - - C C - - C C R - - - ?R - F - R R - - - - - - - - ?R - C - - - C - - -

26-1

P C - - C C - - C C - - - - - - F - - - - - - - - - - - - - C F - R C - R -

25-2 t M C - - C F R - C C - - - - - R - - R R - - - - - - - ?R - - C F - - C - - -
25-2 b M C - - F F - - C C R - - - - - F - - R R - - R - - - - - - C F - - C - - -
25-1 t P C - - C F - - C C - R - - - - - - R R - - - - - - - - - - C R - - C - R -
25-1 b P C R - C F - - C C - - - - - - - - - R - - - R - - - - - - C R - - C - - -
24 t

VP C - - R F - - C C - - - - - - R - - - - - - - - - - - R - A R - - C - - -

24 b

P C - - F F - - C C R - - - - - - R - R - - - - - - - - ?R - C R R R C - - -

23-3 VP F - - F F - - C C - - - - - - - R - R ?R - - - - - - - - - A F R - C - - -
23-2

P C - - F C - - C C - - - - - - - - ?R ?R - - - - - - - - - - C F - - C - - -

NJT

MIDDLE TITHONIAN

23-1

P C - - F F - - C C R - - - - - R - - - - - - - - - - - F - C F - - C - R -

22 t

P C R - C F - - C C R - - - - - R - ?R ?R - - - - - - - - R - C R - - C - - -

22 m M C R - C F - - C C - - - - - R R - - ?R ?R - - - - - - - F - C F - - C - - -
22 b

P F R - F F - - C C - - - - - - R - - - - - - - - - - - F - C R - R C - - -

P F - R R F - - C C R - - - - R ?R - - - - - - - - - - - F - C F - R C - - -

20 t

P F R - R R - - C C R - - - - - - - - - - - - - - - - - F - A F - - C - - R

20 b

P F R - R F - - C C - - - - - R - - - - ?R - - - - - - - F - A R - R C - - -

VP R R - - - - - C C R - - - - - - - - - - - - - - - - - C - A R - R C - - -

VP F R - - ?R - - C C ?R - - - - R R - - - - - - - - - - - C - A R - - C - - -

P C F - R - - - C C - - - - - ?R - - - - - - - - - - - - F - A F - - C - - -

VP C F - R - - - C C - - - - - - - - - - - - - - - - - - C - A R - F C - - -

VP C F - - - - - C C R - - - - R - - - - - - - - - - - - C - A R - R C - - -

VP C F - - - - - C C R - - R - R ?R - - - - - - - - - - - C - A F - - C - - -

M A R - F R - - C C - R R R - - - - - - - - - - - - - - C - A F - R C - - -

11/12 VP C R - - R - - C C R - - - - - - - - - - - - - - - - - C - A F - R C - - -
11

P F F - - R - - C C R - - ?R - R - - - - - - - - - - - - C - A F - - C - - -

VP C - - - - - - C C - - - - - - - - - - - - - - - - - - C - A F - R C - - -

NJT

15b

P C R - - - - - C C - - - - - - - - - - - - - - - - - - C - A F - - C - R -

P A R - - R - - C C - - - - - - - - - - - - - - - - - - C - A F - F C - - -

234

SVOBODOVÁ and KOŠŤÁK

GEOLOGICA CARPATHICA

, 2016, 67, 3, 223–238

Fig.

Correlatio

selected

sequences

showing

the

M20–M17

interval;

imbledon

al.

(2013),

modified.

Revised

from

imbledon

al.

(201

1).

Brodno

columns

based

data

Houša

al.

(2007)*

and

Michalík

al.

(2009)**.

Data

1–4

Tethys

(base

jacobi

Subzone,

base

Alpina

Subzone,

top

M19n.2n, and

base

Ferasini

Subzone)

and

their

approximate

posi

tions at Nordvik (column after Houša et al. 2007).

The Barlya column is based on data provided by Platon

Tchoumatchenco

(unpublished magnetozones by B. Galbrun). Lókút after Grabowski

et al. (2010).

The lowest occurences (LOs) of biostratigrafically significant calcareous nannofossils in the Puerto Escaño section (this work) are indicated in red.

235

CALCAREOUS NANNOFOSSILS OF THE J/K PUERTO ESCAÑO SECTION (S SPAIN)

GEOLOGICA CARPATHICA

, 2016, 67, 3, 223–238

components of the sediment. Three major events representing

marked inputs of terrigeneous material were detected at the

levels of the beds 8, 30-31 and 40-41 (Fig. 3). Positive curve

excursions of oxides SiO

and Al

are accompanied by

marked increases (however, at the lower concentrations) of

FeO, K

O, TiO

and Na

O, as well as marked increases of

238

, Th

232

, Pb

208

and other rare elements (unpublished

data), clearly documenting a terrigeneous input. A positive

(however, not so straightforward) correlation between the in-

crease of calcareous nannoplankton diversity and the terrige-

neous input is indicated in the beds 13, 29-31 and 40-41, but

less marked in the in beds 22 and 35 levels. Higher terrige-

neous input should be evidence of weathering and/or the re-

gression trend. In the studied profile, we should recognize

three major sequences with higher terrigeneous input —

namely bed 8 (and overlying beds 9-14A), beds 29-31 and

the upper part of bed 40 into the lower part of bed 41. The al-

most identical chemical composition of oxides and element

content probably proves the same source area. However, its

position within the Subbetic Basin margin is unknown.

The stable isotope data were precisely analysed by Žák et

al. (2011). The δ

O curve (Fig. 3), shows minor positive ex-

cursions in beds 8, 12, 23, 27-32, 34 and 40 levels, and the

major negative peak has been recorded in bed 26-3 (Fig. 3).

Only the peak in the Bed 12 corresponds to the higher abun-

dance of Polycostella sp. (Fig. 6) and the positive excursion

at level 34 corresponds to abundance increase of Conus

phaera spp. In this respect, we cannot estimate any signifi-

cant correlation between the calcareous nannofossil diversity

development and the temperature changes, as only limited

variations in the δ

O curve have been recorded.

Remarks on the key Bed 28

Bed 28 (J/K boundary sensu Tavera et al. 1994; see also

Pruner et al. 2010) contains a large accumulation of smaller

globular (spherical) Calpionella alpina (Pruner et al. 2010).

Bed 28 (“Saccocoma Bed” herein, Fig. 3) also contains ex-

tremely large, calcitic skeletal debris of Saccocoma skele-

tons, and thus, the reworking of smaller calpionellids cannot

be excluded. However, there is no sign of siliciclastic input

in this horizon, and the highest ratio between Ca-oxide and

minimum SiO

and Al

input is recorded in this bed

(Fig. 3). The opposite trend is visible in the overlying beds,

beds 29-31, where the marked decrease of Ca-oxide and in-

crease of siliciclastics input (represented by SiO

), as well as

an increase of Al

seems to prove a run-off increase due to

an acceleration of weathering and/or rather a regressive se-

quence. The regression hypothesis should be supported by

the presence of a marked increase in crassicollarian abun-

dance (C. parvula — parvula Acme zone, “CPAZ” sensu

Pruner et al. 2010), suggested to have been reworked from

older upper Tithonian limestones and incorporated into the

micrite matrix of the Alpina Subzone (Wimbledon et al.

2013). We agree with this interpretation and we preliminari-

ly suggest the section studied represents a regressive se-

quence (beds 29-31). A similar trend is recorded in the

uppermost part of bed 40 and especially in the lower part of

bed 41, where decrease of Ca-oxide and increase of SiO

and

(as a product of the laterite weathering) input have

been detected (Fig. 3).

Conclusions

We have studied the calcareous nannofossils from the

GA-7 section at Puerto Escaño. Consistent with the results of

Tavera et al. (1994), the dominant genera in the calcareous

nannofossil associations of the section are Watznaueria spp.,

Cyclagelosphaera spp., Nannoconus spp. and Conusphaera

spp. and Polycostella sp.

We have applied the nannofossil zonation of Casellato

(2010), and recognized several nannofossil LOs (Figs. 6, 7):

Microstaurus chiastius, N. globulus minor, N. globulus globu

lus and Cruciellipsis cuvillieri, N. wintereri and finally

N. steinmannii minor, N. steinmannii steinmannii, N. kampt

neri minor and N. kamptneri kampteri. Moreover, the com-

position of the nannofossil assemblage indicated a warm,

carbonate-shelf environment with oligotrophic conditions.

The J/K boundary transition has been approximated with

the LO of N. wintereri to the LOs of N. steinmannii minor,

N. steinmannii steinmannii, N. kamptneri minor and N. kampt

neri kamptneri. This interval is located between beds 27 and

35. Consistent with Pruner et al. (2010), the J/K boundary

lies in approximately the upper middle part of the M19n

magnetozone and includes the Calpionella alpina ‘acme

zone’ (Fig. 2).

The co-occurrence of taxa Biscutum constans, Discorhab

dus ignotus, Diazomatolithus lehmanii and Zeugrhabdotus

erectus with abundant benthic organisms should confirm

their preferences for higher nutrient levels.

The diversity of calcareous nannofossils at higher levels

partly corresponds to the levels with terrigeneous input, well

documented by geochemical methods.

Here we suggest that the increase in abundance of Crassi

collaria parvula (i.e. — Parvula Acme zone, “CPAZ” sensu

Pruner et al. 2010) is the result of reworking from older up-

per Tithonian deposits, in relation to the regressive sequence

between beds 29-31.

Acknowledgements: The authors thanks the Institute of

Geology of the Czech Academy of Sciences for financial

support (Research Plan RV067985831), the Grant Agency of

the Czech Republic (Grant No. GACR 16-09979S), the

Project 7AMB14SK201 supported by the Ministry of Educa-

tion of the Czech Republic and PRVOUK P44. Ladislav Str-

nad (Laboratories of the Geological Institutes, Prague) is

greatly acknowledged for geochemical analysis. We are in-

debted to William Wimbledon (University of Bristol) for lin-

guistic corrections and valuable comments. We are grateful

to the journal reviewers Kristalina Stoykova and an anony-

mous reviewer, for insightful comments on an earlier draft.

References

Aguado R., O’Dogherty L. & Sandoval J. 2008: Fertility changes in

surface waters during the Aalenian (mid-Jurassic) of the

236

SVOBODOVÁ and KOŠŤÁK

GEOLOGICA CARPATHICA

, 2016, 67, 3, 223–238

Western Tethys as revealed by calcareous nannofossils and

carbon-cycle perturbations. Mar. Micropaleontol. 68,

268–285.

Aguado R., Company M., O´Dogherty L., Sandoval J. & Tavera

J.M. 2014: Late Hauterivian–early Barremian calcareous nan-

nofossil biostratigraphy, palaeoceanography, and stable iso-

tope record in the Subbetic domain (southern Spain).

Cretaceous Res. 49, 105–124.

Barbarin N., Bonin A., Mattioli E., Pucéat E., Cappetta H., Gréselle

B., Pittet B., Vennin E. & Joachimski M. 2012: Evidence for

a complex Valanginian nannoconid decline in the Vocontian

basin (South East France). Mar. Micropaleontol. 84–85,

37–53.

Bornemann A., Aschwer U. & Mutterlose J. 2003: The impact of

calcareous nannofossils on the pelagic carbonate accumulation

across the Jurassic-Cretaceous boundary. Palaeogeogr. Palaeo

climatol. Palaeoecol. 199, 187–228.

Bown P.R. 1992: New calcareous nannofossil taxa from the Juras-

sic/Cretaceous boundary interval of sites 765 and 261, Argo

Abyssal Plain. Initial Reports of the Ocean Drilling Program,

Scientific Results 123, 369–379.

Bown P.R. & Concheyro A. 2004: Lower Cretaceous calcareous

nannoplankton from the Neuquén Basin, Argentina. Mar.

Micropaleontol. 52, 51–84.

Bown P.R. & Cooper M.K.E. 1998: Jurassic. In: Bown P.R. (Ed):

Calcareous nannofossil biostratigraphy. Kluwer Academic

Publishers, Cambridge, 34–85.

Bralower T.J., Monechi S. & Thierstein H.R. 1989: Calcareous nan-

nofossils Zonation of the Jurassic-Cretaceous Boundary inter-

val and correlations with the Geomagnetic Polarity Timescale.

Mar. Micropaleontol. 14, 153–235.

Caracuel J. 1996: Association of invertebrates, evolution, sedimen-

tation and biostratigraphical interpretations in of an epioceanic

environment of in the Western Tethys (Upper Jurassic). PhD

Thesis, Universidad de Granada, 1–475 (in Spanish with

English summary).

Caracuel J., Monaco P. & Olóriz F. 2000: Taphonomic tools to

evaluate sedimentation rates and stratigraphic completeness in

Rosso Ammonitico Facies (epioceanic Tethyan Jurassic). Riv.

Ital. Paleontol. Stratigr. 106, 3, 353–368.

Casellato C.E. 2010: Calcareous nannofossil biostratigraphy of Up-

per Callovian-Lower Berriasian successions from Southern

Alps, North Italy. Riv. Ital. Paleontol. Stratigr. 116, 357–404.

Channell J.E.T., Casellato C.E., Muttoni G. & Erba E. 2010: Mag-

netostratigraphy, nannofossil stratigraphy and apparent polar

wander for Adria-Africa in the Jurassic–Cretaceous boundary

interval. Palaeogeogr. Palaeoclimatol. Palaeoecol. 293,

51–75.

Coimbra R., Immenhauser A. & Olóriz F. 2014a: Spatial

geochemistry of Upper Jurassic marine carbonates (Iberian

Subplate). Earth Sci. Rev. 139, 1–32.

Coimbra R., Rodríguez-Galiano V., Olóriz F. & Chica-Olmo M.

2014b: Regression trees for modeling geochemical data — An

application to Late Jurassic carbonates (Ammonitico Rosso).

Comput. Geosci. 73, 198–207.

Coimbra R., Immenhauser A., Olóriz F., Rodríguez-Galiano

V., Chica-Olmo M. & Pufahl P. 2015: New insights into

geochemical behaviour in ancient marine carbonates (Upper

Jurassic Ammonitico Rosso): Novel proxies for interpreting

sea-level dynamics and palaeoceanography. Sedimentology 62,

266–302.

Colombié C., Giraud F., Schnyder J., Götz A.E., Boussaha M., Au-

rell M. & Bádenas B. 2014: Timing of sea level, tectonics and

climate events during the uppermost Oxfordian (planula zone)

on the Iberian ramp (northeast Spain). Palaeogeogr. Palaeo

climatol. Palaeoecol. 412, 17–31.

Erba E. 1987: Mid-Cretaceous cyclic pelagic facies from the Um-

brian-Marchean Basin: What do calcareous nannofossils sug-

gest? Int. Nannoplankton Assoc. Newsletter 9, 52–53.

Erba E. 1992: Middle Cretaceous calcareous nannofossils from the

western Pacific (Leg 129): evidence for paleoequatorial cros-

sings. Proceedings of the Ocean Drilling Program, Scientific

Results 129, 189–201.

Erba E. 1994: Nannofossils and superplumes: the early Aptian

“nannoconid crisis“. Paleoceanography 9, 483–501.

Erba E., Castradori D., Guasti G. & Ripepe M. 1992: Calcareous

nannofossils and Milankovitch cycles: The example of the Al-

bian Gault Clay Formation (southern England). Palaeogeogr.

Palaeoclimatol. Palaeoecol. 93, 47–69.

Giraud F. 2009: Calcareous nannofossil produktivity and carbonate

production across the Middle–Late Jurassic transition in the

French Subalpine Basin. Geobios 42, 699–714.

Giraud F., Reboulet S., Deconinck J.F., Martinez M., Carpentier A.

& Bréziat C. 2013: The Mid-Cenomanian Event in south-

eastern France: Evidence from palaeontological and clay mine-

ralogical data. Cretaceous Res. 46, 43–58.

Grabowski J., Michalík J., Pszczółkowski A. & Lintnerová O.

2010: Magneto-, and isotope stratigraphy around the Jurassic/

Cretaceous boundary in the Vysoká Unit (Malé Karpaty

Mountains, Slovakia): correlations and tectonic implications.

Geol. Carpath. 61, 4, 309–326.

Halásová E., Vašíček Z., Jansa L., Reháková D. & Skupien P. 2012:

Lower Cretaceous succession and biostratigraphy near over-

thrust plane of Silesian Nappe (Ostravice River Channel, Outer

Western Carpathians, Czech Republic). Bull. Geosci. 87, 2,

383–406.

Hardas P. & Mutterlose J. 2007: Calcareous nannofossil assemblages

of Oceanic Anoxic Event 2 in the equatorial Atlantic: evidence

of an eutrophication event. Mar. Micropaleontol. 66, 52–69.

Herrle J.O. 2003: Reconstructing nutricline dynamics of mid-Creta-

ceous oceans: evidence from calcareous nannofossils from the

Niveau Paquier black shale (SE France). Mar. Micropaleontol.

47, 307–321.

Hill M.E. 1975: Selective dissolution of mid-Cretaceous (Ceno-

manian) calcareous nannofossils. Micropaleontology 21,

227–235.

Hoedemaeker P.J., Krs M., Man O., Parés J.M., Pruner P. & Ven-

hodová D. 1998: The Neogene remagnetization and petromag-

netic study of the Early Cretaceous limestone beds from the

Río Argos (Caravaca, Province Murcia, SE Spain). Geol. Car

path. 49, 1, 15–32.

Houša V., Krs M., Krsová M., Man O., Pruner P. & Venhodová D.

1999: High-resolution magnetostratigraphy and micropaleon-

tology across the J/K boundary strata at Brodno near Žilina,

western Slovakia summary results. Cretaceous Res. 20,

699–717.

Houša V., Krs M., Krsová M., Man O., Pruner P., Venhodová D.,

Cecca F., Nardi G. & Piscitello M. 2004: Combined magneto-

stratigraphic, paleomagnetic and calpionellid investigations

across Jurassic/Cretaceous boundary strata in the Bosso

Walley, Umbria, central Italy. Cretaceous Res. 25, 771–785.

Houša V., Pruner P., Zakharov V.A., Košťák M., Chadima M.,

Rogov M.A., Slechta S., Mazuch M. 2007. Boreal-Tethyan

Correlation of the Jurassic-Cretaceous boundary interval by

magneto- and biostratigraphy. Stratigraphy and Geological

Correlation 15, 3, 297–309.

Kessels K., Mutterlose J. & Ruffell A. 2003: Calcareous nannofos-

sils from the late Jurassic sediments of the Volga Basin (Rus-

sian Platform): evidence for productivity-controlled black

shale deposition. Int. J. Earth Sci. 92, 743–757.

Lakova I., Stoykova K. & Ivanova D. 1999: Calpionellid, nannofos-

sil and cacareous dinocyst bioevents and integrated biochro-

237

CALCAREOUS NANNOFOSSILS OF THE J/K PUERTO ESCAÑO SECTION (S SPAIN)

GEOLOGICA CARPATHICA

, 2016, 67, 3, 223–238

nology of the Tithonian to Valanginian in the Western

Bal kanides, Bulgaria. Geol. Carpath. 50, 2, 151–168.

Lees J.A., Bown P.R., Young J.R. & Riding J.B. 2004: Evidence for

annual records of phytoplankton productivity in the Kim-

meridge Clay Formation coccolith stone bands (Upper Juras-

sic, Dorset, UK). Mar. Micropaleontol. 52, 29–49.

Lees J.A., Bown P.R. & Mattioli E. 2005: Problems with proxies?

Cautionary tales of calcareous nannofossil paleoenvironmental

indicators. Mar. Micropaleontol. 51, 4, 333 – 343.

Lees J.A., Bown P.R. & Young J.R. 2006: Photic zone palaeoenvi-

ronments of the Kimmeridge Clay Formation (Upper Jurassic,

UK) suggested by calcareous nannoplankton palaeoecology.

Palaeogeogr. Palaeoclimatol. Palaeoecol. 235, 110–134.

Lowery Ch.M., Corbett M.J., Leckie R.M., Watkins D., Miceli

Romero A. & Pramudito A. 2014: Foraminiferal and nannofos-

sil paleoecology and paleoceanography of the Cenomanian-

Turonian Eagle Ford Shale of southern Texas. Palaeogeogr.

Palaeoclimatol. Palaeoecol. 413, 49–65.

Lukeneder A., Halásová E., Kroh A., Mayrhofer S., Pruner P., Re-

háková D., Schnabl P., Sprovieri M. & Wagreich M. 2010:

High resolution stratigraphy of the Jurassic-Cretaceous boun-

dary interval in the Gresten Klippenbelt (Austria). Geol. Car

path. 61, 5, 365–381.

Marino M.C., Andreini G., Baldanza A., D’arpa C., Mariotti N.,

Pallini G., Parisi G. & Petti F.M. 2004: Middle Jurassic–early

Cretaceous integrated biostratigraphy (ammonites, calcareous

nannofossils and calpionellids) of the Contrada Diesi section

(south-western Sicily, Italy). Riv. Ital. Paleont. Stratigr. 110,

357–372.

Mattioli E., Pittet B., Suan G. & Mailliot S. 2008: Calcareous nan-

noplankton changes across the early Toarcian oceanic anoxic

event. In the western Tethys. Paleoceanography 23, PA3208.

Melinte M. & Mutterlose J. 2001: A Valanginian (Early Creta-

ceous) „boreal nannoplankton excursion“ in sections from Ro-

mania. Mar. Micropaleontol. 43, 1–25.

Michalík J. & Reháková D. 2011: Possible markers of the Jurassic/

Cretaceous boundary in the Mediterranean Tethys: A review

and state of art. Geosci. Frontiers 2, 4, 475–490.

Michalík J., Reháková D., Halásová E. & Lintnerová O. 2009: The

Brodno section – a potential regional stratotype of the Jurassic/

Cretaceous boundary (Western Carpathians). Geol. Carpath.

60, 3, 213–232.

Mitchell S. F. 2005: Eight belemnite biohorizons in the Cenoma-

nian of northwest Europe and their importance. Geol. J. 40,

363–382.

Mutterlose J. & Wise Jr. S.W. 1990: Lower Cretaceous nannofossil

biostratigraphy of ODP LEG 113 Holes 692B and 693A, Con-

tinental slope off east Antarctica, Weddell Sea. Proceedings of

the Ocean Drilling Program, Scientific Results 113, 325–351.

Mutterlose J. & Kessels K. 2000: Early Cretaceous calcareous nan-

nofossils from high latitudes: implications for palaeobiogeog-

raphy and palaeoclimate. Palaeogeogr. Palaeoclimatol.

Palaeoecol. 160, 347–372.

Mutterlose J., Bornemann A. & Herrle O. 2005: Mesozoic calca-

reous nannofossils – state of the art. Paläontologische

Zeitschrift 79/1, 113–133.

Olóriz F. 1978: Kimmeridgian and lower Tithonian in the central

part of Betic Cordillera (Subbetic Zone). Paleontology. Bio-

stratigraphy. PhD Thesis, University of Granada, nº 84, v. I-II,

1–758 (in Spanish with English summary).

Olóriz F. & Tavera J.M. 1989: The significance of Mediterranean

ammonites with regard to the traditional Jurassic-Cretaceous

boundary. Cretaceous Res. 10, 221–237.

Olóriz F. & Tavera J.M. 1990: The Jurassic-Cretaceous boundary in

Southern Spain. Some eco-stratigraphical considerations.

Transactions Institute of Geology and Geophysics. Siberian

Branch of the Academy of Sciences, CCCP (USSR), Hayka

(Nauka), Mockba (Moscow) 699, 64–77.

Olóriz F., Caracuel J. E., Marques B. & Rodríguez-Tovar F.J. 1995:

Associations of the Tintinnids of the Ammonitico Rosso facies

in the Siera Norte (Mallorca). Revista Española de Paleon

tología Nº Extra Homenaje al Dr. Guillermo Colom, 77–93 (in

Spanish with English abstract).

Olóriz F., Reolid M. & Rodríguez-Tovar F.J. 2004: Microboring

and taphonomy in Middle Oxfordian to lowermost Kimmerid-

gian (Upper Jurassic) from the Prebetic Zone (southern Iberia).

Palaeogeogr. Palaeoclimatol. Palaeoecol. 212, 181–197.

Oszczypko N., Malata E., Švábenická L., Golonka J. & Marko F.

2004: Jurassic-Cretaceous controversies in the Western Car-

pathian Flysch: the “black flysch” case study. Cretaceous Res.

25, 89–113.

Perch-Nielsen K. 1985: Mesozoic calcareous nannofossils. In: Bolli

H.M., Saunders J.B. & Perch-Nielsen K. (Eds.): Plankton

Stratigraphy. Cambridge University Press, Cambridge,

329–426.

Pruner P., Houša V., Olóriz F., Košťák M., Krs M., Man O.,

Schnabl P., Venhodová D., Tavera J.M. & Mazuch M. 2010:

High-resolution magnetostratigraphy and biostratigraphic zo-

nation of the Jurassic/Cretaceous boundary strata in the Puerto

Escaño section (southern Spain). Cretacous Research 31,

192–206.

Reháková D., Halásová E. & Lukeneder A. 2009: The Jurassic-Cre-

taceous boundary in the Gresten Klippenbelt (Nutzhof, Lower

Austria): Implications for Micro- and Nannofacies analysis.

Ann. Naturhist. Mus. Wien 110 A, 345–381.

Roth P.H. 1981: Mid-Cretaceous calcareous nannoplankton from

the Central Pacific: implications for paleoceanography. In:

Thiede, J. et al. (Eds.): Initial Report sof the Deep Sea Drilling

Project 62, 471–489.

Roth P.H. 1983: Jurassic and Lower Cretaceous calcareous nanno-

fossils in the western North Atlantic (Site 534): biostratigra-

phy, preservation, and some observations on biogeography and

paleoceanography. In. Sheridan R.E., Gradstein F.M. et al.

(Eds.): Initial Report sof the Deep Sea Drilling Project 76. U.S.

Government Printing Office, Washington, 587–621.

Roth P.H. & Bowdler J. 1981: Middle Cretaceous nannoplankton

biogeography and oceanography of the Atlantic Ocean. In:

Warme J.E., Douglas R.G. & Winterer E.L. (Eds.): The Deep

Sea Drilling Project: a Decade of Progress. Society of Economic

Paleontologists and Mineralogists Special Publications 32,

517–546.

Roth P.H. & Krumbach K.P. 1986: Middle Cretaceous calcareous

nannofossil biogeography and preservation in the Atlantic and

Indian Oceans: Implications for paleoceanography. Mar.

Micropaleontol. 10, 235–266.

Street C. & Bown P.R. 2000: Palaeobiogeography of early Creta-

ceous (Berriasian–Barremian) calcareous nannoplankton. Mar.

Micropaleontol. 39, 265–291.

Suchéras-Marx B., Mattioli E., Gieraud F. & Escarguel G. 2015:

Paleoenvironmental and paleobiological origins of coccolitho-

phorid genus Watznaueria emergence during the late Aale-

nian-early Bajocian. Paleobiology 41, 3, 415–435.

Švábenická L. 2001: Late Campanian/late Maastrichtian penetra-

tion of high-latitude calcareous nannoflora to the outer western

Carpathian depositional area. Geol. Carpath. 52, 1, 23–40.

Tantawy A.A.A., Keller G. & Pardo A. 2009: Late Maastrichtian

Volcanism in the Indian Ocean: Effects on Calcareous Nanno-

fossils and Planktic Foraminifera. Palaeogeogr. Palaeoclima

tol. Palaeoecol. 284, 63–87.

Tavera J.M., Aguado R., Company M. & Olóriz F. 1994: Integrated

biostratigraphy of the Durangites and Jacobi Zones (J/K

boundary) at the Puerto Escaño section in the southern Spain

238

SVOBODOVÁ and KOŠŤÁK

GEOLOGICA CARPATHICA

, 2016, 67, 3, 223–238

(Province of Cordoba). Geobios, Mémoire Special nº 17,

469–476.

Thierstein H.R. 1980: Selective Dissolution of Late Cretaceous and

Earliest Tertiary calcareous nannofossils: experimental evi-

dence. Cretaceous Res. 2, 165–176.

Tremolada F., Bornemann A., Bralower T.J., Koeberl Ch. & van de

Schootbrugge B. 2006a: Paleoceanographic changes across the

Jurassic/Cretaceous boundary: The calcareous phytoplankton

response. Earth Planet. Sci. Lett. 241, 361–371.

Tremolada F., Erba E. & Bralower T.J. 2006b: Late Barremian to

early Aptian calcareous nannofossils paleoceanography and

paleoecology from the Ocean Drilling Program Hole 641C

(Calicia Margin). Cretaceous Res. 27, 887–897.

Wiese F., Košťák M. & Wood C. J. 2009: The Upper Cretaceous be-

lemnite Praeactinocamax plenus (Blainville 1827) from Lower

Saxony (Upper Cenomanian, Northwest Germany) and its dis-

tribution pattern in Europe. Paläontologische Zeitschrift 83, 2,

309–321.

Wimbledon W.A.P. 2014: Warsaw Remarks - Berriasian Progress.

Volumina Jurassica 12, 1, 107 - 112.

Wimbledon W.A.P., Casellato C.E., Reháková D., Bulot L.G., Erba

Assipetra infracretacea (Thierstein, 1973) Roth, 1973

Biscutum constans (Górka, 1957) Black, 1967

Conusphaera mexicana (Trejo, 1969) subsp. mexicana Bralower in Bralower et al., 1989

Conusphaera mexicana (Trejo, 1969) subsp. minor (Bown & Cooper, 1989) Bralower in Bralower et al., 1989

Cruciellipsis cuvillieri (Manivit, 1956) Thierstein, 1971

Cyclagelosphaera argoensis Bown, 1992

Cyclagelosphaera deflandrei (Manivit, 1966) Roth, 1973

Cyclagelosphaera margerelii Noël, 1965

Diazomatolithus lehmanii Noël, 1965

Discorhabdus ignotus (Górka, 1957) Perch-Nielsen, 1968

Faviconus multicolumnatus Bralower in Bralower et al., 1989

Hexalithus noeliae (Noël, 1956) Loeblich & Tappan, 1966

Lithraphidites carniolensis Deflandre, 1963

Microstaurus chiastius (Worsley, 1971) Bralower et al, 1989

Nannoconus sp. Kamptner, 1931

Nannoconus erbae Casellato, 2010

Nannoconus globulus (Brönnimann, 1955) subsp. globulus Bralower in Bralower et al. 1989

Nannoconus globulus (Brönnimann, 1955) subsp. minor Bralower in Bralower et al. 1989

Nannoconus infans Bralower in Bralower et al. 1989

Nannoconus kamptneri (Brönnimann, 1955) subsp. kamptneri Bralower in Bralower et al., 1989

Nannoconus kamptneri (Brönnimann, 1955) subsp. minor Bralower in Bralower et al. 1989

Nannoconus puer Casellato, 2010

Nannoconus steinmannii (Kamptner, 1931) subsp. minor Deres & Achéritéguy, 1980

Nannoconus steinmannii (Kamptner, 1931) subsp. steinmannii Deres & Achéritéguy, 1980

Nannoconus wintereri Bralower & Thierstein in Bralower et al., 1989

Polycostella beckmannii Thierstein, 1971

Rhagodiscus asper (Stradner, 1963) Reinhardt, 1967

Watznaueria barnesiae (Black in Black & Barnes, 1959) Perch-Nielsen, 1968

Watznaueria britannica (Stradner, 1963) Reinhardt, 1964

Watznaueria communis Reinhardt, 1964

Watznaueria fossacincta (Black, 1971a) Bown in Bown & Cooper, 1989

Watznaueria manivitiae (Bukry, 1973) Moshkovitz & Ehrlich, 1987

Zeugrhabdotus cooperi Bown, 1992b

Zeugrhabdotus embergeri (Noël, 1958) Perch-Nielsen, 1984

Zeugrhabdotus erectus (Deflandre in Deflandre & Fert, 1954) Reinhardt, 1965

E., Gardin S., Verreussel R.M.C.H., Munsterman D.K. & Hunt

CH.O. 2011: Fixing a basal Berriasian and Jurassic/Cretaceous

(J/K) boundary — is there perhaps some light at the end of the

tunnel? Riv. Ital. Paleont. Stratigr. 117, 295–307.

Wimbledon W.A.P., Reháková D., Pszczółkowski A., Casellato

C.E., Halásová E., Frau C., Bulot L.G., Grabowski J., Sobień

K., Pruner P., Schnabl P. & Čížková K. 2013: An account of

the bio- and magnetostratigraphy of the Upper Tithonian–

Lower Berriasian interval at Le Chouet, Drôme (SE France).

Geol. Carpath. 64, 6, 437–460.

Zakharov V.A., Rogov M.A., Dzyuba O.A., Žák K., Košťák M.,

Pruner P., Skupien P., Chadima M., Mazuch M. & Nikitenko

B.L., 2014: Palaeoenvironments and palaeoceanography

changes across the Jurassic/Cretaceous boundary in the Arctic

Realm: case study of the Nordvik section (North Siberia, Rus-

sia). Polar Research 33, 19714.

Žák K., Košťák M., Man O. Zakharov V. A., Rogov M. A., Pruner P.,

Rohovec J., Dzyuba O. S. & Mazuch M. 2011: Comparison of

carbonate C and O stable isotope records across the Jurassic/

Cretaceous boundary in the Tethyan and Boreal Realms.

Palaeogeogr. Palaeoclimatol. Palaeoecol. 299, 83-96.

Appendix A — Taxonomic index in alphabetical order