GeolCarp_Vol52_No3_147

GEOLOGICA CARPATHICA, 52, 3, BRATISLAVA, JUNE 2001

147—158

MICROFOSSIL ASSEMBLAGES AS BATHYMETRIC INDICATORS

OF THE TOARCIAN/AALENIAN “FLECKENMERGEL”-FACIES

IN THE CARPATHIAN PIENINY KLIPPEN BELT

JAROSŁAW TYSZKA

Institute of Geological Sciences, Cracow Research Centre, Polish Academy of Sciences, ul. Senacka 1, 31-002 Kraków,

Poland; ndtyszka@cyf-kr.edu.pl

(Manuscript received November 30, 2000; accepted in revised form March 15, 2001)

Abstract: Cluster and principal component analyses of Toarcian-Aalenian microfossils from a uniform Fleckenmergel
facies, found in two different parts of the Pieniny Klippen Basin have been undertaken on the basis of the percentage
abundance of benthic foraminifers and the standardized number of all microfossils per 100-grams of sediment. The latter
type of input data appeared to separate two paleobathymetric settings much better than relative benthic foraminiferal
data alone. Comparison of the foraminiferal assemblages with fossil and recent analogues suggests not so strong paleodepth
differences within the basin ranging from the middle/outer neritic for the Czorsztyn and the outer neritic and/or upper
bathyal for the Branisko Zone. The shallower part of the basin is dominated by Spirillina associated with other epibenthic
foraminifers, enhanced abundance of ostracods, crinoid and other echinoderm fragments. Higher abundance of crinoid
fragments and sponge spicules within the shallower (Czorsztyn) zone suggests a slightly higher energetic environment
with very weak bottom currents. Predominance of nodosariids associated with tubular agglutinated foraminifers
(astrorhizids), as well as lower abundance of benthic foraminifers and ostracods linked to a higher abundance of radiolaria
are supposed to indicate deeper basinal settings. These distributional trends are most likely related to trophic conditions,
and only indirectly to the paleobathymetry of the basin.

Key words: Jurassic, paleoecology, benthic foraminifers, multivariate statistics, Krempachy Marl Fm.

Introduction

For a number of years, geologists have applied various marine
paleodepth (e.g. sedimentologic, ichnologic, paleontologic, and
geochemical) proxies. However, they never give precise depths
because they usually do not directly depend on them. Benthic
foraminifers serve as an example of such a proxy, related to wa-
ter depth and water masses (e.g. Schnitker 1974; Lutze & Coul-
born  1983).  Their  distribution  is  most  strongly  influenced  by
surface  water  productivity  and  organic  carbon  flux  to  the  sea
bed (e.g. Altenbach & Sarnthein 1989). Depth in itself does not
affect the distribution of marine organisms. It seems that espe-
cially  light,  nutrients  or  food,  oxygenation,  temperature,  CO

saturation, substrate type and current activity are closely linked
to certain paleodepths, but vary from basin to basin and within a
basin. A paleobathymetric proxy should therefore be used main-
ly through covariation between depth and other factors, espe-
cially oxygen and organic flux (Van der Zwaan et al. 1999). It is
clear that paleobathymetric reconstructions should never be
based on individual marker species, but preferably on quantita-
tive assemblage characteristics (Van der Zwaan et al. 1999). The
best way to achieve that in the fossil record, is to have insight
into a multidimentional distributional pattern of fossils, apply-
ing multivariate analytical methods.

The Toarcian-Aalenian Krempachy Marl Formation (Fig. 1)

is used to test applicability of multivariate methods and poten-
tial paleobathymetric proxies. The formation is an example of
a  Fleckenmergel-type  facies,  which  lacks  any  clear  macro-
scopic  paleodepth  indications.  This  facies,  developed  in  al-
most  the  whole  Pieniny  Klippen  Basin  during  that  time
(Birkenmajer 1977), was widespread in the Early and Middle

Jurassic Tethys. It is interpreted as basinal deposits sedimented
in  poorly  ventilated,  semi-enclosed  troughs  developed  under
extensional tectonic regime (Bernoulli & Jenkyns 1974). Dur-
ing the Callovian, the Pieniny Klippen Basin was already very
deep with high paleodepth contrasts (Fig. 1b) (after Birkenma-
jer 1977). The question is whether the studied Toarcian-Aalen-
ian basin was already as deep as later, and whether its architec-
ture  was  so  differentiated.  I  will  concentrate  only  on  two
different parts of the basin, the Czorsztyn and Branisko suc-
cessions/paleobathymetric  zones,  because  these  successions
have  been  studied  in  more  detail.  There  are  several  possible
paleobathymetric  interpretations  (Fig.  2A—E):  model  A  con-
siders very shallow depths in both zones; model B shows com-
parable shallow depths with a slightly deeper Branisko Zone;
model  C  highly  differentiates  both  zones  and  resembles  the
Callovian reconstruction (Fig. 1b); model D considers compa-
rable  although  very  deep  depths  (e.g.  bathyal);  and  model  E
suggests  that  the  Czorsztyn  Zone  was  distinctly  deeper  than
the Branisko Zone. These models cannot be tested using sedi-
mentological and ichnological characteristics, thus, the micro-
fossil quantitative analysis is used here.

The aim of this study is to extract paleoecological, especial-

ly paleobathymetric signals from a relatively uniform facies
(Fleckenmergel), to test paleodepth models, and to evaluate
the applicability of selected multivariate methods.

Geological setting

The Pieniny Klippen Belt (PKB) represents the axial, heter-

ogeneous, highly deformed tectonic zone of the Western Car-

148 TYSZKA

pathians  (Fig.  1)  separating  the  Outer  Carpathian  (Magura),
and the Central Carpathian units (Birkenmajer 1977). Several
successions of facies depict different paleobathymetric zones
within  the  PKB  from  the  ‘north’  to  the  ‘south’  (Fig.  1b,c)
(Birkenmajer 1988). Uppermost Early Jurassic (Pliensbachian,
?Toarcian)  is  poorly  recorded  in  the  PKB  (Horwitz  1936;
Birkenmajer & Myczyński 1994). Aalenian and Bajocian sedi-
ments  are  better  preserved  and  represent  the  Fleckenmergel
and Fleckenkalk facies formed by grey spotty marls and lime-
stones  (Krempachy  Marl  Formation  and  Podzamcze  Lime-
stone Formation) interrupted by dark-grey to black claystones,
mudstones  and  marls  (Skrzypny  Shale  Formation  and  Har-
cygrund Shale Formation). These ”black shale” facies devel-
oped at the Aalenian/Bajocian transition record a regional an-
oxic event (Tyszka 1994b) (Fig. 1c).

The Krempachy Marl Formation is characterized by thin-

bedded grey to grey-bluish spotted marlstones and limestones.

The thickness of this unit usually varies between 10 and 30 m
(Birkenmajer  1963,  1977).  An  about  100  m  thick  sequence
from the Krempachy type locality is probably tectonically re-
peated (Birkenmajer pers. commun.). It occurs in the Czorsz-
tyn, Czertezik, Niedzica, Branisko, and probably Pieniny suc-
cessions (Fig. 1b,c). The Czorsztyn and Branisko successions
have been studied here. In the Czorsztyn Succession the age of
the  formation,  was  indicated  by  ammonites,  as  the  latest
Pliensbachian to early Aalenian (Birkenmajer 1977; Myczyńs-
ki  1973).  In  the  Branisko  Succession,  the  formation  ranges
from  the  Lower  Aalenian  to  the  lowermost  Upper  Aalenian
(Leioceras  opalinum  to  Ludwigia  murchisonae  zones)  only
(Myczyński 1973). Benthic foraminifers at least support Late
Toarcian  to  earliest  Early  Aalenian  age  documented  by  the
Falsopalmula  tenuistriata  Subzone  as  the  lower  unit  of  the
Lenticulina d’orbignyi Zone (Tyszka 1999). Spotty structures
are  interpreted  as  trace  fossils  dominated  by  Planolites,  and

Fig. 1. a – Location map with sampled localities (1—3) cropping out the Krempachy Marl Formation. 1 – Krempachy; 2 – Biała Woda; 3
– Podubocze. The grey zone schematically depicts the Pieniny Klippen Belt (PKB). b – Palinspastic reconstruction of the Pieniny Klippen
Basin during the Callovian with positions of stratigraphic/facial successions representing different paleobathymetric zones (Birkenmajer
1977, modified and simplified). c – Lithostratigraphy of Early-Middle Jurassic rocks in the Pieniny Klippen Belt (adapted from Birkenma-
jer 1977; modified).

MICROFOSSIL ASSEMBLAGES AS BATHYMETRIC INDICATORS 149

Chondrites associated with Thalassinoides and very rare Zoo-
phycos and Teichichnus. Maximum burrow diameters (MBD)
vary from 5—12 mm, usually 6—8 mm, in the Czorsztyn Succes-
sion, and 4—8 mm in the Branisko Succession. The rock con-
sists of abundant filaments of the bivalve Bositra buchi.

Our knowledge of the microfaunal composition of this for-

mation  is  limited.  Birkenmajer  &  Pazdro  (1963)  discovered
numerous ostracods and foraminifers dominated by lenticulin-
ids and associated with Eoguttulina liasica Strickland and two
species  of  Reinholdella  within  the  ”Opalinus  Beds”  (=The
Krempachy  Marl  Formation  of  Birkenmajer  1977)  of  the
Niedzica Succession. Scheibnerová (1968) figured fifteen fora-
miniferal taxa and four ostracod taxa from a single locality of
the Czorsztyn Succession in the Orava valley (Slovakia).

Material and methods

Eighteen samples have been collected from three localities:

Krempachy (Czorsztyn Succession), Biała Woda (Czorsztyn
Succession), and Podubocze (Branisko Succession), (Fig. 1;
App. 1; for exact sample locations see Tyszka 1995, Figs. 6, 7).
They represent the best accessible outcrops of the Krempachy
Marl Formation in Poland (recently Podubocze was flooded by
an artificial lake). The samples collected were dried, weighed
out (250 g), and then disintegrated in a sodium solution. Disin-

tegrated samples were sieved through a 105 µm mesh sieve.
The MVSP Version 3.0” (Kovach 1998) was used for the clus-
ter analysis (CLA) and principal component analysis (PCA)
of quantitative microfossil data.

Data sets.

Two types of quantitative data sets were used in

both analyses: (1) percentages of all benthic foraminiferal
taxa, and (2) abundance of all benthic microfossils per 100 g
sediment, including the number of foraminifers per 100 g sed-
iment and an ostracod valves/carapaces ratio. In order to avoid
overestimation of ostracod tests, ostracod number (N) has
been recalculated using a formula, where C represents the car-
apaces number, and V the valves number: N = C + V/2.

PCA recommends an equal or lower amount of variables

than cases. In order to fulfil this requirement, the number of
variables has been reduced to eighteen. Most common micro-
fossils are kept separately and directly transferred to input
files (see App. 2 and 3). Rare taxa or groups, which show tax-
onomic or morphological affinity, have been added together
according to codes presented within the data tables (see App.
2 and 3).

Cluster analysis.

CLA offers a set of numerical techniques,

which divide the object of study into discrete groups usually
presented as dendrograms. Here, the clustering, based on the
cosine theta distance, also known as the normalized Euclidean
distance, links every pair of cases/clusters with the lowest
distance.

∑













−

In these two formulae, C represents cosine theta distance, n

gives the total number of variables, i and j represent two rows
(cases) of the data matrix, k represents the column (variable),
and therefore x

would be the datum in the k

column of row

i. The cluster formed by two cases is considered a single ob-
ject, which is compared and linked to other objects (subclus-
ters). The Unweighted Pair Group Average method (UPGMA)
is used, which employs the average linkage technique. In this
method, the average point (centroid) is closer to the group
with more points (Kovach 1998). Q- and R-mode analyses
have been conducted.

Principal component analyses.

PCA remains one of the

most  widely  used  ordination  techniques  in  paleoecology
(Spicer & Hill 1979). The aim of PCA is to find meaningful
axes  in  the  multidimensional  space  defined  by  the  variable
(taxon abundance). The result of an analysis is to plot the PC-
loadings against usually the first two or three principal axes in
the form of a scatter plot. By performing either an R-mode or
a  Q-mode  PCA  both  variable  and  sample  scores  can  be  ob-
tained, which creates a dual ordination of the original data ma-
trix.  PCA  was  invented  to  present  a  continuum  that  may
sometimes be assigned to an environmental gradient. Never-
theless, it also detects discontinuities, and thereby groups the
data. The clustering on scatter plots (biplots) may then be used
as a basis for dividing the sampled fossils into assemblages,
each exhibiting a more homogeneous structure (Spicer & Hill
1979).  It  means  that  variable  points  in  close  proximity  will
show the correlation between variables, which may be inter-
preted  as  a  result  of  the  same  paleoecological,  sedimentary,

Fig. 2. Possible hypothetical paleodepth models A—E of the studied
paleobathymetric zones within the Pieniny Klippen Basin during
the Toarcian—Aalenian (discussed in this paper).

where

∑

150 TYSZKA

and/or taphonomic process. The variables may not all be di-
rectly comparable, so it would seem necessary to convert all of
them to standardized form (Davis 1986). In order to avoid spe-
cies abundance and sedimentation rate problems standardized
data are used here.

Results

Biofacies/Microfossils.

Quantitative microfossil data are

presented in App. 2 and 3. Preservation of microfossils is simi-
lar in both studied successions. The biofacies of the formation
within the Czorsztyn Succession is characterized by abundant
benthic foraminifers and ostracods, common echinoderm frag-
ments, including frequent crinoid remains and echinoid spines,
and  sponge  sclerites  (triactines,  monactines,  hexactines,  and
more  complex  ones)  (Fig.  3).  Besides  benthics,  relatively
abundant  radiolaria  (1000—10000  specimens  per  250  g-sam-
ple)  have  also  been  found.  In  general,  benthic  foraminifers
from  the  Czorsztyn  Succession  are  more  abundant  than  in
Branisko. They are dominated by calcareous foraminifers, es-
pecially Spirillina (25.4—52.2 %; mean 42.5 %), Laevidentali-
na (12—24 %) and Lenticulina (9—22 %). Patellina and Rhein-
holdella are rare (up to 5 %) but quite characteristic. Elongated
morphogroups  depict  various  species  of  Laevidentalina,
Pseudonodosaria,  and  Eoguttulina.  Lenticulina  constitutes  9
to 22 %. Other nodosariids are rare and compose no more than
4 %. Hyperammina, Ammobaculites, Trochammina, Reophax
and adherent Tolypammina represent scarce agglutinated fora-
minifers. Smooth-carapace ostracods are quite abundant (200—
350 individuals per 100 g sediment).

Laevidentalina (22—56 %), Lenticulina (9—47 %), and Spir-

illina (1—46 %, 21.4 % on average) predominate in the Branis-
ko  Succession.  Samples  enhanced  in  Lenticulina  exhibit  the
lowest  number  of  Spirillina.  Elongated  nodosariids  are  rela-
tively more frequent than in the Czorsztyn Succession. Patelli-
na  and Rheinholdella  do  not  occur  at  all.  Agglutinated  fora-
minifers show a slightly higher number of tubular forms, such
as Hyperammina, Rhabdammina. Ostracods are less abundant
(100—200 valves and carapaces per 100 g sediment) than in the
Czorsztyn  Succession.  Scarce  sponge  spicules  and  echino-
derm fragments exhibit the same pattern. In contrast, radiolaria
(preserved as calcitic casts) are more abundant in the Branisko
than in the Czorsztyn Succession.

Microfaunal association described by Scheibnerová (1968)

from the same ‘formation’ (so-called ”Opalinus Beds”) of the
Czorsztyn Succession in the Orava valley shows 70 % of fora-
minifers and 30 % of smooth-walled ostracods that is similar
to our results. Nodosariids are also very abundant with a domi-
nation  of  elongated  forms,  such  as  Frondicularia  sulcata
Bornemann, Nodosaria fontinensis Terquem, Dentalina vetus-
tissima d’Orbigny. The lack of spirillinids and high proportion
of agglutinated foraminifers (up to 40 %) dominated by Am-
modiscus  incertus  (d’Orbigny)  and  Trochammina  inflata
(Montagu) are surprising. Especially Ammodiscus “forms the
essential part of the association” (Scheibnerová 1968). Actual-
ly,  agglutinated  tubular  and  planispiral  Ammodiscus  is  mor-
phologically analogous to calcareous Spirillina. Our forms are
calcareous  and  do  not  comprise  agglutinated  tests  (Fig.  3).
There  are  several  possible  explanations  of  such  differences,

but they cannot be tested without comparative studies of the
Orava valley microfauna.

Cluster analysis.

The analysis of percentage foraminiferal

data separates variables (benthic foraminiferal taxa) into two
clusters, which represent different assemblages (Fig. 4a). Pa-
tellina,  Reinholdella,  Spirillina,  Eoguttulina,  ophthalmidiids,
and Pseudonodosaria are most closely linked within the first
cluster. All of them represent calcareous foraminifers and, ex-
cept  for  Pseudonodosaria,  do  not  belong  to  the  nodosariids.
However, nodosariids clearly predominate in the second, very
wide cluster, which links some agglutinated foraminifers, such
as  tubular  astrorhizids,  Trochammina,  Ammobaculites,  and
Subreophax. Sample clustering reveals two main clusters sur-
prisingly  linking  all  the  Czorsztyn  Succession  samples  with
selected Branisko Succession samples (Fig. 4b). The Czorsz-
tyn Succession samples representing the same locality (Biała
Woda: BK-1, -2, -3, -11) are most closely associated within a
single subcluster, which is associated with a Branisko Succes-
sion sample (PDK-17).

The cluster analysis of standardized abundance data, includ-

ing all benthic microfossils per 100 g sediment and an ostra-
cod valves/carapaces ratio, shows a different pattern. Variables
have revealed two main loosely linked clusters (Fig. 5a). One
of them is separated into two subclusters, where Patellina, Re-
inholdella,  Spirillina  (including  rare  ophthalmidiids),  Eogut-
tulina are closely associated with abundant foraminifers, ostra-
cods,  crinoid  and  other  echinoid  fragments.  These  closely
linked  variables  represent  well-separated  microfossil  assem-
blage pronounced within samples from the Czorsztyn Succes-
sion. Other subclusters are not so distinct, loosely linking as-
trorhizids  with  nearly  all  elongated  nodosariids,  and  other
agglutinated foraminifers with Nodosaria regularis and Ram-
ulina, holothurian sclerites, and valves/carapaces ratios. These
variables  characterize  the  microfossil  assemblage  fitting  into
samples from the Branisko Succession. Clustering of samples
clearly separates the Czorsztyn Succession samples from the
Branisko Succession ones (Fig. 5b).

Principal component analyses.

The same procedure with

the input data has been applied. In this case, non-standardized
percentage  values  bring  about  domination  of  most  common
taxa, such as Spirillina, Laevidentalina, and Lenticulina, and
to  some  extent  Pseudonodosaria  and  Eoguttulina  (Fig.  6a).
The  distribution  of  these  taxa  is  responsible  for  most  of  the
variability within the data set. The first axis accounts for about
66 % of the variance, and the first three axes for 95 % of the
variance. The first vs. the second axis plot separates four clus-
ters  of  samples  representing  either  mixed  Czorsztyn  with
Branisko  successions  or  just  the  Branisko  Succession  alone.
Samples from the Biała Woda Valley (BK) are most closely
linked due to similar proportions of Spirillina, Laevidentalina,
and Lenticulina. These samples have shown the highest load-
ings on the first axis and medium values on the second axis.
They  also  link  a  single  Branisko  Succession  sample,  with  a
high proportion of Spirillina (Fig. 6a). In fact, the high and sta-
ble relative abundance of Spirillina is mostly responsible for
such variability along the first axis. The second cluster, which
contains some Czorsztyn Succession (from the Krempachy lo-
cality) and Branisko Succession samples, shows medium val-
ues on both axes. Another two clusters spread along the sec-
ond axis. The highest second axis loadings are characteristic

MICROFOSSIL ASSEMBLAGES AS BATHYMETRIC INDICATORS 151

Fig. 3. Benthic foraminifers (1—16) and selected microfossils (17—21) from the Krempachy Marl Formation. Scale bar = 100

m. 1 – Spir-

illina elongata Bielecka & Pożaryski, spiral view, sample PDK-17. 2 – Spirillina infima Strickland, spiral view, sample PDK-15. 3 – Pa-
tellina sp., peripheral view; sample BK-4. 4 – Morphogroup C-2, Ramulina spandeli Paalzow, sample PDK-18. 5 – Nodosaria regularis,
sample PDK-2. 6 – Pyramidulina columnaris (Franke), sample PDK-10. 7 – Pyramidulina dispar (Franke), sample PDK-1. 8 – Pseudon-
odosaria bajociana (Terquem), sample PDK-1. 9 – Laevidentalina subplana Franke, sample PDK-18. 10 – Laevidentalina sp., sample
PDK-11. 11 – Astacolus anceps (Terquem), sample PDK-18. 12 – Falsopalmula deslongchampsi (Terquem), sample PDK-10. 13 – Fal-
sopalmula tenuistriata (Franke), sample PDK-10. 14 – Marginulinopsis dictyodes dictyodes (Deecke), sample PDK-10. 15 – Uncoiled
Lenticulina d‘orbignyi (Roemer) sample PDK-10. 16 – Lenticulina polygonata (Franke), sample PDK-18. 17 – Sponge macroscler; sam-
ple PDK-2. 18 – Ornamented ostracod, ? Lophocythere sp., left valve, sample PDK-2. 19 – Echinoid spine, KSK-6. 20—21 – Radiolaria,
(20) sample PDK-2; (21) sample PDK-18.

152 TYSZKA

different  successions  than  relative  foraminiferal  data,  which
depict  higher  variability  within  successions  than  variability
between the two successions (Figs. 4—6). It suggests that fora-
miniferal  assemblages  reveal  close  affinities  in  both  succes-
sions and points to not too strong paleoenvironmental differ-
ences. On the other hand, percentage values always close the
data, changing their structure. Any highly fluctuating variable
causes fluctuations of other, more stable components. For in-
stance, analyses of percentage data indicate closer affinity of
relatively  abundant  Lenticulina  to  agglutinated  foraminifers
and most of the nodosariids (Figs. 4, 6a). The results based on
abundance data show an opposite trend, that is closer associa-
tion with Patellina, Spirillina and other microfossils (Figs. 5,
6b). Comparison of Spirillina vs. Lenticulina distribution pre-
sented in percentages and abundances (per 100 g sediment) ex-
plains this contradiction (App. 2 and 3). Percentage data reveal
distinctly  higher  proportions  of  Lenticulina  within  some
Branisko  samples  (see  Appendix  2;  e.g.  PDK-1,  -2,  -3,  -5).
Abundance  data  give  a  comparable  number  of  Lenticulina
specimens in both successions with slightly higher values for
the Czorsztyn Succession samples (App. 2). This distribution-
al pattern becomes clear when comparing values for Spirillina,
which  are  on  average  much  higher  in  the  Czorsztyn  Succes-

for Laevidentalina dominated samples, the lowest of which are
associated with high proportions of Lenticulina (samples
PDK-1, PDK-5). The third axis does not reveal any distinct
pattern (App. 4).

Analysis of standardized abundance of all microfossils

clearly  separates  two  microfossil  assemblages  deriving  from
both successions (Fig. 6b,c). Five factors (axes) are significant
and explain 85.1 % of the variance within the data set. The for-
aminiferal  abundance,  Eogutullina,  Spirillina,  ostracod,  and
crinoid  fragment  abundance  (including  other  echinoderm  re-
mains) show the highest (> 0.3) loadings for Axis 1, which ac-
counts for 41.5 % of the variance. Axis 2 depicts the highest
loadings (0.3—0.5) for all agglutinated foraminifers, calcareous
Nodosaria  regularis  +  Ramulina,  and  Laevidentalina.  This
axis explains only 16.1 % of the variance. Axis 3 shows the
highest loadings for Patellina and Reinholdella and the most
negative for sponge spicules and echinoid spines (App. 5).

Interpretation and discussion

Comparison of multivariate analytical results.

Analyses of

standardized abundance data better separate samples from two

Fig. 5. Dendrogram from cluster analysis based on abundance of all
microfossils (per 100 g sediment) from the Krempachy Marl Forma-
tion (for other notes see Fig. 5). a – Clustering by variable (taxon/
groups) with a key for variables from Fig. 6b. b – Clustering by
sample.

Fig. 4. Dendrogram from cluster analysis based on percentage abun-
dance of benthic foraminifers from the Krempachy Marl Formation.
Distance is measured by the cosine theta coefficient. Unweighted
method of average linkage (UPGMA) is used. a – Clustering by
variable (taxon/groups). b – Clustering by sample.

MICROFOSSIL ASSEMBLAGES AS BATHYMETRIC INDICATORS 153

sion creating an artefact of a relatively lower proportions of
Lenticulina. Thus, Lenticulina distribution is stable in both
successions and does not distinctly associate with any succes-
sion.

Paleobathymetry

. The formation studied reveals benthic

foraminiferal assemblages dominated by calcareous foramini-
fers, which suggest paleodepths fairly above the CCD (calcite
compensation depth) and exclude deep bathyal or abyssal
depths for both the Czorsztyn and Branisko paleobathymetric

zones. This rules out hypothetical paleobathymetric models C
and D (Fig. 2). Nevertheless, the studied zones somewhat dif-
fer in the composition of their microfauna, which probably in-
dicates paleoenvironmental and also paleobathymetric differ-
ences. All these biofacies differences are compiled in Table 1.
High abundance of planispiral Spirillina is the most character-
istic feature of the whole formation. This epibenthic foramini-
fer is clearly dominant in the Czorsztyn Succession deposits,
but also relatively common in those of the Branisko Succes-
sion. Multivariate analyses indicate that Spirillina is closely
associated with other epibenthic forms, such as Patellina and
some epistominids (Reinholdella), as well as with some elon-
gated, probably endobenthic forms, like Eoguttulina (see Figs.
4—6). The distribution of these taxa also shows close affinity to
other variables, that is to the total benthic foraminiferal abun-
dance, mostly contributed by much higher abundance of Spir-
illina, as well as, to the abundance of ostracods, crinoid frag-
ments, other echinoderm remains, and sponge spicules.

Gordon (1970) interpreted floods of Jurassic Spirillina as in-

dicative of shallowing. Abundant Spirillina infima was report-
ed  by  Morris  (1982)  from  shallow  water  coralliferous  marls
and  bioturbated  limestones  associated  with  Middle  Jurassic
oolitic  facies  of  Wales.  He  considered  this  taxon  to  form  a
”primary weed fauna” associated with the filamentous algae.
Nagy  (1992)  reported  spirillinids  from  the  Yons  Nab  Beds
(Bajocian, North Sea deltas) reflecting ”a shallow marine en-
vironment,  presumably  within  the  photic  zone”.  Samson
(1997)  summarized  Jurassic  Spirillinidae  as  a  characteristic
group for the infralittoral super-biotope within the photic zone
(down to a. 100 m). Johnson (1977) described Spirillina infi-

Table 1: Summary of selected sedimentologic and micropaleonto-
logic features of the Krempachy Marl Formation between both
Czorsztyn and Branisko successions.

CZORSZTYN
SUCCESSION

BRANISKO
SUCCESSION

similarities

lithology

spotty marlstones and marly limestones

coloration

grey to bluish grey

trace fossils

extensive bioturbation, medium to large maximum

diameters, common Planolites & Chondrites,

frequent Thalassinoides

differences

dominating foram.
morphogroups

epibenthic forms

endobenthic forms

benthic foraminifers:

very abundant

abundant

Patellina

frequent

absent

Rheinholdella

frequent

absent

Spirillina

very abundant

common to abundant

Eoguttulina

common

single to frequent

nodosariids

frequent to common

very abundant

astrorhizids

single

single to frequent

ostracods

abundant

common

crinoid fragments

frequent

single

other echinoderm remains

abundant

frequent

sponge spicules

abundant

frequent

radiolaria

abundant

very abundant

interpretation

paleodepth

middle/outer neritic

outer neritic to upper
bathyal

oxygenation

dysoxic

trophic regime

mesotrophic

food supply

vertical (?seasonal fluxes)
and lateral (weak currents)

vertical

Fig.  6.  a  –  Principal  Component  Analysis  biplot  for  axes  1  and  2
based  on  percentage  abundance  of  benthic  foraminifers  from  the
Krempachy Marl Formation. b – PCA plots for axes 1 and 2 based
on  abundance  of  all  microfossils  (per  100  g  sediment)  from  the
Krempachy Marl Formation (key for variables can be found on Fig.
5a).  c  –  plot  of  samples  revealing  two  clusters  separating  Czorsz-
tyn and Branisko successions.

154 TYSZKA

ma and Reinholdella from the inner to the outer neritic (Toar-
cian-Domerian  of  Wales).  Herrero  (1993)  reported  Toarcian
outer neritic assemblages from the Cordillera Iberica dominat-
ed by the Lagenina (Vaginulinidae family) subordinately asso-
ciated  with  Spirillinina.  Monaco  et  al.  (1994)  described  the
Aalenian Spirillina Unit (up to 50 % of spirillinids associated
with nodosariids) from Central Italy and interpreted as middle
shelf with regressive trends. Spirillinids also dominate within
Oxfordian neritic biohermal sponge facies from southern Po-
land  and  the  French  Jura  (Barwicz-Piskorz  1989;  Gaillard
1984  cited  by  Barwicz-Piskorz  1989),  as  well  as,  within  the
shallow  water  Kimmeridgian  marlstones  of  central  Poland
(Barwicz-Piskorz  &  Tarkowski  1984).  Patellina  and  Rein-
holdella  have  usually  been  reported  from  shallow  neritic
depths (Gordon 1970; Gradstein 1983; Stam 1986; Copestake
&  Johnson  1989;  Riegraf  &  Luterbacher  1989).  Jurassic
Eoguttulina may also indicate a relatively shallow marine en-
vironment (Brouwer 1969; Birkenmajer & Tyszka 1996). On
the other hand, Gradstein (1983) reported Jurassic assemblag-
es dominated by nodosariids and some small agglutinated for-
aminifers  associated  (but  not  dominated!)  with  rare  to  com-
mon  Spirillina  and  Epistomina  from  abyssal  depths  of  the
Blake-Bahama  Basin.  Similar  deep  oceanic  (abyssal)  occur-
rences of Upper Jurassic Spirillina were described from DSDP
sites by Luterbacher (1972), Bartenstein (1974), and Kuzniets-
ova (1974). According to Riegraf & Luterbacher (1989), the
Upper  Jurassic  ”deep  water  fauna”  in  the  DSDP  boreholes
contains mostly nodosariids and small agglutinated forms as-
sociated with variable amount of epistominids, Ophthalmidi-
um, and Spirillina.

Recent analogues are given by Davies (1970) and Brasier

(1975) who described Spirillina on algae in modern lagoons.
Berthold (1976), as well as Kitazato (1988) included Spirillina
vivipara Ehrenberg and Patellina corrugata Williamson to va-
grant phytal forms mainly living on seaweeds. Seaweeds re-
quire light that may govern distribution of these epiphytal for-
aminifers (Kitazato 1988; Murray 1991).

In conclusion, Spirillina and related taxa were reported from

nearly  all  depths,  but  assemblages  dominated  by  these  taxa
tend to indicate shallow settings, usually the inner to middle
neritic. Thus, all above references suggest that the Patellina,
Spirillina  and  associated  microfossils,  deriving  from  the
Czorsztyn  Succession,  indicate  relatively  shallow,  open  sea
paleoenvironment,  which  may  be  placed  around  the  middle
neritic zone. This is also supported by their association with
abundant, poorly sorted, and morphologically diverse echino-
derm  (mainly  crinoidal)  remains  which  were  probably  not
transported from other distant areas. Głuchowski (1987) inter-
preted  optimum  conditions  for  Middle  Jurassic  crinoid  com-
munities as comparatively shallow, below the wave base, but
still within the photic zone.

Relatively low differences in the faunal composition be-

tween two zones point out quite low paleobathymetric differ-
ences. Slightly enhanced proportions of astrorhizids, domina-
tion of nodosariids, a lower number of crinoidal fragments,
and a higher abundance of radiolaria suggest deeper location
of the Branisko Zone and exclude a further two tested paleo-
bathymetric models, that is models A and E (Fig. 4). Nodosa-
riids from the Oxford Clay of England have already been men-
tioned as indicators of deeper (epicontinental) water, even if

they are not restricted to it (Barnard et al. 1981). Foraminiferal
assemblages of the comparable “Fleckenmergel” facies from
the Branisko Succession have already been analysed (Tyszka
1994a).  Those  assemblages  are  dominated  either  by  aggluti-
nated tubular foraminifers (astrorhizids) or nodosariids associ-
ated  with  astrorhizids.  They  indicate  much  deeper  settings,
however  above  the  CCD,  identified  as  the  upper  to  lower
bathyal. Furthermore, it seems that Lenticulina alone is proba-
bly not a good paleodepth proxy. Jurassic assemblages domi-
nated  by  Lenticulina  have  been  identified  from  all  depths
above  the  CCD  (see  Morris  1982;  Gradstein  1983;  Tyszka
1994a,b).

All above references and comments do not suggest any

strong paleobathymetric variability. The Czorsztyn paleo-
bathymetric zone can be estimated within the inner to middle
neritic. The Branisko Zone was probably slightly deeper and
ranged around the outer neritic or the uppermost bathyal.
Thus, model B is the most probable reconstruction of the Toar-
cian-Aalenian Klippen Basin (Fig. 2B).

Trophic and oxic regime.

Dense bioturbations, a high pro-

portion  of  endobenthic  foraminifers,  and  common  radiolaria
do not indicate any strong shortage of nutrients and food. All
studied sections have revealed quite monotonous trace fossil
assemblages with Thalassinoides, Planolites, and Chondrites.
Such  ichnofossil  assemblages  are  typical  of  low  oxygenated
environments  (Bromley  &  Ekdale  1984;  Savrda  &  Bottjer
1986; Tyszka 1994a). Relatively high maximum burrow sizes,
ranging  between  6  and  12  mm,  a  common  occurrence  of
Thalassinoides,  and  appearance  of  skeletal  fossils  such  as
echinoderm and holothurian remains indicate moderate dysox-
ic conditions of bottom water and surficial part of the substra-
tum. Long-term temporal and spatial changes in bottom water
oxygenation are not recorded by trace fossils and macrofaunal
skeletal  fragments.  Actually,  foraminiferal  morphogroups
show a spatial variability pattern, that is the Czorsztyn Zone is
dominated  by  epibenthic  forms  (mostly  spirillinids),  and  the
Branisko Zone by endobenthic forms (elongated nodosariids).
The TROX of Jorissen et al. (1995) and TROX-2 of Van der
Zwaan et al. (1999) models and some earlier references imply
either control by elevated redox boundary or shortage of nutri-
tions as controlling factors of epibenthic dominance (see Van
der Zwaan et al. 1999 for refs.). Such a more elevated position
of the redox boundary in a shallower setting would be the easi-
est explanation, because organic flux is usually higher at shal-
lower depths, and a higher organic matter content causes oxy-
gen utilization and rising of the redox boundary (e.g. Van der
Zwaan  et  al.  1999).  Deep  and  complex  tierings  of  burrows
throughout all successions do not support such an explanation.

Stam (1986) assumed that the distribution and abundance of

spirillinids might depend not so much on the amount of food
but  more  on  the  quality  of  food.  It  can  be  speculated  that
epibenthic  spirillinids  preferred  mesotrophic  conditions  with
episodical,  probably  seasonal  fluxes  of  food  related  to  phy-
toplankton blooms. Spirillinids as surface dwellers were prob-
ably best suited for fast response and reproduction after sea-
sonal organic matter influxes. This seasonal scenario seems to
be  supported  by  the  high  dominance  of  Spirillina  associated
with the enhanced abundance (per 100 gram sediment) of all
benthic  foraminifers.  Such  averaged  fossil  assemblage  may
represent a very dynamic community with a highly fluctuating

MICROFOSSIL ASSEMBLAGES AS BATHYMETRIC INDICATORS 155

population size. The community may have been highly over-
dominated by spirillinids during high organic fluxes. Periods
with a lower organic matter influx were probably favoured by
endobenthic forms and specialized epibenthos with very low
proportions  of  spirillinids.  On  average  in  the  fossil  record,
these  communities  show  averaged  proportions  oscillating
around 50 % in the Czorsztyn and 20 % in the Branisko zones.
This lower proportion of Spirillina in the Branisko Zone prob-
ably reflects a lower seasonal organic matter influx due to re-
cycling  of  organic  matter  during  sinking  through  the  deeper
water column and/or due to lower primary productivity.

Abundance of ostracods follows trends in abundance of

benthic foraminifers. Their higher abundance in the Czorsztyn
Succession may indicate either a higher productivity or a low-
er sedimentation rate, which caused condensation of microfos-
sils. Considering discussion presented below, strong differenc-
es in the sedimentation rate between the two successions are
doubtful. The ostracods/foraminifers ratio reaches on average
0.45, but varies between 0.23 and 0.92 that is comparable to
the highest values recorded in the moderately to weakly dys-
aerobic biofacies of the Podzamcze Limestone Formation, but
much higher than the lowest values (0.01—0.05) characteristic
for strongly dysaerobic biofacies (Tyszka 1994a). It means
that, in general, ostracods were less resistant to suboxic condi-
tions than benthic foraminifers. Their valves/carapaces ratio
shows high variability and seems to be enhanced in the Branis-
ko Succession, but, actually, this pattern is caused by two un-
usually high values (App. 2).

The Czorsztyn Succession preserved an enhanced abun-

dance of crinoid fragments and sponge spicules (especially at
the Krempachy locality), therefore, skeletal fragments of sus-
pension feeders. It indicates weak bottom currents carrying
nutritients, essential for their survival. On the other hand, con-
ditions were far from optimal for densely populated crinoidal
meadows, which developed in the shallower part of the basin
later, that is during the Bajocian and Bathonian (see Birken-
majer 1963; Głuchowski 1987). It is likely that stagnation of
the basin and bottom water oxygen-depletion did not favour
crinoidal meadows.

Sedimentation rate.

It is important to be aware of suscepti-

bility to species abundance when abundances of fossils within
unit volumes of rock are ordinated in PCA. In this case, any
variation in abundance arising from varying rates of sedimen-
tation will be introduced into the final pattern (Spicer & Hill
1979). If variability of sedimentation rate was the only reason
for changes in microfossil abundance, one could expect stable
(constant)  proportions  of  microfossils.  The  negative  correla-
tion of radiolaria vs. foraminifera abundance in the Czorsztyn
and Branisko successions suggests that sedimentation rate dur-
ing deposition of the Krempachy Marl Formation was not so
different  in  the  two  successions.  It  also  means  that  the  en-
hanced abundance of foraminifers associated with shallow wa-
ter indicators is not just an artefact of a lower sedimentation
rate in the Czorsztyn Succession. The foraminiferal productiv-
ity was most likely higher in the shallower part of the basin,
possibly due to higher (?seasonal) food fluxes. Unfortunately,
absolute sedimentation rates cannot be calculated due to poor
time  control  and  tectonically  obscured  stratigraphic  bound-
aries. On the basis of the duration of ammonite zones and re-
ported thickness of the formation, it can be roughly estimated

that the sedimentation rate in both successions ranged between
several up to a dozen or so millimetres per ka.

Conclusions

The results of multivariate analyses based on percentage

abundance of benthic foraminifers and standardized number of
all microfossils per 100 gram sediment have revealed that the
latter data appear to separate paleobathymetric settings much
better than the percentage of benthic foraminiferal data alone
(Figs. 4—6). It is therefore worthwhile to extend quantitative
study to all microfossils and include them into multivariate
statistics.

Comparison of the foraminiferal assemblages with fossil

and recent analogues suggests relatively small paleodepth dif-
ferences within the basin estimated at the middle/outer neritic
for the Czorsztyn and the outer neritic and/or upper bathyal for
the  Branisko  paleobathymetric  zone.  Spirillina  is  the  most
dominant  constituent  of  the  foraminiferal  assemblage  in  the
shallower part of the basin being still frequent or even com-
mon in the Branisko Zone. Association of abundant Spirillina
with  Patellina  and  Reiholdella  indicates  shallow  neritic
depths. The predominance of nodosariids associated with tu-
bular agglutinated foraminifers (astrorhizids) suggests deeper
basinal  settings.  Enhanced  abundance  of  ostracods,  crinoid,
and other echinoderm fragments appears to be a shallow water
indicator. On the other hand, higher abundance of crinoid frag-
ments  and  sponge  spicules  within  the  shallower  (Czorsztyn)
zone points to a higher energetic environment with very weak
lateral currents.

All these proxies depend on organic matter flux. The benthic

association suggests mesotrophic conditions with moderate or-
ganic matter influx. Moderate primary productivity is also in-
dicated by relatively abundant radiolaria, which are very sensi-
tive paleoproductivity proxies. The possibility of seasonal
fluctuations on primary productivity and their effect on a fora-
miniferal distributional pattern is also speculated.

Acknowledgments: The author wishes to thank Prof. Dr K.
Birkenmajer for critical reading of the manuscript and sugges-
tions. RNDr. Katarína Holcová and Prof. RNDr. Milan Mišík
are gratefully acknowledged for reviews.

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MICROFOSSIL ASSEMBLAGES AS BATHYMETRIC INDICATORS 157

Appendix 1. Locations: Krempachy (Czorsztyn Succession). This Krempachy Marl Formation type locality is poorly outcropped on the right
bank of the Białka River, south of the Kramnica Klippe (Birkenmajer 1963) (Fig. 1a). According to Birkenmajer (1963, 1977), about 100 m thick
sequence was accessible before. At present, just a small, one meter thick, ditch dug provided access to a few samples (KSK-2, KSK-6). Out-
croped deposits represent grey to grey-bluish spotty marly limestones intercalated by similar grey spotted shaly marlstones.

Biała Woda (Czorsztyn Succession)

. The formation crops out in the stream-bed (left bank) of the Biała Woda Valley, to the east of the Smolegowa

Skała klippe (Fig. 1a). The locality exposes about 1.5 m thick profile of the bluish grey marly limestones (10—15 cm thick beds) intercalated with
grey marlstones (5—10 cm thick). Beds dip in 194/64. Four samples (BK-1, BK-2, BK-3, BK-11) have been analysed.

Podubocze (Branisko Succession)

. The outcrop was exposed to the east of the Czorsztyn Castle (Fig. 1a), in a small ENE-WSW valley (recently

flooded by an artificial lake). A 6—8 m thick section was outcropped along the valley, on a distance approximately 40 m. Thin (8—15 cm) layers of
grey and grey-bluish spotted limestones are intercalated with thin bands of spotted marly limestones and marlstones. Beds dip in 180/19, 161/22, and
157/34 and are most likely tectonically overturned as is indicated by trace-fossil structures. Twelve samples (PDK-) have been analysed (App. 2 and 3).

Appendices

Appendix 2. Abundance of all identified microfossils per 100 g sediment. The second column depicts the ‘code’ which is applied into row input
data to reduce the number of variables for multivariate statistical analyses: (*) rows used directly as input data; (a—f) sum of variables from se-
lected rows, including as follow: (a) other agglutinated foraminifers (except for astrorhizids); (b) Lenticulina with Astacolus; (c) other nodosari-
ids; (d) Ramulina with Nodosaria regularis; (e) Spirillina with ophthalmidiids (planispiral, with tubular chambers); (f) crinoid and other echino-
derm remains (excluding echinoid spines).

Succession

CZORSZTYN

BRANISKO

cod

-1

-2

-3

-1

KSK

-2

KSK

-6

-1

-2

-3

-5

-7

-8

-1

foram.abund./100g

750 1240

742

637

485

624

289

187

258

213

346

672

438

418

198

392

426

160

Ammobaculites

6.4

1.6

9.6

3.2

4.8

1.6

0.8

astrorhizids:

11.2

19.2

9.6

3.2

22.4

17.6

14.4

3.2

22.4

30.4

22.4

38.4

4.8

22.4

4.8

Conotrochammina

1.6

Evolutinella

3.2

1.6

Reophax

9.6

1.6

3.2

Subreophax

1.6

4.8

1.6

4.8

Textularia

1.6

Tolypammina

4.8

1.6

Trochammina

6.4

3.2

6.4

3.2

1.6

3.2

4.8

9.6

4.8

2.4

Verneulinoides

2.4

3.2

Astacolus

35.2

28.8

22.4

9.6

19.2

10.4

6.4

3.2

14.4

25.6

14.4

12.8

11.2

Citharina

3.2

0.8

6.4

3.2

1.6

3.2

1.6

Eoguttulina

20.8

67.2

19.2

41.6

0.8

1.6

3.2

12.8

4.8

1.6

0.8

Falsopalmula

3.2

6.4

1.6

3.2

11.2

4.8

6.4

Frondicularia

4.8

3.2

1.6

3.2

6.4

12.8

1.6

0.8

Laevidentalina

99.2

213

112

59.2

147

59.2

65.6

57.6

106

378

166

160

83.2

115

94.4

Lenticulina

118

114

86.4

89.6

134

107

43.2

70.4

99.2

57.6

76.8

28.8

73.6

68.8

31.2

Lingulina

1.6

6.4

1.6

Marginulina

2.4

Marginulinopsis

4.8

3.2

1.6

9.6

3.2

1.6

3.2

1.6

3.2

2.4

Nodosaria

3.2

4.8

19.2

4.8

6.4

4.8

3.2

2.4

Nodosaria regularis

4.8

12.8

3.2

1.6

3.2

6.4

4.8

1.6

22.4

4.8

ophthalmidiids

4.8

3.2

6.4

3.2

1.6

Patellina

12.8

57.6

38.4

12.8

1.6

Planularia

5.6

6.4

1.6

Pseudonodosaria

12.8

38.4

6.4

9.6

83.2

6.4

3.2

9.6

14.4

3.2

4.8

1.6

0.8

Pyramidulina

6.4

9.6

3.2

6.4

3.2

9.6

6.4

1.6

4.8

0.8

Ramulina

3.2

1.6

6.4

3.2

1.6

3.2

9.6

6.4

1.6

5.6

Reinholdella

9.6

38.4

41.6

28.8

3.2

Saracenaria

0.61

1.29

0.8

Spirillina

373

602

387

323

123

176

30.4

81.6

25.6

134

112

73.6

43.2

41.6

114

195

27.2

Vaginulina

1.6

3.2

4.8

1.6

4.8

1.6

0.8

Vaginulinopsis

6.4

3.2

4.8

22.4

11.2

3.2

12.8

11.2

4.8

6.4

ostracod carapaces

194

314

195

173

258

259

119

114

150

179

147

130

123

106

120

104

62.4

ostracod valves

92.8

43.2

44.8

70.4

28.8

49.6

81.6

36.8

33.6

35.2

54.4

75.2

38.4

27.2

30.4

17.6

18.4

OSTRACOD SUM

240

335

218

208

272

283

144

154

169

196

165

157

161

99.2

119

135

113

71.6

valves/carapaces ratio

0.48

0.14

0.23

0.41

0.11

0.19

0.42

0.72

0.24

0.19

0.24

0.42

0.61

0.48

0.26

0.25

0.17

0.29

SPONGE spicules

14.4

35.2

12.8

36.8

57.6

89.6

4.8

6.4

4.8

3.2

14.4

12.8

4.8

9.6

CRINOID fragments

12.8

19.2

25.6

12.8

4.8

12.8

0.8

1.6

3.2

other

ECHINODERM

frag.

65.6

123

112

35.2

70.4

0.8

11.2

28.8

6.4

11.2

3.2

1.6

ECHINOID spines

6.4

12.8

14.4

19.2

0.8

12.8

1.6

14.4

9.6

1.6

12.8

4.8

1.6

4.8

6.4

0.8

ECHINODERMS SUM

86.4

149

144

60.8

75.2

102

2.4

20.8

17.6

20.8

3.2

12.8

9.6

0.8

HOLOTHURIAN sclerites

2.4

9.6

4.8

juvenile BIVALVES

1.6

3.2

BIVALVE fragments

1.6

3.2

6.4

4.8

3.2

FISH teeth

3.2

RHYNHOLITHS

1.6

158 TYSZKA

Appendix 5. Principal components matrix for all microfossil abun-
dance (per 100 g) data.

Appendix 3. Relative proportions of all foraminifers (in percent). The second column depicts the ‘code’ which is applied into row input data: (*)
rows used directly as input data; (a—d) sum of variables from selected rows, including as follow: (a) other agglutinated foraminifers (except for
astrorhizids); (b) other nodosariids; (c) Marginulina with Marginulinopsis; (d) Vaginulina with Vaginulinopsis.

Appendix 4. Principal components structure matrix for the benthic
foraminiferal percentage data. The table presents the correlations of
the taxa to the components (axis).

Succession

CZORSZTYN

BRANISKO

cod

-1

-2

-3

-1

KSK

-2

KSK

-6

-1

-2

-3

-5

-7

-8

-1

Ammobaculites

0.9

0.1

3.3

1.7

1.5

1.1

0.4

0.5

astrorhizids

1.5

1.3

0.5

0.7

3.6

6.1

7.7

1.2

6.5

4.5

5.1

9.2

2.4

5.7

1.1

2.5

Conotrochammina

0.4

Evolutinella

0.4

Reophax

2.0

0.9

2.0

Subreophax

0.6

0.7

0.4

1.2

Textularia

0.4

Tolypammina

0.6

0.1

0.4

Trochammina

0.9

0.3

0.7

0.5

2.2

1.7

0.6

0.8

0.5

1.1

2.3

1.2

1.5

Verneulinoides

0.8

2.0

Astacolus

4.7

2.3

3.0

1.5

3.3

3.1

3.6

4.3

2.5

3.0

0.9

4.8

3.3

6.1

7.3

3.3

2.6

5.0

Citharina

0.3

0.7

0.5

0.3

3.4

0.9

0.4

0.8

0.4

1.0

Eoguttulina

2.8

5.4

2.6

6.5

8.6

5.1

0.3

0.9

0.6

1.5

3.7

0.7

1.1

0.4

3.8

0.5

Falsopalmula

0.4

1.3

1.0

1.4

0.6

1.5

2.6

1.1

3.2

2.0

1.5

Frondicularia

0.6

0.3

0.5

0.7

0.5

0.6

0.9

0.5

1.5

3.1

0.8

0.4

0.5

Laevidentalina

13.2

17.2

15.1

12.2

23.6

26.3

31.6

25.5

27.1

30.6

56.2

38.0

38.3

41.9

29.4

22.2

35.0

Lenticulina

15.8

9.2

11.6

13.6

18.5

21.5

37.1

23.1

27.3

46.6

9.3

8.6

17.5

13.4

14.5

18.8

16.2

19.5

Lingulina

0.3

1.0

0.8

Marginulina

0.8

Marginulinopsis

0.6

0.4

0.5

0.3

1.5

1.1

1.7

0.6

0.9

0.2

1.8

0.8

1.5

Nodosaria

0.4

1.3

0.5

1.0

3.1

1.9

0.7

1.9

1.6

0.8

1.5

Nodosaria regularis

0.6

1.0

0.5

0.3

0.5

5.5

3.4

1.9

2.3

0.2

1.8

5.4

1.2

ophthalmidiids

0.6

0.3

0.4

0.5

1.3

1.2

1.5

0.4

Patellina

1.7

4.6

5.2

2.0

0.3

Planularia

1.9

0.2

0.4

Pseudonodosaria

1.7

3.1

0.9

1.5

17.2

1.0

2.2

1.2

2.8

2.1

0.7

0.8

2.4

0.4

0.5

Pyramidulina

0.9

0.8

0.5

1.7

1.0

1.1

1.2

0.9

1.2

2.2

1.5

0.8

1.1

0.5

Ramulina

0.4

0.1

1.3

0.5

0.9

0.6

1.5

1.4

1.5

0.4

1.9

3.5

Reinholdella

1.3

3.1

5.6

4.5

0.7

Saracenaria

0.4

0.8

1.0

Spirillina

49.7

48.5

52.2

50.8

25.4

28.2

1.4

16.2

31.7

12.0

38.9

16.7

16.8

10.3

21.0

29.0

45.9

17.0

Vaginulina

0.3

0.7

2.6

0.6

0.8

0.2

0.4

1.2

0.4

0.5

Vaginulinopsis

1.1

0.5

0.9

0.5

1.0

3.6

3.9

0.5

2.9

2.7

2.4

2.0

1.5

4.0

SUM

100

Axis 1

Axis 2

Axis 3

Eigenvalues

352.657

124.015

30.304

Percentage

66.264

23.303

5.694

Cum. Percentage

66.264

89.567

95.261

PCA variable loadings

Axis 1

Axis 2

Axis 3

ASTRORHIZIDS

-0.078

0.089

-0.097

other

AGGLUT. FORAM.

-0.065

-0.062

-0.089

Astacolus

-0.042

0.042

-0.037

Eoguttulina

0.085

-0.045

-0.233

Laevidentalina

-0.451

0.662

0.407

Lenticulina

-0.26

-0.719

0.508

Marginulina/-opsis

-0.012

-0.011

-0.01

Nodosaria

-0.001

0.02

-0.031

Nodosaria regularis

-0.054

-0.052

-0.034

NODOSARIIDS

(others)

-0.05

0.024

-0.099

OPHTHALMIDIIDS

0.002

-0.026

-0.004

Patellina

0.061

-0.001

Pseudonodosaria

0.026

-0.061

-0.576

Pyramidulina

-0.006

0.012

-0.051

Ramulina

-0.014

-0.001

0.011

Reinholdella

0.066

-0.005

Spirillina

0.833

0.144

0.392

Vaginulina/-opsis

-0.039

-0.01

-0.056

Axis 1

Axis 2

Axis 3

Axis 4

Axis 5

Eigenvalues

7.475

2.901

2.419

1.351

1.171

Percentage

41.526

16.114

13.438

7.508

6.504

Cum. Percentage

41.526

57.64

71.077

78.585

85.089

PCA variable loadings

Microfossils

Axis 1

Axis 2

Axis 3

Axis 4

Axis 5

FORAM.

Abundance

0.33

0.112

0.222 0

0.079

astrorhizids

0.001

0.468

0.144

0.094

0.353

other AGGL. forams.

-0.026

0.42

-0.041

0.063

-0.585

Eoguttulina

0.343

-0.039

-0.068

-0.021

0.006

Laevidentalina

0.09

0.327

0.279

-0.183

0.49

Lenticulina+Astacolus

0.267

0.187

-0.036

0.056

-0.139

elong. nodosariids

0.229

0.252

-0.342

-0.204

0.019

Ramulina+N.regularis

0.002

0.504

0.047

0.005

-0.192

Patellina

0.29

-0.089

0.324

0.066

-0.15

Pseudonodosaria

0.217

0.069

-0.278

-0.35

-0.227

Reinholdella

0.274

-0.173

0.302

0.149

-0.105

Spirillina (+ophthalm.)

0.319

-0.092

0.246

0.085

-0.046

OSTRACODS

(valves/2)

0.336

-0.011

-0.12

-0.026

0.046

Valves/Carapaces ratio

-0.141

0.262

0.125

0.459

0.016

SPONGE

spicules

0.246

0.051

-0.379

0.011

0.21

ECHINOID

spines

0.141

-0.035

-0.418

0.404

0.287

CRINOID

+other

ECHINOD.

0.336

-0.084

0.122

0.021

-0.041

HOLOTHURIAN

sclerites

-0.085

0.024

0.169

-0.614

0.11