GEOLOGICA CARPATHICA, OCTOBER 2008, 59, 5, 411—424
www.geologicacarpathica.sk
Paleoecology of benthic foraminifera of the Baden-Sooss
section (Badenian, Middle Miocene, Vienna Basin, Austria)
KATALIN BÁLDI and JOHANN HOHENEGGER
Department of Paleontology, University of Vienna, Althanstrasse 14,
1090 Vienna, Austria;
kati.baldi@siemelink.net; johan.hohenegger@univie.ac.at
(Manuscript received December 13, 2007; accepted in revised form June 12, 2008)
Abstract: A quantitative analysis of benthic foraminifera was carried out on a scientific core from the type locality of the
Paratethyan stage Badenian (Middle Miocene) and results were compared to stable isotopes (
δ
18
O and
δ
13
C) and magnetic
susceptibility. Two approaches were applied to reconstruct the paleoenvironment. First, several indices (inbenthics %,
oxyphylic %, foraminiferal numbers, diversity) and paleodepth proxies were calculated, second, multivariate statistics
(Detrended Correspondence Analysis, cluster analyses combined with the indicator value method) were carried out. High
correlations between environmental indices and core-depth manifest some trends. Five biofacies units were recognized,
and the lowermost unit I encompasses a laminated interval coinciding with a biofacies subunit. The most indicative and
abundant species is Uvigerina semiornata, a dysoxy tolerant inbenthic. The following unit II can be characterized as a
stable period of improved oxygenation and lowered nutrient supply. Unit III is distinguished by Trifarina angulosa pre-
ferring well-aerated turbulent bottom water and sandy substrate, while accompanied by some low salinity tolerant species.
A non-stratified water column with high terrigenous input is accepted for this unit. Unit IV is characterized by low
diversities and the deep inbenthic indicator species Bulimina elongata, Fursenkoina acuta, tolerant of oxygen deficiency
and benefiting from unlimited food supply in a well-stratified sea. The high foraminiferal numbers are the consequence of
low terrigenous input not diluting the highly productive opportunistic species in the sediment. The indicator species of unit V,
the reticulate Bolivina viennensis, has ecological needs similar to the previous unit. Environmental changes from unit IV
to V are the increased terrigenous input and oscillation, indicating less stable conditions. The biofacies unit boundaries
always coincide with major shifts of the isotope and susceptibility record. It is concluded, that the physical environmental
parameters controlling benthic foraminiferal distribution are primarily influenced by proximity to land.
Key words: Middle Miocene, Paratethys, Vienna Basin, statistics, paleowater proxy, benthic foraminifera, Badenian
stratotype.
Introduction
The Badenian of the Vienna Basin has been of great interest
for foraminiferal experts since the pioneer monograph of Al-
cide d’Orbigny (1846). The work of d’Orbigny founding the
basis of foraminiferal taxonomy has been revised by Marks
(1951) and Papp & Schmid (1985). Since the times of
d’Orbigny, two lines of interest have developed among fora-
miniferal experts. Some wanted to reconstruct the flow of
time recorded in the sediments by studying foraminifera,
while others wanted to reconstruct the environment of the
past. A significant example of this first kind is the benthic
foraminiferal zonation of Grill (1941) of the Vienna Basin
used for correlation in the entire Central Paratethys. The ba-
sic work defining the Badenian, its correlation and strato-
types (Papp et al. 1978) gives proper attention to the
foraminifera of the studied area. The most recent summariz-
ing work of similar magnitude is Cicha et al. (1998) focusing
on Central Paratethyan foraminifera. There are several works
dealing with the correlation of the Central Paratethys based
on foraminifera (Rögl 1996). Contributions from a paleoeco-
logical point of view applying a fully quantitative approach
are of Rupp (1986) on the Vienna Basin, and partly Mandić
et al. (2002), while Spezzaferri et al. (2002), Spezzaferri
(2004) and Hohenegger (2005) dealt with material from the
neighbouring Styrian Basin.
The aim of the present work is the reconstruction of the pa-
leoenvironment based on the recently deepened Baden-Sooss
borehole close to the stratotype section (Fig. 1). Fortunately,
most benthic species of the Middle Miocene Paratethys have
present-day living counterparts, thus paleoenvironmental re-
constructions can rely on actualistic principles. Recent contri-
butions to benthic foraminiferal ecology, concerning the
in-sediment microhabitat distribution in relation to food avail-
ability and bottom water oxygenation (e.g. Jorissen et al.
1995; de Stigter et al. 1998; Kaiho 1999; Den Dulk et al.
2000; Fontanier et al. 2002; Altenbach et al. 2003; Kouwen-
hoven & Van der Zwaan 2006; Hayward et al. 2007) and sta-
tistical methods were employed (Dufrêne & Legendre 1997;
McCune & Mefford 1999; Hammer & Harper 2005). In the
present work, it is hoped that benthic paleoecology results can
be confirmed by independent geochemical and geophysical
evidence. The independent records available of the studied
section are the oxygen and carbon stable isotopes (
δ
18
O and
δ
13
C) and magnetic susceptibility. Thus, the samples were
prepared in the present work especially for statistical analysis,
and the species concept applied here used generalizing, ‘lump-
ing’ species (Báldi 1999).
412
BÁLDI and HOHENEGGER
Methods
Sample processing
Benthic foraminiferal analyses were carried out on a sam-
ple-set taken in regular intervals of 1.25 m from the bottom of
the core at 102 m to 8 m at the top. The sediments were rather
undisturbed in the lower part (40 to 102 m), and tectonically
affected in the upper part (8 to 40 m; Hohenegger et al. 2007).
The samples were dissolved in pure water, then sieved
through a set of standard sieves. Fractions from sieves were
combined to make the ‘fine fraction’ (63—125
µm), the ‘me-
dium fraction’ (125
µm—2 mm) and ‘coarse fraction’
( > 2 mm). All fractions were paid a quick look under a bin-
ocular microscope to exclude the possibility of missing in-
formation by analysing only the medium fraction. The fully
quantitative analysis of benthic foraminifera was carried out
on the medium fraction (125
µm—2 mm). This fraction was
split with a microsplitter to the appropriate size of more than
200 specimens of benthic foraminifera. All benthic foramin-
ifera were picked from the split and mounted on a microslide
with a layer of glue (Tragacantha). This glue makes it possi-
ble to turn specimens with wet brush to see different, taxo-
nomically important views of foraminifera. Planktonic
foraminifera were counted under the microscope in the same
split used for the benthic foraminiferal analysis.
The quantitative analyses of benthic foraminifera were
carried out on 74 samples eligible for statistics containing
more than 200 benthic foraminiferal specimens. Altogether,
102 taxonomic categories were distinguished including 68
on the species level (Table 1; Cicha et al. 1998; Báldi 1999).
Strictly all specimens were counted, even broken or badly-
preserved. Counting broken specimens resulted in occasion-
al decimals, expressing the picked fragments proportion to
the whole test. Sometimes, when determination to the gener-
ic level was impossible, the categories of ‘undetermined cal-
careous’ or ‘agglutinated’ specimens were used. These
‘undetermined’ categories also include the extremely spo-
radic occurrences of determinable species, that is few speci-
mens in all the material. However, the percentage of
‘undetermined calcareous’ or ‘agglutinated’ specimens was
kept below 5.99 % of all benthic foraminifera in any particu-
lar sample.
The basic lists can be found as an Electronic Supplement of
this paper in web version at http://www.geologicacarpathica.sk.
Quantitative methods
The analyses of benthic foraminifera were directed to-
wards determining the environmental changes, possible
trends and underlying gradients in the studied time interval.
Two sorts of approach were applied:
1. Indices and proxies were calculated for environmental
reconstruction.
2. For multivariate statistics the Detrended Correspon-
dence Analysis (DCA) and cluster analyses were carried out
2
Fig. 1. a – Tectonic map of the
Vienna Basin and location of
the studied borehole Baden-
Sooss. b – Schematic sedimen-
tological log of the borehole
Baden-Sooss (after Hohenegger
et al. 2008).
413
PALEOECOLOGY OF BENTHIC FORAMINIFERA OF THE BADEN-SOOSS SECTION (BADENIAN, AUSTRIA)
to contribute to our understanding of benthic assemblages
coupled with environmental shifts or gradients.
Indices and proxies
The inbenthic and oxyphylic indices are based on the dif-
ferent microhabitat preferences of benthic foraminifera in re-
lation to their ecological needs and tolerances (Corliss 1985;
Corliss & Chen 1988; Jorissen et al. 1995; Fontanier et al.
2002; Altenbach et al. 2003; Hayward et al. 2007). These in-
dices are the percentages of taxa following an inbenthic
mode of life, or needing well aerated bottom water in the
case of oxyphylic species. Considering the in-sediment dis-
tribution of each recognized taxa, where possible, a decision
was made on the inbenthic or oxyphylic nature of the partic-
ular species (Corliss 1985; Corliss & Chen 1988; Kaiho
1994, 1999; de Stigter et al. 1998; Den Dulk et al. 2000;
Kouwenhoven & Van der Zwaan 2006), see details in Báldi
(2006). The selection of taxa considered inbenthic or oxy-
phylic are to be found denoted by asterisks next to their
name in Table 1. Variables based on percentages were trans-
formed using the arcsine-root transformation. The benthic
foraminiferal number in 100 g sediments was calculated
from the proportion of split size yielding the > 200 specimens
benthic sample to the total original dry sediment sample
weighed on an analytical scale.
Table 1: Total count of species and species groups and their presence in samples. Inbenthic taxa are denoted by single asterisks, while oxy-
phylic taxa are denoted by double asterisks.
Species
Specimen
number
Number
of
samples
Species
Specimen
number
Number
of
samples
Haplophragmoides sp.
4 4
Cibicides ungerianus (d’Orbigny)**
229 49
Martinotiella communis (d’Orbigny)
55.8 34
Dentalina brevis (d’Orbigny)
4 3
Sigmoilopsis sp.
115 48
Dentalina communis (d’Orbigny)
27 15
Spiroplectammina carinata (d’Orbigny)
529 68
Dentalina elegans (d’Orbigny)
131 24
Spiroplectammina spp.
43 8
Ehrenbergia serrata Reuss
30 10
Textularia cf. mexicana
130 41
Elphidium spp.
204 62
Textularia sp.
68 19
Fissurina spp.
69 41
Textularia spp.
171 58
Fursenkoina acuta (d’Orbigny)*
152 47
Vulvulina spp.
104 44
Gavelinopsis praegeri (Heron-Allen & Earland)
550 70
Glandulina sp.
6 5
Adelosina spp.**
49 17
Globobulimina pyrula (d’Orbigny)*
239 53
Cornuspira plicata (Czjzek)
13 7
Guttulina sp. (d’Orbigny)
10 9
Pyrgo spp.**
34 21
Gyroidina parva (Cushman & Renz)
32 14
Quinqeloculina spp.**
144 47
Gyroidina soldanii (d’Orbigny)
82 28
Sigmoilinita tenuis (Czjzek)**
163 60
Gyroidina umbonata (Silvestri)
271 56
Spiroloculina excavata (d’Orbigny)**
49 25
Hanzawaia boueana (d’Orbigny)**
266 65
Triloculina spp.**
21 10
Heterolepa dutemplei (d’Orbigny)**
503 70
Miliolid spp.** 119
45
Hoeglundina elegans (d’Orbigny)
210 49
Lagena spp.
89 30
Alabamina sp.
130 36
Lenticulina arcuata (d’Orbigny)**
17 9
Allomorphina trigonia (Reuss)*
13 12
Lenticulina calcar (Linné)**
34 15
Ammonia beccarii (Linné)
68 34
Lenticulina sp.**
245 61
Amphicoryna badenensis (d’Orbigny)
53 30
Marginulina hirsuta (d’Orbigny)
29 17
Asterigerinata planorbis (d’Orbigny)
222 59
Melonis pompilioides (Fichtel & Moll)*
738 74
Astrononion cf. italicum (Cushman & Edwards)
120 46
Nodosaria raphanistrum (Linné)
19 12
Biapertorbis biaperturatus Pokorny
145 33
Nodosaria sp.1
40 31
Bitubulogenerina reticulata Cushman
22 20
Nodosaria sp.2
19 16
Bolivina antiqua (d’Orbigny)*
16 13
Nonion commune (d’Orbigny)
1052 60
Bolivina hebes (Macfadyen)
63 21
Nonionella turgida (Williamson)
15 13
Bolivina plicatella (Cushman)
325 38
Oolina spp.
17 13
Bolivina scalprata var. Miocenica (Macfadyen)
246 68
Oridorsalis umbonatus (Reuss)
56 29
Bolivina spathulata (Williamson)*
29 6
Pappina parkeri (Karrer)*
24 15
Bolivina spp.
67 30
Pullenia bulloides (d’Orbigny)*
179 59
Bolivina undetermined (contorted forms)
10 9
Pullenia quinqueloba (Reuss)*
35 19
Bolivina viennensis (Marks)*
255 49
Reussella spinulosa (Reuss)
18 13
Bulimina aculeata (d’Orbigny)*
797 68
Rosalina spp.
40 24
Bulimina costata (d’Orbigny)
694 49
Siphonina reticulata (Czjzek)**
13 9
Bulimina elongata (d’Orbigny)*
665 58
Sphaeroidina bulloides (d’Orbigny)*
99 33
Cancris auriculus (Fichtel & Moll)
26 15
Stilostomella adolphina (d’Orbigny)
109 36
Cassidulina carinata (Silvestri)
73 31
Trifarina angulosa (Williamson)
400 57
Cassidulina laevigata (d’Orbigny)
531 67
Trifarina bradyi (Cushman)
29 16
Cassidulina oblonga (Reuss)*
231 41
Uvigerina aculeata (d’Orbigny)*
235 18
Cassidulina spp.
19 9
Uvigerina acuminata (Hosius)*
37 13
Cassidulina subglobosa (Brady)*
468 69
Uvigerina acuminata f. macrocarinata*
8 3
Ceratocancris hauerina (d’Orbigny)
270 52
Uvigerina semiornata (d’Orbigny)*
327 33
Cibicides kullenbergi (Parker)**
49 20
Uvigerina spp.*
24 10
Cibicides lobatulus (Walker & Jacob)**
1147 74
Valvulineria complanata (d’Orbigny)*
803 72
Cibicides spp.
17 10
414
BÁLDI and HOHENEGGER
Different aspects of diversity were examined in the present
work:
– dominance (D) taking values from 0 to 1 (Simpson
1949; Magurran 2004)
where n
j
indicates abundance of the j
t h
species and m the
number of species. When it takes the maximum value 1, one
single species dominates the whole assemblage.
– Shannon index (H)
reaching maximum values when a few individuals of
many taxa are present in a sample, a sort of measure of entro-
py (Shannon 1948).
– Fisher-
α
m =
α ln (l+n/α)
a classical measure of species richness able to compensate
for variable sample sizes. Fisher’s
α diversity is calculated
using the formula given by Fisher et al. (1943) with the con-
stants from Williams (1964: fig. 125). All diversity indices
were calculated by the program PAST (Hammer & Harper
2005).
Paleowater depth as an essential factor determining the dis-
tribution of marine life was estimated by two independent
proxies. The first proxy is based on the ratio of planktonic and
benthic foraminifera (abbreviated as P/B-ratio in the follow-
ing) in the sediment (Van der Zwaan et al. 1990, 1999), while
the second one is an extension of the transfer equation devel-
oped by Hohenegger (2005) including species abundance by
needing the depth ranges of taxa as an input. In this equa-
tion, the location parameters l of the j
t h
species is weighted on
the one side by abundance n
j
and by dispersion d
j
on the other.
Multivariate analyses
First, Detrended Correspondence Analysis (DCA) was
used to find relations between samples and species to sup-
port environmental interpretation (program PAST, Hammer
& Harper 2005). Second, samples were clustered into non-
overlapping classes by Ward’s Method based on relative Eu-
clidean distances (program PC-ORD – McCune & Mefford
1999). Since species normally cannot be grouped into non-
overlapping clusters, the indicator value method (Dufrêne &
Legendre 1997) has been used for finding characteristic spe-
∑
=
−
=
m
j
j
j
n
n
n
n
H
1
)
/
(
ln
)
/
(
)
(
/
)
(
1
1
1
1
∑
∑
=
−
=
−
=
m
j
j
j
m
j
j
j
j
d
n
d
l
n
depth
∑
=
=
m
j
j
n
n
D
1
)
/
(
2
Fig. 2. Benthic foraminiferal indices and proxies plotted against depth in core with biofacies units on the left. From left to right cumulative
percentages of inbenthic taxa, oxyphylic taxa, paleowater depth proxies with planktonic/benthic (PB) ratio and depth gradient analysis,
benthic foraminiferal number calculated in 100 g dry sediments, species diversities of Fischer-
α and of Shannon (H) and Dominance.
415
PALEOECOLOGY OF BENTHIC FORAMINIFERA OF THE BADEN-SOOSS SECTION (BADENIAN, AUSTRIA)
cies determining the sample clusters. This method has hard-
ly ever been used in paleontology, though it was successful-
ly employed in biology (Dai et al. 2006; Nahmani et al.
2006). It combines information on the concentration (A
jk
) of
species j in a particular cluster k and the faithfulness of oc-
currence of a species in a particular cluster (B
jk
), where
with r representing the number of samples in cluster k and
g representing the number of clusters. Transforming relative
abundance A
jk
of species j in cluster k into presence/absence
values yields B
jk
. This method produces indicator values
(IndVal
jk
) for each species j in each cluster k by
),
/
(
/
)
/
(
1
1
1
∑∑
∑
=
=
=
=
g
k
r
i
ijk
r
i
ijk
jk
r
n
r
n
A
Table 2: Calculated indices and proxies against core depth in Fig. 2.
IndVal
jk
= A
jk
. B
jk
. 100,
which are tested for statistical significance using the Mon-
te Carlo technique. The indicator value reaches its maximum
(i.e. 100 %), when all individuals are found in a single clus-
ter and when the species is present in all samples of that par-
ticular cluster.
Results
Indices and proxies
The indices and proxies (Fig. 2) are based on the counting
results and percentages (Table 2, Fig. 3). Correlations were
D
epth i
n c
or
e (m
)
Nu
mb
er
of
b
en
th
ic f
or
amin
if
era
12
5 µ
m
t
o 2 m
m
in
10
0 g
s
ed
im
en
t
P
er
ce
ntag
e o
f
inbe
nthic
t
axa
on
to
ta
l be
nthi
c for
am
ini
fe
ra
P
erc
en
ta
ge
of
o
xyp
hy
lic
ta
xa
on
to
ta
l be
nthi
c for
am
ini
fe
ra
P
erc
en
ta
ge
of
p
lan
kt
on
ic
for
am
in
if
er
a
Nu
mb
er
of
b
en
th
ic sp
ec
ie
s
in
cl
us
iv
e unde
te
rm
in
ed
g
ro
ups
D
epth i
n c
or
e (m
)
Nu
mb
er
of
b
en
th
ic f
or
amin
if
era
12
5 µ
m
t
o 2 m
m
in
10
0 g
s
ed
im
en
t
P
er
ce
ntag
e o
f
inbe
nthic
t
axa
on
to
ta
l be
nthi
c for
am
ini
fe
ra
P
erc
en
ta
ge
of
o
xyp
hy
lic
ta
xa
on
to
ta
l be
nthi
c for
am
ini
fe
ra
P
erc
en
ta
ge
of
p
lan
kt
on
ic
for
am
in
if
er
a
Nu
mb
er
of
b
en
th
ic sp
ec
ie
s
in
cl
us
iv
e unde
te
rm
in
ed
g
ro
ups
8.41
17532 47.8
31.9
52.9 35
56.41
10877 30.2 19.0 74.1 47
10.02
10880 35.8
20.1
24.2 45
57.61
11711 24.2 23.2 79.8 46
11.21
8285 41.2
19.0
23.6 33
58.81
34384 28.2 16.5 72.6 48
12.44
16121 39.9
27.9
29.4 33
60.01
10073 22.7 24.5 82.5 58
13.62
5936 46.0
26.1
37.9 41
61.41
9509 20.0 19.2 59.2 52
14.81
5641 41.6
31.2
31.8 33
62.61
14295 19.0 24.0 76.2 54
16.41
5493 38.3
21.8
15.6 30
63.82
21333 21.2 24.3 50.4 46
17.61
33639 34.5
39.1
41.8 33
65.01
4422 27.6 19.3 78.4 53
18.88
11616 43.1
28.9
29.0 37
66.42
8975 39.5 16.0 76.9 49
20.02
12585 42.3
35.0
30.8 40
67.61
8812 31.5 25.0 75.0 50
22.41
16486 43.5
12.2
16.9 25
68.81
12030 29.1 22.2 75.5 57
23.61
65030 45.7 6.8
27.5 30
70.01
9719 35.1 22.0 56.9 46
24.86
32156 39.8
13.1
31.3 23
71.42
14222 35.8 18.1 72.9 40
26.41
55242 48.3 9.3
38.1 25
72.61
10628 34.6 17.3 77.3 55
27.62
27858 41.1
15.5
32.2 28
73.81
7538 25.5 17.5 67.0 52
28.86
22135 43.5
20.8
53.9 41
75.01
7487 29.4 26.2 86.1 49
30.01
38847 44.4
14.9
55.3 30
76.41
13392 22.7 26.5 68.1 68
31.22
41189 45.4
21.8
12.6 32
77.61
7697 30.5 21.9 65.4 55
34.01
9762 33.5 5.9
63.5 39
78.81
7754 28.6 36.6 74.9 54
35.21
8758 33.2
31.1
59.3 47
80.01
18988 22.9 32.9 64.4 60
36.81
3398 29.5
30.0
57.5 56
81.41
14122 22.3 28.4 70.3 62
37.61
4205 27.1
25.7
49.5 54
82.61
9691 19.8 29.7 67.6 54
38.81
4533 21.8
27.0
55.7 56
83.83
3408 34.7 27.7 75.3 52
40.01
5277 48.6
13.3
33.7 43
85.01
4967 32.8 24.7 65.1 60
41.21
2528 26.8
30.2
59.4 51
86.41
8163 30.6 33.5 72.6 50
42.41
11084 37.2
21.7
72.3 53
87.61
11396 17.7 36.5 70.1 61
42.61
5065 27.6
19.7
63.6 39
88.81
2709 26.5 33.5 76.6 59
43.81
5747 32.3
20.1
72.4 45
90.01
189 23.7 34.3 70.0 57
45.01
5257 31.4
22.0
68.0 60
91.41
5653 35.1 27.9 73.4 51
46.41
1939 28.5
21.5
66.2 44
92.63
8696 23.4 26.6 78.0 53
47.61
2199 31.0
22.9
58.3 50
93.81
6838 27.1 26.6 70.5 55
48.81
6121 33.1
18.0
67.2 50
95.01
4724 27.8 29.4 41.9 55
50.01
2205 29.4
15.6
76.2 44
96.41
8234 30.7 26.7 79.5 52
51.41
8648 41.1
12.1
73.7 37
97.61
15298 34.3 28.2 71.7 51
52.61
9356 48.6
10.2
61.3 43
98.81
13022 29.4 17.0 80.3 55
53.81
14171 35.0
14.7
65.7 52
100.01
13077 33.9 21.6 79.1 53
55.01
29757 35.7
17.2
68.4 56
101.41
1292 49.0 24.3 37.6 35
416
BÁLDI and HOHENEGGER
Fig. 3. a – The relative frequencies (percentages) of the most common taxa (>10 %) plotted against depth in core and illustrated by SEM;
bar length = 100
µm. Bolivina hebes (Macfadyen) from 95.01 m, Bolivina viennensis Marks from 42.41 m, Bulimina aculeata (d’Orbigny)
from 95.01 m, Bulimina elongata (d’Orbigny) from 28.81 m, B. costata (d’Orbigny) from 95.01 m, Cassidulina laevigata (d’Orbigny)
from 8.41 m, Cibicides lobatulus (Walker & Jacobs) from 95.01 m, Dentalina elegans (d’Orbigny) from 8.41 m, Globobulimina pyrula
(d’Orbigny) 95.01 m. b – bar length = 100
µm. Heterolepa dutemplei (d’Orbigny) from 16.41 m, Melonis pompilioides (Fichtel & Moll)
from 42.41 m, Nonion commune (d’Orbigny) from 95.01 m, Trifarina angulosa (Williamson) from 95.01 m, Uvigerina semiornata
(d’Orbigny) from 8.41 m, Uvigerina aculeata (d’Orbigny) from 95.01 m, Valvulineria complanata (d’Orbigny) from 95.01 m,
Spiroplectammina carinata (d’Orbigny) from 42.41 m.
a
b
417
PALEOECOLOGY OF BENTHIC FORAMINIFERA OF THE BADEN-SOOSS SECTION (BADENIAN, AUSTRIA)
calculated among environmentally significant indices and
proxies (inbenthics %, oxyphylic %, paleowater depth prox-
ies, benthic foraminiferal number in 100 g sediment and dif-
ferent diversity indices and core depth; Table 3). Calculating
correlation with core depth makes it possible to verify up-
ward trends. Generally, the indices showed significant corre-
lation with core depth meaning definite trends through the
Multivariate statistics
Detrended Correspondence Analyses
(DCA)
This analysis was carried out on speci-
men counting. In the present study, the
first two axes were considered and plot-
ted against each other. These plots
(Fig. 4) showed five clouds or sample
groups (1, 2, 3, 4, 5), where each group
contains samples of a rather distinct core
depth.
Group 1) consists of all samples be-
tween 82.6 and 88.8 m with the excep-
tion of the sample from 85.01 m. Group
2) comprises samples from three distinct
intervals. These are the lowermost sam-
ples of the core between 90.0 and
101.4 m, then between 76.4 and 81.4 m,
and between 35.2 and 41.2 m. Group 3)
Fig. 4. Presentation of samples by the first and second axes of DCA (Detrended
Correspondence Analysis) and subjective clustering into 5 groups marked by colours.
De
pth (m
)
in
be
nthi
c (%)
ox
yp
hy
li
c (
%
)
P
/B
p
ale
ow
at
er
d
ep
th
N
o. of
b
en
th
ic i
n 10
0 g
Do
m
in
anc
e
Sha
nno
n
Fis
ch
er-
α
D
ep
th
gr
ad
ie
nt
an
al
ys
es
Depth (m)
0.00 0.03 0.00 0.01 0.00 0.00 0.00 0.07
inbenthic (%)
0.53
0.00 0.00 0.00 0.00 0.00 0.00 0.00
oxyphylic (%)
–0.25 –0.40
0.68 0.00 0.00 0.00 0.00 0.24
P/B paleowater depth
–0.66 –0.44 0.05
0.01 0.00 0.00 0.00 0.08
No. of benthic in 100 g
0.32 0.35 –0.38 –0.31
0.00 0.00 0.00 0.29
Dominance
0.44 0.57 –0.54 –0.47 0.40
0.00 0.00 0.81
Shannon
–0.61 –0.67 0.52 0.57 –0.49 –0.94
0.00 0.72
Fischer-α
–0.65 –0.68 0.45 0.60 –0.50 –0.76 0.91
0.80
Depth gradient analyses –0.21 0.39 –0.14 0.20 –0.12 –0.03 0.04 0.03
Table 3: Correlation matrix, where in lower triangle Pearson’s r to be found, while in up-
per triangle the probability of uncorrelated columns. Calculated in PAST (Hammer &
Harper 2005).
contains most samples between 42.4 and 75.0 m, and an oc-
casional occurrence of this group expressed in sample
34.0 m above the mentioned interval. Group 4) has samples
from the interval 22.4 to 30.0 m. Samples from the upper
part of the core belong to group 5) between 8.4 and 20.0 m,
with a few additional samples below 85.0 m and at 31.2 m
(Fig. 4).
studied interval. An increase of inbenthic
(r = 0.53) and a slightly decreasing oxyphylic
(r = —0.25) trend was observed up the core. Both
paleowater proxies demonstrated a significant
shallowing trend (P/B: r = —0.66, depth gradient:
r = —0.21) through the examined time interval,
also the number of benthic specimens in 100 g
sediment increased (r = 0.32). The highest corre-
lations are found, not surprisingly, among the
different diversity indices. Nevertheless, diversi-
ties showed very strong trends up the core (Domi-
nance: r = 0.44, Shannon: r = —0.61, Fisher-
α:
r = —0.65), meaning enhancing dominance and
declining diversity with time. The oxyphylic and
inbenthic taxa (r = —0.4) show significant nega-
tive correlation, as expected. The inbenthic in-
dex has high values when diversity is low and
dominance is high (inbenthic to Dominance:
r = 0.57, inbenthic to Shannon: r = —0.67, inbenth-
ic to Fisher-
α: r=—0.68) while oxyphylic taxa
show the opposite correlation to inbenthics
(oxyphylic to Dominance: r = —0.54, oxyphylic to
Shannon: r = 0.52, oxyphylic to Fisher-
α: r=0. 45).
The two depth proxies behave differently to di-
versity. Depth estimation based on the P/B-ratio
highly correlates with diversity (with Domi-
nance r = —0.47, Shannon r = 0.57, Fisher-
α
r = 0.60), while the depth gradient analysis based
estimations shows negligible correlation with di-
versity (with Dominance r = —0.03, Shannon
r = 0.04, Fisher-
α r=0.03; see results in Table 3).
418
BÁLDI and HOHENEGGER
Detrended Correspondence Analyses (DCA) were also ap-
plied to recognize species groups, but there were no distinct
well-defined groups because of overlapping clusters in R-mode
classification analysis.
However, looking at the bivariate plot of the first two axis
(Fig. 5), species on the left are deep inbenthic, low oxygen
tolerant forms (Bulimina elongata, Fursenkoina acuta, B.
spathulata), while oxyphylic species of epibenthic, or even
epiphytic habitat (Hanzawaia boueana, Cibicides lobatulus,
Cibicides spp.) tend to appear on the right side of the plot.
Several species suspected to be deepwater markers appear on
the right upper side of the plot (Siphonina reticulata, Cibi-
cides ungerianus, Cibicides kullenbergi) as expected of oxy-
phylic deep dwelling taxa (Fig. 5).
Cluster Analyses (CA)
The first step to attain indicator values of species according
to the method described in Dufrêne & Legendre (1997) is to
carry out a cluster analysis. Ward’s method based on the arc-
sine-root transformed percentage database without cut-off lev-
els to ignore rare species was used. Generally, the
recognizable clusters (Fig. 6) appear to include samples from
particular intervals of core depth. Eight clusters (a, b, c, d, e, f,
g, h) can be separated with the cut-off level 49 % of informa-
tion remaining. The uppermost samples of the borehole be-
long to cluster a) from 8.4 to 20.0 m (except – 16.4 m) and to
another interval from 35.21 to 37.61 m and two additional
samples at 45.0 m and 75.0 m. Cluster b) samples fall into one
single distinct interval from 80.0 to 91.4 m. The cluster c) is to
be found from 76.4 to 78.8 m and from 38.8 to 43.8 m. Clus-
ters d) and e) are closely related and both scatter in the
depth interval from 53.8 to 73.8 m, where cluster d) con-
sists of 11 samples (53.8 m, 55.0 m, 60.0 m, 63.8 m, 66.4 m,
67.61 m, 68.81 m, 70.01 m, 71.415 m, 72.61 m, 73.81 m),
while cluster e) comprises six samples (56.4 m, 57.6 m,
58.8 m, 61.4 m, 62.6 m, 65.0 m). The samples of cluster f) are
the lowermost samples of the borehole from the interval of
Fig. 5. Presentation of species by the first and second axes of DCA (Detrended Correspondence Analysis) and their position related to sam-
ple groups. Position of indicator species of clusters gained by Ward’s method (Table 4; Fig. 6) are accentuated by size and cluster colours
related to Fig. 6.
419
PALEOECOLOGY OF BENTHIC FORAMINIFERA OF THE BADEN-SOOSS SECTION (BADENIAN, AUSTRIA)
92.6 to 101.4 m. Cluster g) samples are from 22.4 to 31.2 m
and the additional sample of 16.4 m. The 7 samples of clus-
ter h) are from 46.4 to 52.6 m with the extra sample of
34.0 m (Fig. 6).
Combining methods – Ecofacies units
The most indicative species of each group with an indica-
tor value higher then 20 % are listed in Table 4. Based on the
multivariate methods (DCA and indicator values) five main
benthic foraminiferal ecofacies units (I—V) can be distin-
guished. Sample groupings of both methods seem to depend
on core depth, thus these groupings were translated to inter-
vals along the borehole, referred in meters. The boundaries
Fig. 6. Cluster analyses of samples by Ward’s method based on relative Euclidean distances.
of these main units are chosen, where
sample groupings of both methods coin-
cide suggesting a distinct boundary.
However, the main units contain smaller
subunits found by DCA and cluster
analysis (Fig. 7).
Unit I: This biofacies unit ranges
from 81.41 to 101.4 m containing the
lowermost part of the core. Both multi-
variate techniques (DCA and Cluster
Analyses) indicate a shorter interval en-
compassed in the subunit from 80.0 to
91.4 m (Cluster b)) or from 81.4 to
88.81 m (DCA group 1)). In this encom-
passed interval, indicated by darker
shades in Fig. 7 the species Uvigerina
semiornata has a high indicator value
(Table 4) and high abundance (Fig. 3),
while Siphonina reticulata, in spite of
the identically high indicator value, has
only a few specimens in the whole core
(Fig. 3). The number of inbenthic spe-
cies is relatively high in this subinterval
accompanied by a decrease in the abun-
dance of oxyphylic taxa, but oxyphylic
forms are rather abundant ( > 30 %) in
the whole unit. Therefore, the diversity
indices Fisher-
α and Shannon are slight-
ly high (Fig. 2).
Unit II: This biofacies unit is recog-
nized between 53.8 and 75.0 m. The
only indicator species of this period is
Bolivina hebes from the upper part (Ta-
ble 4). Generally, the inbenthic index is
higher in the lower than in the upper
part, while oxyphylic percentages are
rather stable. Depth estimations are fluc-
tuating, particularly the P/B ratio tends
to oscillate. Both diversity indices are
moderately high, indicating low domi-
nance (Fig. 2).
Unit III: The biofacies unit ranges
from 35.2 to 52.6 m. The species of
Nonion commune and Trifarina angu-
losa have high indicator values in the
lower part (cluster h)) accompanied by the ecologically im-
portant, otherwise rare taxa of Ammonia beccarii, Quinque-
loculina spp., div. miliolids. The indicator species of the
upper part (cluster c)) are Bulimina costata, Globobulimina
pyrula and Trifarina angulosa (Table 4). The last mentioned
T. angulosa as an indicator species of both clusters has spe-
cial significance in this ecofacies unit. It is not just indicative
of the whole period, but it is also common in this interval
reaching even 10 % of the assemblage. The inbenthic index
fluctuates in this interval showing two maxima at 52.6 and
40.0 m, and these peaks have corresponding oxyphylic mini-
ma. The number of benthic foraminifera is generally low.
Diversity indices are oscillating, slightly lower than before,
and higher dominance characterizes this interval (Fig. 2).
420
BÁLDI and HOHENEGGER
Unit IV: This biofacies unit ranges from 22.4 to 34.0 m.
The species Bulimina elongata (Indicator Value = 62),
Fursenkoina acuta (I.V. = 35), Bolivina spathulata (I.V. = 33),
Nonion commune (I.V. = 30) are highly indicative in this in-
terval as reflected in the high cumulative abundance of these
species on the inbenthic curve. The species Cassidulina lae-
vigata has a slightly lower indicator value, but reaches maxi-
mum abundance in this interval (Fig. 3). This biofacies is
rather distinct in many respects compared to other units. It is
characterized by high inbenthic and very low oxyphylic per-
centages. In this interval, the P/B-ratio estimates depths
shallower than the gradient method. The number of benthic
foraminifera is particularly high in this biofacies unit. This
interval is similarly distinctive in respect to diversity. The
Fisher-
α and Shannon diversity indices are low revealing
low species richness, thus high dominance is characteristic
in this biofacies unit (Fig. 2).
Unit V: This biofacies unit is found in the uppermost part
of the core from 20.0 m to the top at 8.4 m. The most indica-
Indicator values of clusters
Indicator values of clusters
Cluster a
Cluster
e
Bolivina viennensis (Marks)*
40
Bolivina hebes (Macfadyen)
52
Lenticulina arcuata (d’Orbigny)**
34
Spiroloculina excavata (d’Orbigny)**
29
Dentalina elegans (d’Orbigny)
27
Textularia sp.
29
Cibicides lobatulus (Walker & Jacob)**
23
Ceratocancris hauerina (d’Orbigny)
26
Uvigerina semiornata (d’Orbigny)*
23
Nonionella turgida (Williamson)
25
Bitubulogenerina reticulata Cushman
22
Bulimina costata (d’Orbigny)
23
Melonis pompilioides (Fichtel & Moll)*
21
Textularia spp.
21
Cluster
f
Heterolepa dutemplei (d’Orbigny)**
20
Uvigerina aculeata (d’Orbigny)*
92
Sphaeroidina bulloides (d’Orbigny)*
35
Cluster b
Ceratocancris hauerina (d’Orbigny)
33
Siphonina reticulata (Czjzek)**
41
Cibicides ungerianus (d’Orbigny)**
25
Uvigerina semiornata (d’Orbigny)*
41
Textularia cf. mexicana
24
Trifarina bradyi (Cushman)
36
Vulvulina spp.
21
Hoeglundina elegans (d’Orbigny)
35
Biapertorbis biaperturatus Pokorny
20
Cornuspira plicata (Czjzek)
33
Stilostomella adolphina (d’Orbigny)
32
Cluster
g
Cibicides spp.
27
Bulimina elongata (d’Orbigny)*
62
Bolivina plicatella (Cushman)
26
Fursenkoina acuta (d’Orbigny)*
35
Triloculina spp.**
26
Bolivina spathulata (Williamson)*
33
Hansawaia boueana (d’Orbigny)**
24
Nonion commune (d’Orbigny)
30
Globobulimina pyrula (d’Orbigny)*
23
Heterolepa dutemplei (d’Orbigny)**
29
Rosalina
22
Cassidulina laevigata (d’Orbigny)
28
Bolivina undetermined (contorted forms)
21
Asterigerinata planorbis (d’Orbigny)
22
Undetermined calcareous specimens
21
Cluster
h
Cluster c
Nonion commune (d’Orbigny)
38
Bulimina costata (d’Orbigny)
32
Trifarina angulosa (Williamson)
31
Globobulimina pyrula (d’Orbigny)*
30
Lenticulina calcar (Linné)**
26
Trifarina angulosa (Williamson)
28
Cassidulina oblonga (Reuss)*
25
Bulimina aculeata (d’Orbigny)*
23
Cluster d
Quinqeloculina spp.**
22
Alabamina sp.
25
Ammonia beccarii (Linné)
21
Lenticulina sp.**
25
Miliolid spp.** 20
Spiroplectammina carinata (d’Orbigny)
24
Pullenia bulloides (d’Orbigny)*
23
Textularia cf. mexicana
23
Cassidulina oblonga (Reuss)*
21
Gyroidina umbonata (Silvestri)
21
Bulimina aculeata (d’Orbigny)*
20
Table 4: List of species with higher then 20 indicator values in their cluster groups. Indica-
tor values are calculated according to Dufrêne & Legendre (1997). Species with high indica-
tor values ( > 30) are indicated by darker shades.
tive species of this interval is the retic-
ulate bolivinid species Bolivina vien-
nensis, endemic to the Paratethys and,
secondly, the species Lenticulina arcu-
ata. The latter species has low abun-
dances in the material, thus it could be
ignored for ecological interpretation
(Table 4). This unit is characterized by
rather high percentages of inbenthic
and oxyphylic species. The P/B ratio is
similar to the biofacies unit IV below
in estimating shallower paleowater
depth than the gradient analyses. This
interval has moderately high diversity
and thus low dominance (Fig. 2).
Discussion
Trends, proxies and correlation
The correlation of indices and proxies
with core-depth is generally significant,
indicating a changing environmental
scenery during deposition (Table 3).
The growing number of inbenthic
forms and decreasing number of oxy-
phylic taxa upward in the section
points to a eutrophication trend cou-
pled with the stress of low oxygen
content in the bottom water. Species
richness measured by the Fisher-
α
and Shannon index decreased, while
dominance
increased
with
time
(Fig. 2). These trends in diversity are
interpreted
as
enhancing
stress
through the studied time interval, and
can probably be related to the general
eutrophication trend throughout the
Badenian described in Báldi (2006).
Paleowater depth estimations are based on two different
proxies, independent of each other (Van der Zwaan et al.
1990; Hohenegger 2005). The proxy of Van der Zwaan et al.
(1990) is based on the P/B ratio, while the proxy of Ho-
henegger (2005) is based on depth ranges of Recent benthic
foraminifera, thus strongly depending on an actualistic ap-
proach. Both proxies showed a shallowing trend, however a
much stronger trend is postulated according to the plankton-
ic/benthic method. The general trend of shallowing through
time in the Badenian is in accordance with Filipescu & Gir-
bacea (1997) and Báldi et al. (2002). Both proxies behaved
also differently concerning diversity. The P/B-ratio showed
deep water when diversity was high and dominance low,
while the gradient analyses showed no significant correlation
to these indices (Table 3). Thus accepting diversity changes
as a measure of stress, the low diversity means high stress,
most likely to happen near-shore rather than in an open ma-
rine environment. The P/B method is sensitive for eutrophy
causing high productivity surface water (meaning a high
421
PALEOECOLOGY OF BENTHIC FORAMINIFERA OF THE BADEN-SOOSS SECTION (BADENIAN, AUSTRIA)
Fig. 7. The recognized biofacies units (I—V) based on groups ob-
tained by DCA (1—5) and Cluster Analysis (a—f). Samples are de-
noted by black triangles if an actual sample belongs the sample
group. When a sample belongs to a different group, then the triangle
is empty and the group’s assignment is given next to the triangle.
number of planktonic foraminifera) and oxygen deficiency at
the bottom culling benthic life (Van der Zwaan et al. 1990).
In the studied material, episodic lamination and the high
number of planktonic foraminifera ( > 50 % of all foramin-
ifera) without the occurrence of deep water marker benthic
taxa suggests, that paleowater depth is overestimated by the
P/B-ratio method. This overestimation is most likely to be
pronounced in the deeper part of the core. However, under-
estimation of paleowater depth by the P/B ratio through re-
worked foraminiferal tests from shallower parts of the basin
is a possibility, while gradient analyses excludes the depth
ranges of shallow living species (Hohenegger 2005).
The number of benthic foraminifera correlated to core
depth shows an increasing number of benthic foraminifera
with time (Table 3). The foraminiferal number is determined
by two processes, where the one is dilution in sediment de-
pending on sedimentation rates, while the other is benthic
standing stocks and turn over controlling benthic productivi-
ty. Here it is assumed that the latter mentioned benthic pro-
ductivity is responsible of this weak trend through time, as
the number of highly productive inbenthic taxa also increase
with time. The observed eutrophication and shallowing trends
are in accordance with the general trends described throughout
the whole Badenian (Báldi 2006). However, the high number
of benthic foraminifera from 22.4 to 34.0 m – discussed lat-
er as biofacies unit IV – is also related to changes in sedi-
mentation rates.
Ecofacies and geochemistry
Unit I: Periods of stronger lamination are observed in this
interval between 82 m and 92 m. This lamination coincides
with shorter subunits encompassed by the biofacies as docu-
mented by both multivariate methods (Cluster b) and DCA
group 1)). An inbenthic maximum coupled with a relatively
low number of oxyphylic taxa (from 82.6 to 87.6 m) also ex-
ists in this interval (Fig. 2). According to benthic foramin-
ifera, episodic dysoxy or anoxy combined with high food
availability conditions (organic matter) is highly probable
during deposition. These are optimal conditions for the dom-
inating (Fig. 5) and indicative (Table 4) inbenthic species
Uvigerina semiornata. According to the stable isotope
record in this interval, the
δ
18
O values are more negative
measured both in the planktonic and benthic forms, suppos-
edly from higher freshwater influx from land (Fig. 8). This
fact is supported by the magnetic susceptibility data showing
higher terrigenous influence (Fig. 9). The relatively high
number of oxyphylic taxa ( > 30 %), contributing to the rela-
tively high diversity in the laminated part needs further ex-
planation (Fig. 3). There are two alternatives:
1. well oxygenated periods were sampled, missing the per-
haps barren laminated layers;
2. oxyphylic forms were reworked from shallower parts of
the basin.
Signs of reworking were not visible on the foraminiferal
tests, yet it cannot be excluded that marine sediments from
shallower parts of the basin arrived (Holcová 1999). Based
on the benthic foraminiferal results accomplished with the
geophysical and geochemical parameters, we can suppose
the episodic dysoxy and/or anoxy of this period is related to
increased terrigenous influence in the form of lowered salini-
ty surface water.
Unit II: The only indicative species Bolivina hebes is an
endemic species with still unknown ecological demands (Ta-
ble 4). Concerning isotope results, this biofacies unit marked
a rather stable period with slightly increasing
δ
18
O values
measured in planktonic foraminifera and high values mea-
sured in the benthic forms, indicating normal marine salini-
ties (Fig. 8). This refers to a period of stability, lowered food
supply, and better-oxygenated bottom water conditions than
in the previous period. Concerning magnetic susceptibility,
it was also a stable period of moderate terrestrial input
(Fig. 9). Generally, stable conditions and lowered food sup-
ply of this period resulted in relatively high diversities. How-
ever, the oscillations in the P/B paleowater depth proxies
have no plausible explanation at the moment (Fig. 2).
Unit III: A general negative trend of
δ
18
O reaching a min-
imum in the upper part at 36.8 m, especially pronounced in
422
BÁLDI and HOHENEGGER
the benthic
δ
18
O, with a corresponding
δ
13
C record, charac-
terizes this interval. Extracting the planktonic
δ
18
O from the
benthic
δ
18
O as a measure of water column stratification is
created here, referred to as
∆δ
18
O. This difference shows a
definite minimum of the entire studied section at 42.6 m,
meaning minimal density stratification during deposition of
this biofacies (Fig. 8). The magnetic susceptibility shows
maximum terrestrial input throughout the whole core, with
the highest values in the lower part of the unit (Fig. 9). Some
– not too common indicator species (Table 4; Fig. 3) of the
lower part (cluster h)) such as Ammonia beccarii, Quinque-
loculina spp. and div. miliolids are low salinity tolerant taxa.
The species Trifarina angulosa, indicative for most of this
unit (cluster h) and c)), is associated with lower salinities
sometimes, however, more consequently with coarser sub-
strates, well oxygenated, high energy bottom waters (Harloff
& Mackensen 1997; Hayward et al. 2002, 2004; Klitgaard-
Kristensen et al. 2002). On the basis of the peak of Trifarina
angulosa, we suppose well-aerated, turbulent bottom waters
at the time of deposition. This is in accordance with the re-
duced stratification of the period revealed by the
∆δ
18
O mini-
mum. The rather low benthic foraminiferal numbers
characterizing this period are due to dilution in sediment by
Fig. 8. Stable isotope record plotted against depth in core and the recognized biofacies units on the left. From left to right: The stable iso-
tope record of
δ
18
O and
δ
13
C measured in Globigerinoides trilobus and in Hoeglundina elegans. The delta-delta record was calculated to
express differences of surface and bottom water, where the
∆δ
18
O
=
δ
18
O
benthic
—
δ
18
O
planktonic
, while
∆δ
13
C
=
δ
13
C
benthic
—
δ
13
C
planktonic
.
Fig. 9. Magnetic susceptibility record plotted against depth in core
and the recognized biofacies units on the left.
423
PALEOECOLOGY OF BENTHIC FORAMINIFERA OF THE BADEN-SOOSS SECTION (BADENIAN, AUSTRIA)
terrigenous input, also providing the coarse substrate neces-
sary for Trifarina angulosa.
Unit IV: The indicator species Bulimina elongata and
Fursenkoina acuta (Table 4) are typical deep inbenthic spe-
cies possessing elongate tests (Corliss 1985; Corliss & Chen
1988; de Stigter et al. 1998), which are tolerant for episodic
dysoxy, while profiting from high food availability living
close to the redox front in the sediment. Bolivina spathulata
(synonym to Bolivina dilatata in de Stigter et al. 1998 and
Den Dulk et al. 2000) is also a deep inbenthic, while Cassidu-
lina laevigata is observed to be tolerant for reduced oxygen
and enhanced organic carbon content in muddy sediments
(Rogerson et al. 2006). The other indices like oxyphylic per-
centages are low, confirming lowered oxygen levels. The di-
versity indices Shannon (H) and Fisher-
α are low, while
dominance is high due to the low oxygen levels at the bot-
tom causing stress for benthic life. The number of benthic
foraminifera is raised ten folds in this biofacies unit (Fig. 2).
This could be the combined result of r-strategist highly pro-
ductive inbenthic forms (Bulimina elongata, Fursenkoina
acuta and Bolivina spathulata) and the low terrigenous input
shown by magnetic susceptibility not diluting the foramin-
iferal stock in the sediment (Fig. 9).
The stable isotope results (Fig. 8) provide additional infor-
mation about how the highly eutrophic bottom water condi-
tions developed in this biofacies. The carbon isotope record
measured in the planktonic foraminifera (Globigerinoides
trilobus) indicates rather highly productive surface water
conditions sustaining food for the benthic community. The
only available
∆δ
18
O data point of this interval shows a
highly stratified water column, which contributed to oxygen
deficiency at the bottom by restricting vertical circulation.
Unit V: The indicator species, the reticulate Bolivina vien-
nensis, is abundant enough to contribute significant informa-
tion about the environment. Reticulate bolivinids prefer
oxygen depleted bottom water with sustained food supply
and sluggish circulation (Hayward et al. 2002). The stable
isotopes and magnetic susceptibility highly oscillate during
this period (Figs. 8, 9). The
∆δ
18
O is rather high, thus we
can suppose that the intensive stratification of the water col-
umn characterizing the previous unit had persisted. How-
ever, the faunal turnover from biofacies unit IV to V is
probably due to less stable conditions reflected as oscilla-
tions and intensified terrigenous input according to suscepti-
bility. The relatively high percentages of oxyphylic taxa are
due to reworking and intensified transport from land similar-
ly to the anoxic event in biofacies unit I (Fig. 2).
Conclusion
1. The benthic foraminifera of the Baden-Sooss borehole
were found to be a sufficient tool for paleoenvironmental re-
constructions. Results based on foraminifera were confirmed
by stable isotope records (
δ
18
O and
δ
13
C) and magnetic sus-
ceptibility (Figs. 8, 9), thus the actualistic approach com-
bined with the applied species concept worked.
2. The different indices and proxies (Fig. 2) showed strong
correlation with core depth, meaning apparent trends (Ta-
ble 3). The applied multivariate techniques resulted in sam-
ple groupings or species assemblages that also characterize
distinct depth intervals of the section (Figs. 4, 6, 7). These
observations suggest that a succession of changing environ-
mental regimes is manifested in the benthic foraminiferal
record.
3. The correlation to core depth revealed an increasing in-
benthic percentages coupled with a slightly decreasing oxy-
phylic number and declining diversity (Table 3, Fig. 2). This
is interpreted as lowering oxygen levels and increasing food
resources causing increasing stress observable through the
studied section. This trend can be related to the general
eutrophication trend throughout the Badenian (Báldi 2006).
The paleowater depth proxies document a shallowing trend.
4. The boundaries of the five recognized biofacies units
(Fig. 7) based on multivariate techniques correlate to shifts in
the stable isotope and susceptibility record (Figs. 8, 9). This
suggests, that changes in the physical environmental parame-
ters (oxygen, food, salinity) inducing a turnover of benthic
foraminiferal assemblages are controlled by proximity to the
land. This idea is in accordance with the general geological
setting of the Baden-Sooss borehole located in a rather shal-
low part of the Paratethys not too far from the land.
Acknowledgments: This contribution benefited from the fi-
nancial support of the Austrian Science Foundation FWF
Project P16793-B06. Thanks are due to the whole group
working in the above project, especially to Christian Rupp
and Stjepan Ćorić (Geological Survey, Wien), Fred Rögl
(Natural History Museum, Wien), Anna Selge, Robert
Scholger (Institute of Geophysics, Montan Universität Le-
oben), Maksuda Khatun, Michael Wagreich (Department of
Geodynamics and Sedimentology, Universität Wien), Peter
Pervesler (Institute of Palaeontology, Universität Wien) and
Nils Andersen (Leibniz Laboratory, CAUniversity Kiel).
Special thanks are also due to Christian Baal (Vienna) for
making SEM micrographs.
References
Altenbach A.V., Lutze G.F., Schiebel R. & Schönfeld J. 2003: Im-
pact of interrelated and interdependent ecologic controls on
benthic foraminifera: an example from the Gulf of Guinea.
Palaeogeogr. Palaeoclimatol. Palaeoecol. 197, 213—238.
Báldi K. 1999: Taxonomic notes on benthonic foraminifera from
SW-Hungary, Middle Miocene (Badenian), Paratethys. Acta
Geol. Hung. 42, 193—236.
Báldi K. 2006: Paleoceanography and climate of the Badenian Central
Paratethys (Middle Miocene 16.4—13.0 Ma) based on foramin-
ifera and stable isotope evidence. Int. J. Earth Sci. 95, 119—142.
Báldi K., Benkovics L. & Sztanó O. 2002: Badenian (Middle Mi-
ocene) basin development in SW Hungary: Geohistory based
on quantitative paleobathymetry of foraminifera. Int. J. Earth
Sci. 91, 490—504.
Cicha I., Rögl F., Rupp C. & Čtyroká J. 1998: Oligocene—Miocene
foraminifera of the Central Paratethys. Abh. Senckenberg.
Naturforsch. Gesell. 549, 1—325.
Corliss B.H. 1985: Microhabitats of benthic foraminifera within
deep-sea sediments. Nature 314, 435—438.
Corliss B.H. & Chen C. 1988: Morphotype patterns of Norwegian
424
BÁLDI and HOHENEGGER
Sea deep-sea benthic foraminifera and ecological implications.
Geology 16, 716—719.
Dai X., Page B. & Duffy K.J. 2006: Indicator value analysis as a
group prediction technique in community classification. South
African J. Botany 72, 589—596.
Den Dulk M., Reichart G.J., Van Heyst S., Zachariasse W.J. & Van
der Zwaan G.J. 2000: Benthic foraminifera as proxies of organic
matter flux and bottom water oxygenation? A case history from
the northern Arabian Sea. Palaeogeogr. Palaeoclimatol. Palaeo-
ecol. 161, 337—359.
De Stigter H.C., Jorissen F.J. & Van der Zwaan G.J. 1998: Bathymetric
distribution and microhabitat partitioning of live (Rose Bengal
stained) benthic foraminifera along a shelf to bathyal transect in
the southern Adriatic Sea. J. Foraminiferal Res. 28, 40—65.
d’Orbigny A.D. 1846: Foraminifères fossiles du Bassin Tertiaire de
Vienne (Austriche). Gide et Comp, Paris, XXXVII +1—312.
Dufrêne M. & Legendre P. 1997: Species assemblages and indicator
species: The need for a flexible asymmetrical approach. Eco-
logical Monographs 67, 345—366. doi:10.2307/2963459
Filipescu S. & Girbacea R. 1997: Lower Badenian sea-level drop on
the western border of the Transylvanian Basin: Foraminiferal pa-
leobathymetry and stratigraphy. Geol. Carpathica 48, 325—334.
Fisher R.A., Corbet A.S. & Williams C.B. 1943: The relationship
between the number of species and the number of individuals
in a random sample of an animal population. J. Animal Ecolo-
gy 12, 42—58.
Fontanier C., Jorissen F.J., Licari L., Alexandre A., Anschutz P. &
Carbonel P. 2002: Live benthic foraminiferal faunas from the
Bay of Biscay: Faunal density, composition, and microhabi-
tats. Deep Sea Research Part I: Oceanographic Res. Pap. 49,
751—785.
Grill R. 1941: Stratigraphische Untersuchugen mit Hilfe von Mik-
rofaunen im Wiener Becken und den benachbarten Molasse-
Anteilen. Öl Kohle 31, 595—603.
Hammer O. & Harper D. 2005: Paleontological data analysis.
Blackwell Publishing, Malden, MA, 1—351.
Harloff J. & Mackensen A. 1997: Recent benthic foraminiferal as-
sociations and ecology of the Scotia Sea and Argentine Basin.
Mar. Micropaleontology 31, 1—29.
Hayward B.W., Neil H.L., Carter R., Grenfell H.R. & Hayward J.J.
2002: Factors influencing the distribution patterns of Recent
deep-sea benthic foraminifera, east of New Zealand, Southwest
Pacific Ocean. Mar. Micropaleontology 46, 139—176.
Hayward B.W., Grenfell H.R., Carter R. & Hayward J.J. 2004:
Benthic foraminiferal proxy evidence for the Neogene palae-
oceanographic history of the Southwest Pacific, east of New
Zealand. Mar. Geol. 205, 147—184.
Hayward B.W., Grenfell H.R., Sabaa A.T. & Neil H.L. 2007: Fac-
tors influencing the distribution of Subantarctic deep-sea
benthic foraminifera, Campbell and Bounty Plateaux, New
Zealand. Mar. Micropaleontology 62, 141—166.
Hohenegger J. 2005: Estimation of environmental paleogradient
values based on presence/absence data: a case study using
benthic foraminifera for paleodepth estimation. Palaeogeogr.
Palaeoclimatol. Palaeoecol. 217, 115—130.
Hohenegger J., Ćorić S., Khatun M., Pervesler P., Rögl F., Rupp C.,
Selge A., Uchmann A. & Wagreich M. 2008: Cyclostratigraph-
ic dating in the Lower Badenian (Middle Miocene) of the Vi-
enna Basin (Austria) – the Baden-Sooss core. Int. J. Earth
Sci., DOI 10.1007/s00531-007-0287-7.
Holcová K. 1999: Postmortem transport and resedimentation of for-
aminiferal tests: Relations to cyclical changes of foraminiferal
assemblages. Palaeogeogr. Palaeoclimatol. Palaeoecol. 145,
157—182.
Jorissen F.J., de Stigter H.C. & Widmark J.G.V. 1995: A conceptu-
al model explaining benthic foraminiferal microhabitats. Mar.
Micropaleontology 22, 3—15.
Kaiho K. 1994: Benthic foraminiferal dissolved-oxygen index and
dissolved oxygen levels in the modern ocean. Geology 22,
719—722.
Kaiho K. 1999: Effect of organic carbon flux and dissolved oxygen
on the benthic foraminiferal oxygen index (BFOI). Mar. Mi-
cropaleontology 37, 67—76.
Klitgaard-Kristensen D., Sejrup H.P. & Haflidason H. 2002: Distri-
bution of recent calcareous benthic foraminifera in the north-
ern North Sea and relation to the environment. Polar Res. 21,
275—282.
Kouwenhoven T.J. & Van der Zwaan G.J. 2006: A reconstruction of
late Miocene Mediterranean circulation patterns using benthic
foraminifera. Palaeogeogr. Palaeoclimatol. Palaeoecol. 238,
373—385.
Magurran A.E. 2004: Measuring biological diversity. Blackwell
Publishing, 1—114.
Mandić O., Harzhauser M., Spezzaferri S. & Zuschin M. 2002: The
paleoenvironment of an early Middle Miocene Paratethys se-
quence in NE Austria with special emphasis on paleoecology
of mollusks and foraminifera. Geobios 35, 193—206.
Marks P., Jr. 1951: A revision of the smaller foraminifera from the
Miocene of the Vienna Basin. Contr. Cushman Found. Foram.
Res. 2, 33—73.
McCune B. & Mefford M.J. 1999: PC-ORD. Multivariate analysis
of ecological data, version 4. MjM Software Design, Gleneden
Beach, Oregon, USA, 1—237.
Nahmani J., Lavelle P. & Rossi J.-P. 2006: Does changing the taxo-
nomical resolution alter the value of soil macroinvertebrates as
bioindicators of metal pollution? Soil Biology & Biochemistry
38, 385—396.
Papp A. & Schmid M.E. 1985: Die fossilen Foraminiferen des tertiären
Beckens von Wien. Abh. Geol. Bundesanst.,Wien 37, 1—311.
Papp A., Cicha I., Seneš J. & Steininger F. 1978: Chronostratigra-
phie und Neostratotypen: Miozän der Zentralen Paratethys. Bd.
VI. M
4
, Badenien (Moravien, Wielicien, Kosovien). VEDA
SAV, Bratislava, 1—594.
Rogerson M., Kouwenhoven T.J., van der Zwaan G.J., O’Neill B.J.,
van der Zwaan C.J., Postma G., Kleverlaan K. & Tijbosch H.
2006: Benthic foraminifera of a Miocene canyon and fan. Mar.
Micropaleontology 15, 295—318.
Rögl F. 1996: Stratigraphic correlation of the Paratethys Oligocene and
Miocene. Mitt. Gesell. Geol. Bergbaustud. Österr. 41, 65—73.
Rupp C. 1986: Paläoökologie der Foraminiferen in der Sandschaler-
zone (Badenien, Miozän) des Wiener Beckens. Beitr. Paläont.
Österr. 12, 1—95.
Shannon C.E. 1948: A mathematical theory of communication. The
Bell System Technical J. 27, 379—423, 623—656.
Simpson E.H. 1949: Measurement of diversity. Nature 163, 688.
Spezzaferri S. 2004: Foraminiferal paleoecology and biostratigra-
phy of the Grund Formation (Molasse Basin, Lower Austria).
Geol. Carpathica 55, 155—164.
Spezzaferri S., Corić S., Hohenegger J. & Rögl F. 2002: Basin-scale
paleobiogeography and paleoecology: An example from Kar-
patian (Latest Burdigalian) benthic and planktonic foramin-
ifera and calcareous nannofossils from the Central Paratethys.
Geobios 35, supplement 1, 241—256.
Van der Zwaan G.J., Jorissen F.J. & de Stigter H.C. 1990: The
depth dependency of planktonic/benthic foraminiferal ratios:
Constraints and applications. Mar. Geol. 95, 1—16.
Van der Zwaan G.J., Duijnstee I.A.P., Den Dulk M., Ernst S.R.,
Jannink N.T. & Kouwenhoven T.J. 1999: Benthic foramini-
fers: proxies or problems?: A review of paleocological con-
cepts. Earth Sci. Rev. 46, 213—236.
Williams C.B. 1964: Patterns in the balance of nature. Academic
Press, London, 1—324.
De
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GE
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CA
CA
RPA
THI
C
A
, 5
9
, 5
(2008);
B
Á
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D
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and
HOHENEGGE
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:
PAL
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OF
B
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BA
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SECTI
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,
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1
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4
www.geologicacar
pathica.sk
Tabl
e 1:
Abunda
nce of
benthic for
aminifera in 100
g sediment of the
Baden-Sooss
core
.
Part 1
from 4
.
De
pt
h i
n
c
o
re (m)
Cera
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s ha
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(d
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Ci
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(W
alker & Ja
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(Parke
r)**
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ic
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Cibic
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p.
Co
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ny
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rev
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(d
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ny
)
De
nt
al
ina e
le
g
an
s
(d
’O
rb
ig
ny
)
E
h
re
n
b
er
g
ia se
rr
at
a
Reuss
El
ph
idi
um
spp
.
Al
ab
ami
n
a
sp
.
Fi
ss
ur
ina
sp
p
.
F
u
rs
en
ko
ina ac
ut
a
(d’
O
rb
ign
y
)*
Gavelinopsis
pra
egeri
(
H
eron-Al
len
&
Earla
nd)
G
lan
dul
ina
sp
.
G
lob
obu
lim
ina
p
yru
la
(
d
’O
rbi
gny
)*
G
lob
uli
na
sp
.
G
utt
ul
ina
sp
. (d’O
rbig
ny
)
Gy
ro
id
ina p
a
rv
a
(Cus
hman
&
Renz)
G
yr
o
idin
a
so
ld
ani
i
(d’
O
rb
ign
y
)
G
yr
o
idin
a
um
bon
ata
(Silve
stri)
H
ans
awai
a
bo
ueana
(
d
’O
rb
ig
ny
)**
He
ter
o
le
pa du
te
m
p
le
i
(d
’O
rbig
ny
)*
*
H
o
eglu
nd
ina
elegan
s
(d
’O
rbig
ny
)
8.410
30
1
1
3
11
1
2
22
10.015
10
3
4
33
1
1
2
6
1 15
11.210
9
5
6
1
14
1
2
1
4
12.440
36
3
15
2
14
1
9
13.620
22
5
3
18
2
2
2
4
1
14.810
26
11
5
2
16
16.410
1
9
2
1
8
4
23
17.610
2
42
2
3
21
24
18.880
4 12
9
5
1 13
9
1
3
6 14
20.020
2
33
4
1
2
9
1
6
18
22.410
5
8
5
1
2
20
23.610
7
7
2
7
1
1
4
24.855
7
2
3
3
10
2
4
26.410
8
3
5
3
3
5
27.615
14
1
4
2
6
5
1
1
11
28.855
2
25
5
17
4
1
1
1
32
30.010
1
22
6
3
1
10
5
2
5
2
10
31.215
13
2
7
5
1
1
1
25
34.010
2
1
7
1
1
2
35.210
55
11
3.5
11 4
2 3
5
16 5 4 1
36.810 2
22 7 4
12 4 6
2
4
3 4 3 6 2
37.610
10
5
2
2
2
1
1
2
13
5
2
1
3
3
38.810
3 14
5
5
1
3
5
1
14
1
2
5
6
1
40.010
1
6
3
3
1
1
4
51
3
4
8
41.210
8
12
3
5
5
1
4
14
5
1
1
1
2
7
5
42.410
2
13
4
1
1
1
14
1
1
4
1
1
42.610
4
9
3
1
1
2
2
14
8
1
5
43.810 3
16 3 2
2 2 1
9
1
2 2 1 4 3
45.010
2 12
5
1
3
3
10
1
1
2
46.410
2
6
5
2
8
3
1
1
3
1
47.610
2
4
1
3
1
7
2
3
1
1
2
48.810
2
8
1
2
6
1
4
3
3
50.010
2
5
4
1
6
2
1
1
51.410
3
1
4
1
2
12
1
1
1
52.610
2
1
1
1
11
4
3
1
4
53.810 6
11
1
2 8 2 2 7
1
2 1 2 1
55.010 6 9
2
1 3 2 3 8 1 2
1
3 3 3 8
56.410 13 13
1
2
11
1
5
3
2
1
1
57.610
5
17
3
2
3
9
5
1
5
5
4
3
58.810 1
13
1
1.5
1 1
7 1 3
2 6 1 1 5
60.010
17
1
1
1
15
1
6
2
9
1
4
4
61.410
6
17
4
1
14
1
6
5
5
9
3
5
62.610 8
18
2
3 1 2 3
17
3 8 6 3 1
63.815
16 3 1 4
2
1 1 2 5
3 1
12 6
65.010 17 10
3
2
1
4
4
3 12
1
5
2
4
7
66.415 4 7 1 3
2 3 6 2
5
1
7
2 4
67.61
10
8
3
1
1
1
2
9
3
7
3
1
4
1
4
7
68.810 4
11 2 1
3 2 3 3 8
1
5 4 9 3
70.010
15
2
3
7
1
10
5
6
2
2
71.415
13
1
6
6
6
5
13
15
5
1
72.61
2 10
4
2
3
4
2
3
1
1
2
6
1
6
2
73.810
8
1
4
1 3 5 1 2
9 3 2 3
75.010 4
36
3
5
10 1 2 9
1
4 8 7 5 1
76.410 4
21 1 6
1
1 1
13 4 2 1
14
3
1
6
16
13
13 7
77.610 6
15
3
1 1 5
1 4
3
2 2 7 4 9 4
78.810
3
11
2
9
2
1
3
3
8
5
6
22
6
80.010 6
31
4 4 1 1
16
2
1 3 6
9
1 2 3
10 9 6
81.410 5
17 2 4
2 1
7 1 1 1 8
10
1
6 2 3
82.610
2 16
3
1
1
6
2
4
1
2
1
6
2
4 12
2
7
83.830
1
24
1
4
2
3
3
1
4
9
1
5
8
85.010
4 14
5
2
3
2
2
2
4
2
1
2
5
4
8
86.410
1 16
4
8
1
1
2
1
3
7
4
4
5
5 10
87.610 4
20 2 1 2 2
3
1 1 5
1
2
12 2 1
88.810 1
21
7 1 1
3
1 2 1 1 2
3
2 6 3 2 6 6
90.010
15 4 4
3
2 2 1 8
7
2 2 1 5 9 4
18
91.410 1
10
3
2 2 4 1 1 7
10
1 3 8 3
14
92.630
8
27
2
5
1
1
6
3
11
2
10
3
93.810
7 25
10
3
2
1
5
3
17
6
1
95.010 15 20
10
2
2
9
9
4 13
3
2
96.410
15
19 2 2
11 2 3 3 3
5
7 2 8 7
97.610 13 27
2
9
2
3
8
8
1
4
3 12
1
1
98.810
11
14 1 2
11 4 2 3
4
1 5 6 6
100.010 12 13
10
3
3
3
2
8
4
6
4
4
101.410
4
6
3 11
2
4
2
4
5
6
7
BÁL
DI
a
nd H
O
H
E
NE
G
G
E
R
: PAL
EO
EC
OL
OG
Y O
F
BE
NTHI
C F
O
R
A
MINIF
ERA
OF
THE
B
A
D
E
N
-SO
O
SS
SE
CTI
O
N; E
L
ECT
R
ONIC
S
U
PPL
EME
N
T
Table 1:
Continued.
Part 2 from 4.
E2
De
pt
h i
n
c
o
re (m)
La
gena
spp
.
Lent
ic
ul
ina
cal
ca
r (L
inné)
*
*
Lent
ic
ul
ina
ar
cu
ata
(d
’O
rbig
ny
)*
*
Lent
ic
ul
ina
sp
.**
Meloni
s pompil
ioides
(Ficht
el
& Moll
)*
A
d
el
os
ina
spp.
**
Pyr
g
o
spp
*
*
Tr
ilo
culi
na
spp
.**
Q
uin
qelo
cu
lin
a
sp
p.
*
*
R
o
sa
li
na
Spir
o
loculina excavata
(d'Orbigny)
**
S
igm
oil
in
ita
t
enu
is
(
C
zjze
k)
**
Mi
li
ol
id
sp
p.
*
*
Ma
rg
inu
lin
a
h
ir
sut
a
(d
'O
rbig
ny
)
No
do
sa
ri
a
sp.
1
No
do
sa
ri
a
sp.
2
No
do
sa
ri
a ra
p
h
a
n
is
tr
um
(L
inné)
No
ni
on co
m
m
u
n
e
(d
'O
rb
ig
ny
)
Non
io
n
ella
tu
rg
ida
(
W
il
lia
ms
o
n
)
O
oli
na
spp
.
O
rid
or
sal
is
umbo
na
tu
s (R
euss)
Pl
an
uli
na
sp
.
Pu
ll
enia
bu
llo
id
es
(d
'O
rb
ig
ny
)*
Pu
ll
enia
qu
inq
uelob
a
(Reu
ss)
*
Reu
sell
a
s
pin
ulo
sa
(
R
eu
ss)
8.410
2 26
4
1
2
1
1
3
10.015
1
2
2 18
1
1
3
9
1
1
1
11.210
2
8
3
2
22
1
12.440
1
23
2
7
5
1
13.620 1
1
17
2
1
4
3
14.810
2
13
3
4
19
2
16.410
2
8
1
2
47
3
17.610
1
15
1
1
3
1
13
1
18.880
2
6
4
1
1
3
20.020
1
1
20
3
1
2
1
2
1
1
3
22.410
2
3
1
50
2
23.610
4
1
1
1
41
24.855 1
1
3
3
49
3
26.410
3
2
37
27.615
4
1
1
38
2
28.855
1 10
1
1
1
2
1 35
1
1
5
30.010
4
36
3
31.215
2
4
1
11
4
1
34.010
1
2
1
1
4
4
5 103 2
1
1
35.210
5 21
5
1
1
1
3
5
1
36.810
3 4 8 4
7 3
2 4 1 1 2
1
1
37.610
2
3
5
5
1
1
3
5
2
2
2
2
38.810
6
5
4
4
1
1
3
2
1
4
1
1
1
1
40.010
1
9
1
1
1
2
1
1
3
4
41.210
3
10
1
1
3
1
7
2
3
42.410
3
3
1
2
4
2
2
2
4
1
1
42.610
2
3
10
1
1
1
2
3
1
2
2
1
43.810 1 1
6
2 4
2 3 2 1
1 1
2 1
45.010 4 4
1 7 2 3 1 3
2 3 3
11
4
46.410
2
2
4
1
2
11
1
5
1
41
1
47.610 1 2
6
10 6 3
5
1 3 6
1 1
39
2 1 1
48.810
1
2
7
3
3
2
2
1
19
1
1
50.010
4
2 14
5
5
8
4
2
63
2
51.410 2
2
10
1
4
34
52.610
6
1
5 16
1
1
3
1
36
1
2
4
53.810
1
6
6
4
1
1
9
3
2
55.010 3
4
10
1
1
3
11
1
56.410
1
7
1
7
1
9
1
1
57.610
3
2
7
3
1
4
10
1
1
1
58.810
4
8
2
1
14
7
60.010 2
8
14
2
2 1 2
1 1
15 2
1
4 2
61.410
1
14
1 1
2 1
1 2
33 1
1
1
62.610
2
2
7 13
4
4
1
1
9
1
1
63.815
19 18
2
4
2
3
1
1
1 12
1
65.010
5
1 13
4
5
2
1
2
19
1
2
2
2
66.415
8
6
1
1
1
1
8
2
6
67.61
7
13 11
2
4
1
5
7
2
68.810
3 12
1
1
2
2
1
6
2
5
70.010
2
1
3 21
4
1
3
1
3
2
2
71.415
3
17
5
17
12
1
72.61
4
4 17
2
2
1
2
1 13
1
2
1
10
73.810
3
1 10
2
5
1
1
7
1
15
2
6
75.010 11
5
25
3 1 2 3 2
2 2
6 1 1
76.410
3 20
1
11
2
4
3
8
4
77.610 5
2
9
1
6
1
3
1
7
2
78.810
2
2
7 16
2
4
2
1
5
1
1
1
80.010 2
12
11
2
2
3
4
1
1
1
1
81.410 2
3
13
1
1
3
1
5
3
1
1
2
82.610 4
5
6
1
1
2
8
3
2
1
1
83.830
4
21
1
2
1
3
3
1
1
2
85.010
2
1
7 10
1
1
1
2
1
13
1
2
86.410
2 3
2 2 2
6 3 2 1 1
1
1
87.610
1
1
9 14
2
1
6
2
2
1
3
4
88.810 2
8
6
3
5
1
1
2
1
1
3
1
90.010
8
1
2
3
2
3
1
2
1
2
91.410
5
14
2
2
1
4
1
1
2
92.630
5
2
1
1
1
2
1
1
2
3
93.810
2
1
2
3
1
1
1
3
1
1
95.010
2 3
6
3 2 3 1 1
2 6 1
4
2
96.410
7
5
2
1
1
6
1
1
97.610
3
3
4
1
6
4
1
4
5
98.810
2
1
2
5
1
4
3
8
2
4
100.010
5
3
5
3
1
4
1
1
16
1
2
1
101.410
3
8
6
2
1
3
1
2
1
BÁL
DI
a
nd H
O
H
E
NE
G
G
E
R
: PAL
EO
EC
OL
OG
Y O
F
BE
NTHI
C F
O
R
A
MINIF
ERA
OF
THE
B
A
D
E
N
-SO
O
SS
SE
CTI
O
N; E
L
ECT
R
ONIC
S
U
PPL
EME
N
T
Tabl
e 1:
Continued.
Part 3 from 4.
E3
De
pt
h i
n
c
o
re (m)
S
iph
oni
na
reti
cu
la
ta
(Czjz
ek)
**
S
pha
eroi
din
a
bu
ll
oides
(d'
O
rb
igny)*
S
ti
los
tom
el
la
ado
lph
in
a
(d
'O
rb
ig
ny
)
Tr
ifa
ri
na
a
ngu
lo
sa
(
W
il
lia
ms
o
n
)
Tr
ifa
ri
na
b
rad
yi
(C
ush
m
an
)
Pa
pp
ina
pa
rkeri
(
K
ar
rer)*
Uv
ige
rin
a s
em
ior
n
a
ta
(d
'O
rb
ign
y)*
Uv
ige
rin
a p
yg
m
ea
(d
'O
rb
ig
ny
)*
Uv
ige
rin
a ve
n
u
st
a
(Fr
an
zen
au
)*
U
vi
g
eri
n
a
acul
ea
ta
(d
'O
rb
igny
)*
U
vi
g
eri
n
a
acumi
nat
a
(Hosius)
*
Uvig
erina
acuminata
f.
macro
carinata*
Uv
ige
rin
a
spp
.*
Va
gi
nul
ops
is
p
edum
(d
'Orbi
gny
)
Va
lv
u
lin
er
ia
co
mpl
ana
ta
(d
'O
rbi
gny
)*
Unde
ter
m
ine
d
ca
lcar
eous
spe
cimens
H
apl
oph
rag
m
oi
des
sp
.
Ma
rt
ino
ti
el
la
co
mmun
is
(d
'O
rbi
gny
)
S
pir
op
lectamm
ina
spp
.
Spiropl
ectammina
car
inata
(d
'O
rbigny)
Textu
lar
ia
sp
.
Textu
lar
ia
spp
.
Textu
lar
ia
cf.
mexic
a
na
S
igm
oil
op
sis
sp
.
V
u
lv
ul
ina
sp
p.
Undet
ermined
agglutinated
specimens
8.410
4
4
8
1
6
2
1
2
3
10.015
2 5
3
6
1
9
3
3
1
1
4
2
11.210
1
14
25
1 1
18
3
2
1
12.440
2
14
9
2
1
15
6
5
3
13.620
5
3
8
27 3
2
8
4
1
1
14.810
3
2
1
23
18
3
2
5
2
2
16.410
1
12
8
2
7
1
2
17.610
3
1
19
5
4
22
2
2
1
18.880
4
1
9
2
7
3
9
1
1
20.020
2
22
17
2
7
2
2
22.410
1
19
2
3
1
23.610
1
1
20
3
1
5
2
2
24.855
9
1
3
5
26.410
9
3
5
1
27.615 1
7
5
3
19
1
28.855
3
6
15
1
23
2
2
4
30.010
2
6
12
5
13
3
4
31.215
20
14
3
33
2
4
1
34.010
1
1 5
4
16 2
2.5
21
9
2
4
35.210
1
1
5
3
18
5
4
1
8
4
2
4
36.810
13
11 2
2
5 3
1
5
7
1
4
37.610
4 2
2
14
9
0.3
3
4
3
38.810 2
1 2
6
19
6
2
3
2
1
40.010
2
10
20
6
3
2
5
1
41.210 1
1 3
7
25
3
1 1
10
2
2
1
2
42.410
1 2
20
1
2
5
3
2
4
1
1
4
1
42.610
2
27
10
1
1
1
2
4
43.810
2
32
1
21
1
5
1
1
4
45.010
1
26
1
1
1
18 9 1
2
5
3 3 1 1 4
46.410
16
2
9
6
8
2
4
1
47.610
2
8
2
12
4
5
4
3
3
4
48.810
1
16
3
13
4
1
9
5
3
2
8
1
4
50.010
4
27
1
2
7
3
6
1
3
51.410
20
1
1
11
6
2
2
52.610
7
28
2
1
13
1
3
2
2
53.810
4
1
1
10
6
1
13
1
1
55.010
1
3
2
7 1
23 5
3
21 4
2 3 2 3
56.410
3
2
15
4
1
3
2
4
2
3
57.610
1
9
1 2
5
2
2
1
3
58.810
1 3
2
2
12
6
10
11
3
3
1
60.010
1 5
7
3
4
4
10
9
1
3
1
2
4
61.410
2
5
14
3
2
6
1
2
4
62.610
3
10
1
18
4
1
8
6
1
1
1
63.815
1
1
1
12
5
5
21
10
5
11
4
65.010
4
2
17
4
2
5
4
5
2
4
66.415
1
19
7
5
1
1
2
5
67.61
1
1
1
5
5
2
6
1
1
1
1
1
68.810
1
2
7
7
1
14
4
2
1
4
1
70.010
1
3
1
9
3
14
3
1
1
2
71.415
5
14
2
3
15
2
4
6
2
72.61
7
1
1
5
5
10
2
2
1
3
73.810
2
2
1
6
25
1
12
5
2
75.010
4
4
3
1
3
1
7
6
76.410
1
5
1
4
11
13
2
11
3
4
3
4
8
3
77.610
1
8
5 2
3
9 4
6 2
2 1 2
4
78.810
5
2
6
2
7
5
7
2
2
2
1
1
3
1
80.010
1
2 11 3
2
12 9
3
2
2
2 1 2
81.410 1
2 2
1
2
6
12
1
3
3
6
1
82.610 2
6
10
3
15
2
9
7
83.830
2 4
7
1
27
12
5
1
3
2
1
4
85.010
2 3
4
4
1
13
12
6
3
1
1
2
2
86.410 3
6 5
4
2
26
7
2
2
2
1
1
2
87.610 1
3 9
2
4
12
11
1
1
1
4
1
88.810 1
10
1
1
23
9
7
6
2
2
1
1
90.010
1
6
13
2
1 6
1
4
5 2 1
2
91.410
1 1
4
9
13
6
2
6
3
3
1
2
2
92.630
6 5 3 2
1
6
4
1 6
1
1 2
7
93.810
2
4.5
8
2
10
2
1
9
1
1
1
1
95.010
2
0.5
4
23
6
1
4
3
3
5
2
4
96.410
1 1
1
12
4
6
2
3
6
1
3
6
97.610
1 2
3
36
4
4
2
2
10
1
2
2
98.810
4
5
15
3
9
1
5
2
5
2
100.010
6
3
30
4 5
1 1 2
2
3
101.410
7
79
1
2
5
1
3
4
7
2
3
BÁL
DI
a
nd H
O
H
E
NE
G
G
E
R
: PAL
EO
EC
OL
OG
Y O
F
BE
NTHI
C F
O
R
A
MINIF
ERA
OF
THE
B
A
D
E
N
-SO
O
SS
SE
CTI
O
N; E
L
ECT
R
ONIC
S
U
PPL
EME
N
T
Tabl
e 1:
Continued
.
Part 4 from 4.
E4