ELSEVIER
Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
Taphonomic experiments on marginal marine foraminiferal
assemblages: how much ecological information is preserved?
John W. Murray a,Ł , Elisabeth Alve b
a
School of Ocean and Earth Science, Southampton Oceanography Centre, European Way, Southampton SO14 3ZH, UK
b Department of Geology, University of Oslo, PO Box 1047 Blindern, N-0316 Oslo, Norway
Received 10 December 1996; revised version received 3 July 1997; accepted 8 June 1998
Abstract
The most important taphonomic processes affecting the composition of benthic foraminiferal assemblages are transport
and destruction of tests. The latter includes dissolution of calcareous tests. We have previously simulated nature
experimentally by dissolving mainly shelf to deep sea samples in weak acid to give acid-treated assemblages (ATA). In
this study we sampled the marginal marine environments of the Skagerrak=Kattegat and found that the main taphonomic
process there is dissolution of calcareous tests. Consequently, the taphonomic processes occurring in this area represents
a perfect natural analogue of what we suspect to be the main mechanism for the formation of some fossil agglutinated
assemblages. The evidence for this is that some living assemblages dominated by calcareous forms become original
dead assemblages (ODA) often consisting entirely of agglutinated species and equivalent to ATAs. The principal ATA is
dominated by Miliammina fusca and this ATA can be derived from 10 different types of ODA including those that are
strongly dominated by various calcareous taxa. This residual M. fusca assemblage represents a broad range of intertidal
to shallow subtidal environments. On the other hand, marsh living assemblages are completely agglutinated and obviously
give rise to agglutinated ODA=ATAs but not necessarily dominated by the same taxa. All the ATAs faithfully record
the low species diversity characteristic of marginal marine environments. The principal conclusion is that in spite of
the severe taphonomic loss, the preserved assemblages record much ecological detail. This is in accordance with similar
investigations of deeper water assemblages and has important consequences for palaeoecology. This study demonstrates
unequivocally that it is always essential to compare data on living and dead assemblages in order to determine the pathways
to fossilisation. The use of total (living plus dead) assemblages in this area would give unrepresentative and misleading
results.  1999 Elsevier Science B.V. All rights reserved.
Keywords: taphonomy; agglutinated foraminifera; ecology; marginal-marine; dissolution
1. Introduction
Palaeoecological studies are normally carried out
by comparing fossil assemblages with their modern
Ł Corresponding
author. Fax: C44-1703-593052; E-mail:
jwm1@mail.soc.soton.ac.uk
analogues. Although fossil assemblages consisting
exclusively of non-calcareous, organo- and ferroagglutinated foraminifera are known from Mesozoic
and Cenozoic rocks (for example, Charnock and
Jones, 1990), the only modern analogues appear to
be either intertidal marshes or seas deeper than the
calcium carbonate compensation depth (Alve and
0031-0182/99/$ – see front matter  1999 Elsevier Science B.V. All rights reserved.
PII: S 0 0 3 1 - 0 1 8 2 ( 9 8 ) 0 0 2 0 0 - 4
184
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
Murray, 1995). It seems unlikely that all fossil examples fit into one or other of these two extreme
environments so that has limited their interpretation.
A reasonable assumption is that some of the fossil
agglutinated assemblages are the residuum of diagenetic dissolution of the calcareous component.
Therefore, in the past few years we have carried out
experimental studies to help to determine the origin
of agglutinated fossil assemblages (Alve and Murray,
1994, 1995; Murray and Alve, 1994, in press). We
simulated natural dissolution of the original dead assemblage (ODA) by treating the samples with dilute
acid and called the resultant residue the acid-treated
assemblage (ATA). The initial studies considered environments ranging from deep sea, through slope
and shelf, to intertidal but with only 8 samples from
shallow marginal marine settings.
The coastal zone of the Skagerrak=Kattegat is an
ideal area in which to investigate shallow marginal
marine subenvironments in more detail. These range
from small areas of marsh with various combinations
of Phragmites, Carex and Salicornia developed in the
sheltered parts of bays, through intertidal flats to subtidal areas, with or without seagrass. The area is microtidal with an astronomical tidal range of 20–30 cm
along the Skagerrak and about 40 cm along the Kattegat coast. However, meteorological effects (surge,
caused by wind and barometric pressure) generally
have a stronger impact on the prevailing sea level
in this area than the astronomical tide implying that
ranges of about 1 m occur several times a year (R.
Braaten, pers. commun., 1997). However, the general
range of variation is probably less than 0.5 m with
the marshes being developed at the upper limit. These
shallow waters generally lack currents and develop
stratification (at least during the summer). At most
stations, there is low input of freshwater from the surrounding land. The annual temperature range is from
at least 0 to 28ºC and salinities ranged from 10 to
30‰ at the time of sampling. From previous studies
by Alexandersson (1979) it is known that carbonate
dissolution is active in this area.
In order to evaluate the ATA results, it was essential to investigate the ecology and distribution
of the living assemblages (Alve and Murray, 1999)
and the taphonomy of the ODAs (Murray and Alve,
1999). The most important taphonomic processes affecting the composition of benthic foraminiferal as-
semblages are transport and destruction of tests, the
latter including dissolution of calcareous tests. Longdistance transport is not significant here; all the dead
taxa are represented by living individuals and there is
no introduction of exotic species. This is consistent
with the small microtidal regime. There is some local
transport due to waves. However, natural dissolution
of calcareous tests is shown to be of major importance in this area. This makes it a very worthwhile
natural laboratory in which to study taphonomy and
the origin of agglutinated assemblages.
2. Material and methods
From 1994 to 1996, 171 samples were collected
from 27 localities along the Skagerrak=Kattegat
coast from the intertidal zone (including marsh) to
a water depth of 6 m (Fig. 1) but two (162, 163)
were not used in this study. Because of the irregular
tidal pattern, it was impossible to know the position
of average sea level in the investigated areas. When
sampling, it was sometimes clear that the water level
was low because there was an exposed tidal flat.
However, for a number of stations from water depths
of less than around 0.5 m at the time of sampling, it
has not been possible to be certain whether they are
normally subtidal or intertidal. The depths recorded
are those at the time of sampling.
In areas exposed at the time of collection, samples
were scraped from the sediment surface (to a depth
of 1–2 cm). The subtidal samples were collected
with a small grab. Most samples were about 100 cm3 .
The grab was carefully operated so that negligible
loss of surface sediment took place. On removal
from the water, it was gently opened in a bowl. The
surface oxidised sediment layer was scooped off and
placed in a container. The samples were preserved in
70% ethanol.
At Tjøme (in the area of samples 86 and 87), a
9 cm core (sta. 172, diameter 6.8 cm, taken in 0.8
m of water in September 1996) was sectioned into 1
cm slices in the field and all slices were preserved in
70% ethanol.
In the laboratory, the foraminiferal samples were
processed by washing on a 63 µm sieve, stained in
rose Bengal for 1 hour, washed again on a 63 µm
sieve to remove excess stain, and dried at 50ºC. The
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
185
Fig. 1. Map showing the 27 sampling areas.
foraminifera in those samples not rich in organic
detritus were concentrated using tetrachloroethylene.
Living and dead assemblages (each of around 250
individuals) were picked from a representative >63
µm portion of each sample. Another >63 µm portion of those samples with a dominance of calcareous
tests was digested in acetic acid pH 2.5 for at least
2 hours (to remove the calcareous material), washed
on a 63 µm sieve (to remove the acid) and dried.
The purpose of acid treatment was to concentrate
the agglutinated tests in samples predominated by
calcareous forms.
Whenever possible, 250 individuals were picked
from each acid-treated sample to give the ATA. For
the 98 ODA samples with a high proportion of agglutinated tests (>50%), it was unnecessary to treat
them with acid; additional agglutinated specimens
were simply picked from the ODA to increase the
agglutinated count to 250 individuals. The term ATA
refers to all residual agglutinated assemblages re-
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J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
sulting from dissolution, regardless of whether the
process was natural or carried out by us. In some
instances (notably marshes), both the living and dead
assemblages were entirely agglutinated.
Most Reophax moniliformis tests were fragmented so it was impossible to decide exactly how
many individuals were present. In order to reduce
over-representation, we decided to count only those
specimens with 3 or more chambers. Another problem is that individuals of Goesella waddensis lacking their initial chambers are impossible to distinguish from R. moniliformis. Consequently, we have
included all G. waddensis in R. moniliformis and
the few instances where we are confident that it
is present are given in Murray and Alve (1999).
Ammotium cassis and Ammobaculites balkwilli were
also commonly fragmented. We therefore counted
only those fragments including the initial part.
Thirty-six samples yielded fewer than 50 individuals (Appendix A) and therefore have been omitted
from the statistical discussion. Species diversity was
calculated using the Fisher alpha index (Fisher et
al., 1943) and the information function (see Murray,
1991) but only on those samples with at least 100
individuals. The similarity values given in Table 2
were calculated using the Sanders method (see Mur-
Fig. 2. SEM photographs of agglutinated taxa. The scale bar is
100 µm for all except Ammotium cassis (V) where it is 1 mm. #
refers to sample no.; the localities are given in Appendix A.
A–C. Balticammina pseudomacrescens Brönnimann, Lutze and
Whittaker, #159.
A.
Showing typical collapsed chamber walls on the spiral
side.
C.
Showing the deep umbilicus with primary apertural rim
and secondary pore apertures.
D, E. Haplophragmoides wilberti Andersen, #126.
F–H. Jadammina macrescens (Brady), #153.
I.
Ammobaculites balkwilli Haynes, #98.
J.
Ammotium salsum (Cushman and Brönnimann), #168.
K.
Eggerelloides scaber (Williamson), #18.
L.
Miliammina fusca (Brady), #122.
M. Ammoscalaria runiana (Heron-Allen and Earland), #100.
N–P. Tiphotrocha comprimata (Cushman and Brönnimann),
#166.
Q–S. Trochammina inflata (Montagu), #126.
T.
Goesella waddensis Van Voorthuysen, #122.
U.
Reophax moniliformis Siddall, #122, note the similarity to
the uniserial portion of T.
V.
Ammotium cassis (Parker), #100.
187
ray, 1991, p. 322). Most of the agglutinated taxa are
illustrated in Fig. 2. Paratrochammina (Lepidoparatrochammina) haynesi is illustrated in Brönnimann
and Whittaker (1986).
3. Results
Appendix A lists all ATA data from the 27 areas
(Fig. 1). Rare occurrences are listed at the bottom of
the table as ‘other agglutinated’. Table 1 gives a list
of species arranged according to water depth distribution.
The only species totally restricted to intertidal areas is Balticammina pseudomacrescens. Other species
which are dominant from intertidal to 0.5 m include
Trochammina inflata and Haplophragmoides wilberti,
whereas Jadammina macrescens can dominate down
to 1 m. Miliammina fusca and R. moniliformis dominate over the greatest depth range, from intertidal to 6
m. All other species, except Ammotium cassis (upper
depth limit 2 m), have a total range extending from the
intertidal zone to subtidal areas and commonly to 6 m.
In intertidal–subtidal areas, 73 ATAs (½50 individuals) are dominated by M. fusca and, of these,
50, ranging in depth from the shoreline to 6 m,
have ½60% of this form (see Appendix A). The
species diversity values are very low: Fisher alpha
0.5–2.7 and H .S/ 0.2–1.9 for all environments (Appendix A). The diversity data presented in this paper
have been plotted together with previously published
data giving a total of 191 samples (Figs. 3 and 4).
The results from the Tjøme core are given in Table 2. The surface (0–1 cm) living assemblage has
57% calcareous forms (Ammonia beccarii, Elphidium oceanensis, Elphidium williamsoni) whereas the
ODA has just 5% (A. beccarii). No living forms were
encountered below the 0–2 cm layer (examination of
the 1–2 cm interval revealed 1 live per 250 dead).
Dead calcareous forms are further reduced down to
4–5 cm and they are absent below this. Therefore,
the ODA is a natural ATA from 5 cm downwards.
4. Discussion
The dead assemblages represent the time-averaged contributions of empty tests from the produc-
188
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
Table 1
Water depth ranges of ATA species
Species
Overall range
(m)
Makes up ½5% of
assemblage
Makes up ½10% of
assemblage
As dominant species
B. pseudomacrescens
T. comprimata
H. wilberti
A. salsum
J. macrescens
T. inflata
M. fusca
R. moniliformis
A. runiana
A. balkwilli
E. scaber
P. (L.) haynesi
A. cassis
IT
IT–2
IT–6
IT–5.5
IT–6
IT–5.5
IT–6
IT–6
IT–6
IT–6
IT–6
IT–6
2–6
IT
IT–0.1
IT–4.5
<0.5–5.5
IT–2
IT–4.5
IT–6
IT–6
IT–6
IT–6
IT–6
IT–5
3–6
IT
IT–0.1
IT–4.5
<0.5
IT–1
IT–4.5
IT–6
IT–6
IT–6
IT–5.5
IT–6
2
3–6
IT
not dom.
IT–<0.5
not dom.
IT–1
IT–<0.5
IT–6
IT–6
1–5.5
2–4
3–6
not dom.
3–4.5
IT D intertidal.
tion of successive living assemblages and subsequent
modification due to postmortem processes. In general, the most significant postmortem processes are
transport (which can lead to loss or gain of tests)
and destruction (especially dissolution of the calcareous tests but also disintegration of fragile agglutinated taxa). In the Skagerrak=Kattegat area, there is
some local transport due to wind and wave activity
and this extends the depth ranges of typical marsh
species into shallow subtidal waters (see Table 1 and
Murray and Alve, 1999). There may be some loss
of agglutinated tests (M. fusca is sometimes rather
fragile). Dissolution is widespread (Alexandersson,
1979) and a significant modifying process (Murray
and Alve, 1999). Those living assemblages which
are dominantly calcareous in many cases give rise
to ODAs rich in agglutinated tests as a result of
dissolution under natural conditions. This is a major difference from other areas that we have studied
(Alve and Murray, 1994, 1995; Murray and Alve,
1994) and makes the Skagerrak=Kattegat highly distinctive.
Fig. 3. Fisher alpha values for ODA and ATA data from this
study and published data (Alve and Murray, 1995). The shaded
marginal marine area includes the new data together with those
published on the Hamble estuary.
Fig. 4. Information function values [H .S/] for ODA and ATA
data from this study and published data (Alve and Murray, 1995).
Shaded area as in Fig. 3.
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
189
Table 2
Data on core 172
Stained
0–1
Core depth (cm):
% values
Ammobaculites balkwilli
Ammoscalaria runiana
Eggerelloides scaber
Jadammina macrescens
Miliammina fusca
Reophax moniliformis
Ammonia beccarii
Ephidium oceanensis
Ephidium williamsoni
Number counted
No. spp
% agglutinated
% calcareous
% similarity with dead
% similarity with total
Dead
0–1
Total
0–1
Dead
2–3
Dead
4–5
Dead
6–7
Dead
8–9
0
3
0
0
36
3
44
0
13
0
32
0
0
50
11
5
0
0
0
28
0
0
48
10
10
0
3
0
32
0
1
52
12
3
0
0
1
30
0
0
62
6
0
0
0
0
25
0
0
72
3
0
0
0
0
23
0
0
74
2
0
0
0
265
7
43
57
48
56
258
6
95
5
–
92
305
7
87
13
92
–
255
6
97
3
98
89
253
6
99
0
89
84
259
5
100
0
80
77
259
5
100
0
76
73
4.1. The origin of the ATAs
In our previous studies on the development of
wholly agglutinated assemblages, we have simulated
natural dissolution on predominantly calcareous assemblages through laboratory experiments. However, an important aspect of this study is that samples
from the Skagerrak=Kattegat already show significant natural dissolution; many living calcareous assemblages become agglutinated ODAs which are
natural ATAs. The results from core 172 demonstrate
very clearly that natural dissolution is proceeding
beneath the sediment surface so that at a depth of 1
cm a predominantly calcareous surface living assem-
Table 3
Different pathways from live through ODA to ATA assemblages for samples where the living assemblages are dominated by agglutinated
taxa (½50 individuals)
Sample No.
No. of samples
Live
ODA
ATA
37, 42, 80, 84,
86, 98, 134, 135,
143, 144, 146,
148, 158, 160,
161, 165, 168
125
124, 127, 155
61, 57
31
111
152, 153, 154,
156, 166, 171
128
169
123
126
159
17
M. fusca
M. fusca
M. fusca
M. fusca
M. fusca
M. fusca
M. fusca
A. runiana
R. moniliformis
R. moniliformis
J. macrescens
J. macrescens
J. macrescens
J. macrescens
A. balkwilli
H. wilberti
B. pseudomacr.
M. fusca
M. fusca
M. fusca
M. fusca
M. fusca
T. inflata
J. macrescens
M. fusca
R. moniliformis
A. runiana
J. macrescens
J. macrescens
H. wilberti
M. fusca
M. fusca
H. wilberti
B. pseudomacr.
M. fusca
M. fusca
M. fusca
M. fusca
M. fusca
T. inflata
J. macrescens
M. fusca
R. moniliformis
A. runiana
J. macrescens
J. macrescens
H. wilberti
M. fusca
M. fusca
H. wilberti
B. pseudomacr.
1
3
2
1
1
6
1
1
1
1
1
190
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
Table 4
Different pathways from live through ODA to ATA assemblages for samples where the living assemblages are dominated by calcareous
taxa (½50 individuals)
Sample No.
No. of samples
Live
ODA
ATA
16
53
64
32, 65, 68, 70,
71, 74, 81, 82,
88, 89, 87
15, 99, 100, 114
1
17, 28
101
115
2, 67
3, 26, 43, 120
48
130, 129
140, 119, 139
47, 49
66
113
1C, 20, 23, 46,
55, 58, 59, 60,
72, 75, 77, 79,
85, 91, 141,
142, 164, 167
44
51
52, 54, 109
73, 76, 90
92
121, 122
45
63
11
13
1
1
1
11
A. beccarii
A. beccarii
A. beccarii
A. beccarii
A. beccarii
A. beccarii
A. beccarii
A. beccarii
A. beccarii
A. beccarii
A. beccarii
E. excavatum
E. excavatum
E. excavatum
E. excavatum
E. excavatum
E. excavatum
E. excavatum
E. excavatum
E. williamsoni
E. williamsoni
E. williamsoni
E. williamsoni
E. williamsoni
E. williamsoni
E. williamsoni
E. williamsoni
E. williamsoni
E. williamsoni
E. oceanensis
H. germanica
H. germanica
C. lobatulus
N. depressulus
A. beccarii
E. williamsoni
R. moniliformis
M. fusca
M. fusca
M. fusca
A. runiana
A. beccarii
E. scaber
E. williamsoni
A. beccarii
A. beccarii
M. fusca
A. beccarii
A. runiana
E. excavatum
A. cassis
H. germanica
E. excavatum
M. fusca
M. fusca
M. fusca
M. fusca
M. fusca
J. macrescens
E. williamsoni
E. williamsoni
E. williamsoni
A. beccarii
M. fusca
J. macrescens
M. fusca
C. lobatulus
C. lobatulus
A.balk.=A. run.
M. fusca
R. moniliformis
M. fusca
M. fusca
M. fusca
A. runiana
R. moniliformis
E. scaber
A. runiana
A. runiana
M. fusca
M. fusca
A. runiana
A. runiana
M. fusca
A. cassis
M. fusca
A. runiana
M. fusca
M. fusca
M. fusca
M. fusca
M. fusca
J. macrescens
R. moniliformis
M. fusca
M. fusca
M. fusca
M. fusca
J. macrescens
M. fusca
M. fusca
M. fusca
4
1
2
1
1
2
4
1
2
3
2
1
1
18
1
1
3
3
1
2
1
1
1
1
blage has become an almost completely agglutinated
assemblage.
Dissolution of calcareous tests has previously
been recorded from shallow subtidal sediments in
Oslo Fjord where sites with >20% calcareous tests
during the spring had <3% during the other seasons
and organic linings were frequent (Alve and Nagy,
1986). Severe dissolution was also reported in subtidal sediments in Long Island Sound, USA (Green
et al., 1993). Here, the calcareous foraminifera disappeared during the course of a year due to frequent occurrence of irrigated burrows and tubes of
macrofauna which prevent the buildup of carbonate
alkalinity (see also Aller, 1982). Dissolution has also
been reported from intertidal sediments in the northern Gulf of California, Mexico (Martin et al., 1995)
and marsh sediments are well known for having poor
preservation of calcareous tests due to postmortem
dissolution (see, for example, Parker and Athearn,
1959; Murray, 1971; Scott and Medioli, 1980a,b;
Williams, 1989; Jonasson and Patterson, 1992).
In the present investigation, the simplest relationship between live assemblages, ODAs and ATAs is
found in areas where the living assemblages are
composed mainly of agglutinated forms and they in
turn give rise to agglutinated ODAs. The dominant
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
Table 5
Different pathways from ODA to ATA for those samples with
living assemblages of <50 individuals
Sample No.
No. of
samples
ODA
ATA
5
78
4, 7, 8, 14,
21, 22, 27,
35, 36, 56,
83, 141, 145,
147, 157
19, 24, 25,
29, 30
6
18
1
1
15
E. williamsoni
R. moniliformis
M. fusca
M. fusca
M. fusca
M. fusca
M. fusca
E. scaber
E. scaber
E. excavatum
A. beccarii
M. fusca
R. moniliformis
M. fusca
M. fusca
M. fusca
M. fusca
M. fusca
E. scaber
E. scaber
M. fusca
E. scabrus
5
1
1
taxon may be the same in both the living assemblage and ODA but in some instances the dominant
taxa are different (Table 3). For example, a living R.
moniliformis assemblage may go to an A. runiana
ODA, or a living J. macrescens assemblage to a H.
wilberti ODA. Also, some living agglutinated assemblages (A. runiana, J. macrescens, A. balkwilli) go
to M. fusca ODAs. This is most probably due to the
presence of a bloom in the dominant living species at
the time of sampling. Although no dissolution may
be apparent, the reality is that these Scandinavian
intertidal environments are not favourable for the
existence of calcareous foraminifera.
The calcareous living assemblages may follow
two alternative routes. In one pathway, both the living assemblage and the ODA are dominated by a
calcareous species (either the same or a different
one) and the ATA by an agglutinated form (Table 4).
For example, an A. beccarii living assemblage may
give rise to an A. beccarii or an E. williamsoni ODA.
These then give rise to different ATAs (see Tables 4
and 5). In the other case, the living calcareous assemblage goes directly to an agglutinated ODA and this
is a naturally formed ATA. The commonest examples are those of living A. beccarii or E. williamsoni
giving rise to an M. fusca ODA=ATA.
It is significant that the most abundant ATA (M.
fusca) can be derived from numerous different parent
living assemblages and ODAs. Indeed, ten assemblages, 7 of them calcareous, follow this pathway (A.
balkwilli, A. runiana, M. fusca, A. beccarii, Cibicides
191
lobatulus, Elphidium excavatum, E. oceanensis, E.
williamsoni, Haynesina germanica, Nonion depressulus). This leaves M. fusca as the most ubiquitous
agglutinated shallow water species.
4.2. Preserved ecological information
Because the marshes are developed on such a
small scale and the area is microtidal, the intertidal
subenvironments are severely compressed and there
is no obvious differentiation into low and high marsh
as seen in more tidally influenced areas. The marsh
ODAs are composed entirely of agglutinated taxa so
they are naturally occurring ATAs.
J. macrescens dominates the marsh assemblages
at Bunnefjorden, Hålkedalskilen and Kalundborg. It
commonly occurs together with M. fusca, T. inflata
and sometimes with B. pseudomacrescens (Hunnebotn), H. wilberti (Kalundborg), or Tiphotrocha
comprimata (Bunnefjord, Hunnebotn, Horten). Of
these, T. comprimata and B. pseudomacrescens have
the most restricted depth range (Table 1).
As discussed above, by far the most abundant
ATA is that of M. fusca which can be derived from
numerous different living assemblages. This implies
that a fossil M. fusca assemblage could represent
a wide range of marginal marine subenvironments,
from marsh edge to 6 m water depth, salinity 15–
30‰ (at the time of sampling but undoubtedly lower
during other seasons), on sediments varying from
carbonate shell sand to clastic sands and organic-rich
muds, with low to high TOC (0.1–12.3%), in a
temperate area (seasonal range from 0 to 28ºC).
In the shallow depth range studied, there are few
obvious depth boundaries in either the ODAs or the
ATAs. Ammotium cassis (shallowest depth limit 2–3
m) is the only agglutinated species encountered which
does not extend into the intertidal zone. Whereas typical live marsh assemblages (J. macrescens, T. inflata,
T. comprimata, H. wilberti, B. pseudomacrescens)
can be readily distinguished from the assemblages
living on bare intertidal flats, and from those ODAs
not too seriously affected by dissolution, it is not possible to distinguish between marsh and shallow subtidal environments using ATAs. This is because J.
macrescens, T. inflata and H. wilberti extend down
to 0.5–1 m water depth. However, they still indicate
close proximity to a marsh.
192
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
4.3. Comparison with previously analysed ATAs
In the first experiments on dissolution, Alve and
Murray (1994) used 8 shallow subtidal and intertidal
samples from the Hamble estuary in southern England. This estuary has a tidal range of 2.0–4.9 m
(neaps to springs) and the salinity in the area studied ranged from 30 to 33‰. Seven ODAs were dominated by calcareous taxa. Four A. beccarii ODAs gave
rise to A. balkwilli ATAs; two H. germanica ODAs
gave rise to E. scabrus ATAs while another gave rise
to a Textularia tenuissima=Paratrochammina (Lepidoparatrochammina) haynesi ATA. A single marsh
sample had an ODA in which J. macrescens was the
dominant species although the assemblage also comprised more than 50% calcareous forms; the ATA was
likewise dominated by J. macrescens.
The results from the Hamble estuary are comparable with those from this study except that here A.
beccarii ODAs gave rise to A. runiana, R. moniliformis and M. fusca as well as A. balkwilli ATAs.
This may be partly a reflection of the larger number of samples studied. However, the H. germanica
ODAs of this study gave rise to M. fusca ATAs
probably because of the more brackish waters (Alve,
1995; table 1).
In terms of species diversity, using either the
Fisher alpha index or the information function, both
studies show that the low diversity ODAs give rise to
low diversity ATAs. This linkage of diversity patterns
between ODAs and ATAs has also been established
in the other experimental studies involving the deep
sea (Murray and Alve, 1994) and shelf seas (Alve
and Murray, 1995). From marginal marine to deep
sea environments there is a progression from low to
high diversity [for Fisher alpha and H .S/] both for
the ODAs and ATAs (Figs. 3 and 4). This means that
even though some ODAs have been seriously altered
by severe dissolution, the residual ATA still retains
the ecological information necessary to recognise the
environment of deposition.
Overall, it appears that in all the environments
studied, from deep sea to intertidal, the ATAs preserve a remarkable amount of ecological information. Therefore, by comparison with our accumulated
database on modern forms it should now be possible to make palaeoecological interpretations of fossil
agglutinated assemblages with more confidence.
4.4. Comment on the use of living, dead and total
assemblages
Some authors add together the living and dead
assemblages to give so-called total assemblages in
the belief that these are more representative of what
becomes preserved in the fossil record (see discussion in Murray, 1982). This study shows unequivocally that the concept of the total assemblage as
a better analogue of a fossil assemblage than the
time-averaged dead assemblage (which has experienced postmortem modification) is incorrect. Only
by investigating the consequences of the taphonomic
processes that affect all tests after death is it possible
to properly determine the pathways to fossilisation.
As we have shown here, in the Skagerrak=Kattegat
area, many calcareous living assemblages give rise to
exclusively agglutinated ODAs that become part of
the fossil record (e.g., below 5 cm in core 172). This
would not have been shown by using total assemblages because they would have been a mixture of
calcareous and agglutinated forms (compare stained,
dead and total data for 0–1 cm; Table 2). Furthermore, the magnitude of the loss of tests through dissolution would have been seriously underestimated.
5. Summary and conclusions
The shallow water environments of the Skagerrak=
Kattegat are unusual in that they are characterised by
naturally occurring carbonate dissolution on a large
scale. This leads to major taphonomic change in the
transition from live to dead assemblage.
In many cases calcareous dominated living assemblages become preserved as agglutinated original dead assemblages (ODAs). In those instances
where natural dissolution has not taken place, we
have treated the samples with acid to simulate the
natural process and this gives rise to acid treated assemblages (ATAs) of agglutinated foraminifera. The
term ATA refers to any residual agglutinated assemblage resulting from dissolution, whether natural or
carried out by us.
The pathways from living assemblage to ODA
to ATA are varied and complex. In the simplest
case, the living assemblages are agglutinated and
they give rise to similar ODAs and ATAs. This is
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
the case for most marshes where the environment
is commonly hostile to the existence of calcareous
taxa. In other types of intertidal and subtidal areas,
calcareous-dominated living assemblages may give
rise to calcareous or agglutinated ODAs the latter
being natural ATAs.
The marsh ATAs are found to a water depth of
0.5–1 m (due to local transport into subtidal waters)
but are still fairly reliable indicators of sea level.
The most widespread ATA is dominated by Miliammina fusca but this can be derived from 10 different living assemblages (including 7 calcareous
ones). It therefore represents a broad range of environments, from intertidal to 6 m, salinity at least 15–
30‰, temperature 0–28ºC, TOC (0.1–12.3%) and
sediments ranging from carbonate sands, through
clastic sands to organic-rich muds.
The living foraminiferal assemblages have low
species diversity and this is faithfully preserved in
both the ODAs and ATAs.
Although there is severe information loss through
taphonomic change (notably dissolution of calcareous tests), the ODAs and ATAs still give a fairly
reliable record of the ecology of the environments.
This has very important implications for the palaeoecological interpretation of fossil agglutinated assemblages.
193
This study demonstrates unequivocally that it is
always essential to compare data on living and dead
assemblages in order to determine the pathways to
fossilisation. The use of total (living plus dead)
assemblages in this area would give unrepresentative
and misleading results.
Acknowledgements
We are grateful to the Natural Environment Research Council for grant GR9=1591‘A’ and the Industrial Liaison fund (Oslo University) for financial
support. The following people helped in various
ways in the field: Pål Glad for help with sampling
in Norway; Eva Larsen, Siggurd Gausdal, Arne M.
Svendsen, Bent Åsnes, Trond Smith, John Ingebrigtsen, and Tjärnö marinbiologiska laboratorium for
provision of boats. Karen Luise Knudsen and MaritSolveig Seidenkrantz are thanked for taxonomic discussions. Barbara Cressey and Barry Marsh assisted
with the preparation of material for SEM and for
photography. We thank Jean-Pierre Guilbault and
an anonymous referee for helpful comments on the
manuscript.
0
24
0
0
0
2
0
9
54
0
10
0
0
1
264
8
1.7
1.26
% values (ATA)
Ammobaculites balkwilli
Ammoscalaria runiana
Ammotium cassis
Ammotium salsum
Balticammina pseudomacr.
Eggerelloides scaber
Haplophragmoides wilberti
Jadammina macrescens
Miliammina fusca
Paratroch. (L.) haynesi
Reophax moniliformis
Tiphotrocha comprimata
Trochammina inflata
Other agglutinated
Total counted
No. spp.
Alpha-index ATA
H .S/ ATA
0
8
0
0
0
6
0
16
69
0
0
0
0
0
258
6
1.2
0.96
0
42
0
0
0
9
0
0
45
0
4
0
0
0
279
5
1.0
1.08
0
18
0
0
0
5
0
8
68
0
1
0
0
0
257
5
1.0
0.97
S
23
2.5
% value (ATA)
Ammobaculites balkwilli
Ammoscalaria runiana
Ammotium cassis
Ammotium salsum
Balticammina pseudomacr.
Eggerelloides scaber
Haplophragmoides wilberti
Jadammina macrescens
Miliammina fusca
Paratroch. (L.) haynesi
Reophax moniliformis
Tiphotrocha comprimata
Trochammina inflata
Other agglutinated
Total counted
No. spp.
Alpha-index ATA
H .S/ ATA
S
21
1
Kilsfjorden (6)
I
20
0.1
Environment:
Sample no.:
Water depth (m):
0
26
0
0
0
10
0
0
64
0
0
0
0
0
231
5
1.0
0.92
Area (area no.):
3
21
0
0
0
0
0
1
26
0
49
0
0
1
273
6
1.2
1.89
S
32
2
Environment:
Sample no.:
Water depth (m):
I
31
0.2
Isefjærfjord (1)
I
27
0.1
Area (area no.):
2
25
0
0
0
14
0
0
57
0
0
0
0
0
258
8
1.7
1.12
S
26
3.5
5
21
0
0
0
67
0
0
8
0
0
0
0
0
262
4
0.8
0.93
S
28
4
0
44
0
0
0
2
0
2
51
0
0
0
0
1
109
5
1.1
0.89
S
22
4
0
10
0
0
0
88
0
0
1
0
0
0
0
0
267
4
0.8
0.42
S
29
5
1
17
0
0
0
52
0
0
28
0
0
0
0
2
255
9
2.0
1.14
S
24
5.5
1
8
0
0
0
91
0
0
0
0
0
0
0
0
300
4
0.7
0.35
S
30
6
3
14
0
0
0
53
0
0
29
1
0
0
0
0
273
5
1.0
1.13
S
25
6
0
27
0
0
0
13
0
6
46
0
4
0
3
0
252
7
1.4
1.40
I
14
0
25
25
0
0
0
23
0
0
18
0
7
0
0
2
141
7
1.3
1.62
S
16
2
24
23
0
0
0
42
0
0
8
0
4
0
0
0
244
5
1.0
1.37
S
17
3
6
11
0
0
0
82
0
0
0
0
1
0
0
0
266
5
1.0
0.64
S
18
4
12
13
0
0
0
74
0
0
1
0
0
0
0
0
347
5
0.8
0.81
S
19
5
0
17
0
2
0
0
0
2
66
0
13
0
0
0
176
5
1.0
1.00
I
84
0.1
3
18
0
1
0
0
0
1
46
0
30
0
0
0
269
7
1.4
1.25
S
85
0.4
9
16
0
0
0
0
0
1
51
0
22
0
0
0
221
7
1.5
1.27
S
86
0.6
S
88
1.5
2
0
19
24
0
0
0
0
0
0
0
1
0
0
0
0
68
68
0
2
11
1
0
0
0
0
0
3
296 268
4
11
0.8
2.3
0.89 0.96
S
87
0.8
Tjøme, NE Gåsholmen (7)
16
43
0
0
0
8
0
0
32
0
0
0
0
0
230
5
1.0
1.25
S
15
1
Kvastadkilen (2)
Relative abundance data for the acid treated assemblages (ATAs)
Appendix A
2
14
0
0
0
2
0
0
78
2
1
0
0
0
284
7
1.4
0.78
S
89
3
0
0
0
0
0
0
0
0
93
7
0
0
0
0
15
2
0
1
0
0
0
5
0
0
91
3
0
0
0
1
282
5
0.9
0.41
S
11
2
3
0
0
1
0
0
2
1
88
0
5
0
0
0
303
7
1.2
0.54
M
167
0
0
0
0
1
0
0
0
2
62
1
33
0
0
0
230
6
1.2
0.87
I
53
0.1
Borre (8)
0
0
0
0
0
0
0
0
77
9
0
0
0
14
22
4
S
10 a
1
5
1
0
3
0
1
2
1
53
0
33
0
0
1
288
9
1.9
1.22
I
52
0.1
0
0
0
0
0
4
0
0
74
9
0
0
0
13
23
5
S
12 a
4
Lyngør, Dype Holla (3)
I
9a
0
8
0
0
9
0
1
0
0
32
0
48
0
0
2
109
6
1.3
1.26
I
51
0.1
0
2
0
0
0
13
0
0
74
9
0
0
0
1
163
6
1.0
0.86
S
13
5
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6
19
0
0
0
1
0
23
31
0
1
0
17
3
199
8
1.8
1.67
S
2
1
1
18
0
0
0
2
0
1
74
4
2
0
0
1
200
8
1.8
0.87
S
3
2
4
7
0
0
0
3
3
0
82
1
2
0
0
0
119
7
1.8
0.78
S
4
2.5
0
0
0
1
0
0
0
42
46
0
0
11
0
0
276
5
0.9
0.75
1
1
0
1
0
1
0
29
60
0
3
3
0
0
408
10
1.8
1.10
M
165
0
0
8
0
3
0
0
0
14
56
0
13
7
0
0
264
8
1.8
1.36
M
164
0
4
4
0
1
0
0
0
6
48
0
27
10
0
0
277
8
1.6
1.43
I
42
0.1
Horten (9), mainland side
0
32
0
0
0
0
0
4
52
8
2
0
0
1
230
10
2.1
1.23
I
M
50 a 166
0
0
0
2
0
0
0
0
0
0
16
2
79
0
0
2
56
5
S
1C
0.5
Lyngør, Tøkersfjord (4)
I
1
0
3
10
0
2
0
1
0
0
60
0
23
0
0
1
101
7
1.8
1.14
S
46
2
1
31
0
0
0
2
1
1
61
0
4
0
0
0
338
7
1.4
0.97
S
5
3
S
8
5
22
20
30
0
0
4
0
1
3
0
20
0
0
0
286
8
1.7
1.62
S
49
3
27
6
42
0
0
19
0
0
3
0
2
0
0
2
292
9
1.9
1.46
S
47
4.5
0
4
9
14
0
0
0
0
0
0
1
4
0
0
2
2
84
71
2
2
1
1
0
0
0
0
0
1
341 275
7
10
1.4
2.0
0.65 1.09
S
7
4
0
2
0
0
0
0
1
1
95
0
0
0
0
0
175
4
0.9
0.23
2
3
0
0
0
0
2
2
90
0
0
0
0
0
255
7
1.4
0.48
S
81
0.4
Hasdalen (5)
I
80
0.1
2 6
8 6
0 0
0 0
0 0
0 1
0 0
2 6
87 81
0 0
0 0
0 0
0 0
1 0
91 84
5 5
S S
82 83
1 1
0
5
0
0
0
0
4
64
22
0
1
3
0
0
139
6
1.3
1.07
I
44
0.1
S
43
2
0
7
11
5
0
4
0
1
0
0
0
3
0
0
68
9
12
67
0
0
2
2
2
1
6
0
0
1
114 272
6
11
1.3
2.2
1.07 1.27
S
45
1
0
43
26
0
0
7
0
1
1
0
20
0
0
0
69
6
S
48
3
Horten (9), island side
4
5
0
0
0
5
0
4
78
2
2
0
0
0
169
7
1.6
0.92
S
6
6
194
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
0
2
0
4
0
0
0
1
92
0
0
0
0
0
266
5
1.0
0.35
% value (ATA)
Ammobaculites balkwilli
Ammoscalaria runiana
Ammotium cassis
Ammotium salsum
Balticammina pseudomacr.
Eggerelloides scaber
Haplophragmoides wilberti
Jadammina macrescens
Miliammina fusca
Paratroch. (L.) haynesi
Reophax moniliformis
Tiphotrocha comprimata
Trochammina inflata
Other agglutinated
Total counted
No. spp.
Alpha-index ATA
H .S/ ATA
0
14
0
0
0
0
0
4
39
0
43
0
0
0
148
4
0.9
1.14
4
19
0
2
0
0
0
3
51
0
21
0
0
0
275
6
1.2
1.29
0
8
0
0
0
0
0
0
68
0
25
0
0
0
118
3
0.7
0.80
2
13
0
2
0
1
0
4
65
0
13
0
0
0
281
8
1.7
1.14
S
75
0.9
2
13
0
0
0
0
3
12
57
0
11
0
2
0
283
7
1.4
1.37
I
76
0.4
% value (ATA)
Ammobaculites balkwilli
Ammoscalaria runiana
Ammotium cassis
Ammotium salsum
Balticammina pseudomacr.
Eggerelloides scaber
Haplophragmoides wilberti
Jadammina macrescens
Miliammina fusca
Paratroch. (L.) haynesi
Reophax moniliformis
Tiphotrocha comprimata
Trochammina inflata
Other agglutinated
Total counted
No. spp.
Alpha-index ATA
H .S/ ATA
I
77
0.2
Gullm.fj., Finnsbobukten (15)
I
79
0.1
0
0 0
1
0 0
0
0 0
2
0 0
0
0 0
0
0 0
0
0 0
0
0 0
95 100 0
0
0 0
2
0 0
0
0 0
0
0 0
0
0 0
248 28 0
6
1 0
1.2
0.26
Environment:
Sample no.:
Water depth (m):
I
78
0.2
Bunnefjord (11)
3
10
0
0
0
1
0
1
74
0
12
0
0
0
261
6
1.2
0.89
S
74
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6
0
0
0
0
0
0
0
88
0
0
0
0
6
16
3
0
0
0
0
0
0
0
89
5
0
0
6
0
0
342
4
0.7
0.44
0
0
0
0
0
0
0
64
0
0
0
31
3
1
327
4
0.7
0.81
0
0
0
0
0
0
0
97
1
0
0
0
2
0
262
4
0.8
0.15
0
0
0
0
0
0
0
94
5
0
0
0
1
0
272
3
0.6
0.25
0
3
0
0
0
6
0
0
87
0
4
0
0
0
121
4
0.9
0.53
I
73
0
7
14
0
0
0
5
0
0
71
0
3
0
0
0
278
7
1.4
0.97
I
72
0
0
15
0
0
0
0
0
2
62
0
20
0
0
0
253
5
1.0
1.01
I
71
0.1
0
10
0
1
0
0
0
0
49
0
38
0
0
0
298
6
1.2
1.05
S
70
0.5
0
16
0
0
0
0
0
16
26
0
35
0
0
6
31
5
2
13
0
2
0
3
0
2
62
0
15
0
0
1
284
10
2.0
1.26
S
S
69 a 68
1
2
Gullmarsfjord, Gullmarsvik (16)
0
0
0
0
0
0
0
25
75
0
0
0
0
0
4
2
S
S
S
S
S
M
M
M
M=I
39 a 38 a 34 a 40 a 41 a 171 152 153 154
1.5 2
2.5 3
4.1 0
0
0.1 0.1
Area (area no.):
0
5
0
2
0
0
0
0
93
0
0
0
0
0
260
4
0.8
0.32
S
35
1
Environment:
Sample no.:
Water depth (m):
I
36
0.4
Sandebukta (10)
I
37
0.2
Area (area no.):
Appendix A (continued)
Hunnebotn (12)
3
14
0
0
0
0
0
8
46
0
30
0
0
0
79
5
S
67
3
0
2
0
0
0
0
0
52
45
0
1
0
1
0
266
6
1.0
0.87
10
11
0
4
0
3
0
1
51
0
18
0
0
0
115
7
1.8
1.43
S
66
4
0
0
0
0
55
0
0
28
4
0
0
13
0
0
256
5
0.9
0.71
0
0
0
3
19
0
0
31
41
0
0
6
0
0
284
6
1.0
1.33
9
0
0
0
0
0
0
2
88
0
0
0
0
0
274
4
0.8
0.44
6
8
0
2
0
0
0
0
44
0
39
0
0
1
269
7
1.4
1.21
I
65
0
7
4
0
1
0
0
0
2
19
0
67
0
0
0
264
6
1.2
0.6
I
64
0
2
1
0
0
0
0
0
10
59
0
27
0
0
0
281
6
1.3
1.05
S
63
0.7
Hafstensfjord (17)
0
0
0
0
57
0
0
5
20
0
0
18
0
0
212
3
0.6
1.10
0
3
0
9
0
0
2
1
0
0
0
1
1
1
87 29
5 48
0
0
2
5
0
0
4
3
0
1
252 108
6
10
1.0
2.7
0.58 1.44
0
33
0
0
0
0
0
0
33
0
17
0
0
17
6
3
4
2
0
1
0
0
0
0
90
0
2
0
0
0
296
7
1.2
0.47
7
4
0
2
0
0
3
7
56
0
21
0
0
0
268
7
1.4
1.33
20
5
0
4
0
0
1
4
47
0
20
0
0
0
333
7
1.4
1.44
0
10
0
2
0
0
0
2
67
0
19
0
0
0
265
5
0.9
0.98
1
21
0
1
0
5
0
0
56
3
12
0
0
2
197
8
1.7
1.29
2
14
0
0
0
3
0
3
71
5
2
0
0
0
265
8
1.7
1.04
jetties (18)
0
40
0
0
0
3
0
3
49
5
0
0
0
0
253
6
1.2
1.10
S
56
1
4
24
0
0
0
4
0
0
56
1
12
0
0
0
252
6
1.1
1.20
S
61
1.8
S
58
3.5
S
60
4
Jonstorp (19)
11
8 33
24
13 13
0
0
0
0
0
0
0
0
0
3
4
3
0
0
0
0
0
0
41
56 41
1
3
1
20
15
8
0
0
0
0
0
0
1
0
0
319 252 275
7
6
6
1.4
1.2 1.2
1.46 1.33 1.36
S
57
2.5
24
11
0
0
0
10
0
0
38
3
15
0
0
0
255
6
1.2
1.57
S
59
5
12
2
0
3
0
0
1
0
66
0
16
0
0
0
303
7
1.4
1.08
0
24
0
2
0
0
0
2
41
0
31
0
0
0
258
5
1.0
1.23
2
1
0
0
0
0
1
2
92
0
1
0
0
0
401
8
1.5
0.43
0
4
0
0
0
0
0
2
64
0
28
0
0
0
269
5
1.0
0.91
7 0
11 0
0 0
2 0
0 0
0 0
1 50
2 0
58 50
0 0
20 0
0 0
0 0
0 0
327 2
7
2
1.4
1.24
19
2
0
3
0
0
3
0
63
0
8
0
0
0
261
8
1.7
1.16
18
0
0
0
0
0
5
0
0
0
0
0
6
0
0
0
67 100
0
0
3
0
0
0
0
0
0
0
324
7
6
1
1.1
1.02
0
0
20
0
0
0
0
0
60
0
0
0
0
0
5
3
S
S
S
S
S
S
S
S
S
S
144 145 148 147 146 138 a 140 139 136 a 137 a
0.3 1
0.5 1.5 1.8 0.2
0.4 0.9 2
4
Kungsbackafjord (18)
0
0
0
50
0
0
0
0
50
0
0
0
0
0
268
3
0.5
0.69
S
I
I
I
62 a 141 142 143
2
0
0
0.2
0
0
0
20
0
0
0
1
78
0
0
1
0
0
250
4
0.8
0.71
S
55
0.5
Tjärnö (14)
I
S
158 54
0
0.2
Hålkedals. (13)
S
M
M
M
I
I
S
M
I
155 170 159 169 160 161 168 156 157
0.3 0
0
0
0
0.02 0.1 0
0
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
195
Environment:
Sample no.:
Water depth (m):
10
28
0
0
0
0
31
0
31
0
0
0
0
0
29
4
0
6
0
2
0
0
19
3
57
0
11
0
0
1
238
8
1.7
1.31
0
35
0
0
0
0
32
0
30
0
0
0
0
3
37
4
0
20
0
0
0
0
0
0
80
0
0
0
0
0
5
2
9
38
0
0
0
0
21
2
26
0
4
0
1
0
172
7
1.6
1.49
0
0
0
0
0
0
0
0
50
0
0
0
0
50
2
2
0
39
0
0
0
0
0
4
31
0
24
0
2
0
51
5
0
52
0
0
0
0
4
1
17
0
24
0
1
0
270
7
1.4
1.23
0
67
0
0
0
0
0
0
17
0
17
0
0
0
6
3
8
50
0
0
0
0
4
0
38
0
0
0
0
0
24
4
13
25
1
0
0
0
1
0
47
0
14
0
0
0
269
7
1.4
1.33
S
S
S
102 a 103 a 98
0.5
2.5
3.5
0
56
0
0
0
0
0
22
11
0
11
0
0
0
9
4
S
129
4
5
35
21
0
0
0
0
0
17
0
22
0
0
0
284
6
1.2
1.49
S
99
5
0
0
0
0
0
0
55
27
5
0
0
0
13
0
260
4
0.8
1.10
M
126
0
9
32
14
0
0
0
0
0
7
0
39
0
0
0
257
6
1.2
1.42
S
100
6
0
0
0
0
0
0
13
41
25
0
0
0
20
0
278
6
1.2
1.34
M
127
0
6
0
0
0
0
0
26
24
12
0
1
0
31
0
270
6
1.2
1.51
I
125
0.3
0
0
0
8
0
0
38
54
0
0
0
0
0
0
13
3
S
97 a
0.5
0
0
0
0
0
0
0
100
0
0
0
0
0
0
1
1
S
96 a
1
Løgstør (25)
0
0
0
0
0
0
73
10
16
0
0
0
1
0
289
4
0.8
0.81
M
128
0
Kalundborg (21)
7
28
0
0
0
7
0
21
10
24
0
0
3
0
29
7
S
95 a
2
0
0
0
3
0
0
19
41
9
0
0
0
27
1
307
7
1.4
1.39
S
124
0.5
0
0
0
0
0
50
13
0
25
0
0
0
13
0
8
4
S
94 a
4.5
35
0
0
3
0
0
1
5
41
0
13
0
1
0
295
9
1.9
1.40
S
123
0.5
3
0
0
4
0
0
3
6
72
0
10
0
0
1
116
8
2.0
1.02
S
119
1.5
4
6
0
0
0
0
53
28
2
0
0
0
6
0
47
6
S
93 a
0.2
3
10
0
0
0
0
1
5
76
0
3
0
0
0
267
8
1.7
0.89
S
92
0.5
Valsted (26)
8
0
0
1
0
0
0
4
58
0
26
0
3
1
285
8
1.8
1.18
S
122
1
The data are presented by geographic area (see Fig. 1 for locations numbered 1–27). M D marsh; I D intertidal; S D subtidal.
a
Samples which yielded fewer than 50 specimens.
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Kalø Vig, Vosnæs pynt (24)
I
I
I
S
104 a 105 a 106 a 101
0
0
0
0.2
Area (area no.):
% value (ATA)
Ammobaculites balkwilli
Ammoscalaria runiana
Ammotium cassis
Ammotium salsum
Balticammina pseudomacr.
Eggerelloides scaber
Haplophragmoides wilberti
Jadammina macrescens
Miliammina fusca
Paratroch. (L.) haynesi
Reophax moniliformis
Tiphotrocha comprimata
Trochammina inflata
Other agglutinated
Total counted
No. spp.
Alpha-index ATA
H .S/ ATA
4
8
0
0
0
0
9
0
71
0
7
0
0
0
246
6
1.2
1.01
% value (ATA)
Ammobaculites balkwilli
Ammoscalaria runiana
Ammotium cassis
Ammotium salsum
Balticammina pseudomacr.
Eggerelloides scaber
Haplophragmoides wilberti
Jadammina macrescens
Miliammina fusca
Paratroch. (L.) haynesi
Reophax moniliformis
Tiphotrocha comprimata
Trochammina inflata
Other agglutinated
Total counted
No. spp.
Alpha-index ATA
H .S/ ATA
S
S
S
S
133 a 132 a 131 a 130
0.6
0.9
1.5
3
Environment:
Sample no.:
Water depth (m):
S
134
0.4
Kildehuse (20)
S
135
0.4
Area (area no.):
Appendix A (continued)
0
10
0
3
0
0
1
3
78
0
4
0
0
0
269
6
1.5
0.82
S
91
1
2
5
0
3
0
0
3
5
65
0
15
0
2
1
282
9
2.0
1.26
S
121
2
0
79
0
0
0
0
0
4
13
0
4
0
0
0
24
4
27
0
0
0
0
69
0
0
0
0
4
0
0
0
26
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
S
S
S
151 a 150 a 149 a
0.4
0.5
1
Fredr. havn (27)
2
0
13
0
0
0
1
0
0
0
0
0
2
0
3
0
75
0
0
0
3 100
0
0
0
0
1
0
125
2
8
1
2.0
0.93
S
90
1
0
16
0
0
0
0
0
6
58
0
13
0
0
6
31
5
5
62
0
0
0
0
9
0
21
0
3
0
0
0
120
5
1.1
1.11
S
S
S
S
118 a 117 a 116 a 115
0.2
1
2.1
3
Brejning, Vejle Fjord (22)
7
0
0 75
0
0
0
0
0
0
0
0
0
0
4
0
68
0
0
0
16 25
0
0
1
0
2
0
268
4
9
2
2.0
1.10
S
120
3
6
58
8
0
0
0
0
0
16
0
12
0
0
1
190
6
1.3
1.25
S
113
4.5
10
59
9
8
0
0
0
0
4
0
7
0
0
1
153
7
1.6
1.35
S
114
5.5
Kalø Vig, Havhuse (23)
23
10
0
14
0
0
0
3
45
0
4
0
0
0
69
6
0
100
0
0
0
0
0
0
0
0
0
0
0
0
5
1
0
20
0
0
0
0
20
0
27
0
13
0
20
0
15
5
0
100
0
0
0
0
0
0
0
0
0
0
0
0
2
1
9
35
0
0
0
0
7
0
39
0
9
0
0
0
54
5
9
43
0
1
0
0
1
0
11
0
35
0
0
0
307
7
1.4
1.29
I
S
S
S
S
S
109 108 a 112 a 107 a 110 111
0.2 0.4
0.5
0.6
2
3
196
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
J.W. Murray, E. Alve / Palaeogeography, Palaeoclimatology, Palaeoecology 149 (1999) 183–197
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