ELSEVIER
Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
Marginal marine environments of the Skagerrak and Kattegat:
a baseline study of living (stained) benthic foraminiferal ecology
Elisabeth Alve a,Ł , John W. Murray b
b
a Department of Geology, University of Oslo, P.O. Box 1047, Blindern, N-0316 Oslo, Norway
School of Ocean and Earth Science, Southampton Oceanography Centre, European Way, Southampton, SO14 3ZH, UK
Received 21 October 1997; accepted 29 April 1998
Abstract
This is the first detailed investigation of the distribution and ecology of living (stained) shallow water (0–6 m)
foraminifera along the Skagerrak–Kattegat coast, eastern North Sea. A total of 25 species (13 agglutinated; 12 calcareous)
are common in the 169 sediment surface samples which were collected from 27 geographic areas. The sediment grain
size and total organic carbon (TOC) content are strongly variable and the salinity and temperature ranges were 10–31‰
and 9–30ºC, respectively, at the time of sampling (July to October) but temperatures down to freezing occur during the
winter. The species are divided into six environmental categories of which the first five comprise euryhaline and the sixth
essentially stenohaline taxa: (1) species associated only with marsh plants, (2) species basically, but not entirely, associated
with marsh plants, (3) species basically, but not entirely, restricted to non-marsh areas, (4) species solely recorded in
non-marsh intertidal to subtidal environments, (5) species restricted to subtidal areas, (6) species basically living in the
most open marine areas. In this region, marshes have a patchy distribution and they are small and compressed due to low
tidal ranges (<40 cm). Balticammina pseudomacrescens (not reported here before) lives in the most elevated, landward,
terrestrial parts of marshes and thus defines the uppermost limit of the influence of marine water. However, the marshes
are generally dominated by Jadammina macrescens and Miliammina fusca at the landward and seaward sides, respectively.
Jadammina macrescens is observed living epiphytically on decaying Carex leaf debris. The most widely distributed
euryhaline species are Elphidium williamsoni, Miliammina fusca, Ammonia beccarii, and Haynesina germanica. The
former two are common only in sediments with a mud content less than about 60%, whereas the latter two are common
even in sediments with >80% mud. Ammoscalaria runiana is common only in coarse-grained sediments (<20% mud)
with low TOC (0.7%). There are no marked biogeographic boundaries within the Skagerrak–Kattegat area but 10 of the
25 commonly occurring species have not been reported from the adjacent Baltic Sea, probably partly due to the brackish
character of the water there. The southern limits of distribution of the northern species, Elphidium albiumbilicatum,
Ammotium cassis, and Ophthalmina kilianensis, are in the Kattegat–Baltic Sea.  1999 Elsevier Science B.V. All rights
reserved.
Keywords: ecology; biogeography; shallow water foraminifera; marsh; Skagerrak–Kattegat
Ł Corresponding
author. Tel.: C47 2285 7333; Fax: C47 2285 4215; E-mail: elisabeth.alve@geologi.uio.no
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 1 3 1 - X
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E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
1. Introduction
2. Area description
There continues to be considerable interest in the
Quaternary history of the Skagerrak–Kattegat (eastern North Sea) region and foraminiferal studies have
contributed significantly (e.g., Nordberg and Bergsten, 1988; Lykke-Andersen et al., 1993; Conradsen
and Heier-Nielsen, 1995). However, interpretations
of past environments depend upon comparison with
modern analogues. Although there are some studies of
the deeper water areas (Conradsen et al., 1994; Alve
and Murray, 1995, 1997), the ecology of the shallow
water foraminiferal assemblages has been little studied as noted below. This is a significant gap in knowledge, for recognition of shoreline deposits and former
sea levels are important aspects of environmental reconstructions especially in relation to palaeoclimate.
In this study, we systematically sampled the
coastal zone of the Skagerrak and Kattegat from
intertidal, with or without marsh, down to 6 m water depth in inner fjordic to open coastal waters,
to establish a detailed record of living (stained)
and dead (unstained) foraminiferal assemblages. The
live, down-core distribution was not investigated as
the main purpose has been to focus on the broader
distributional trends. In the present paper, we address the biogeographic distribution of taxa and the
ecological factors which control these patterns in the
living assemblages. The dead assemblages, with particular attention to the taphonomic processes which
modify the fossil foraminiferal record, are discussed
in Murray and Alve (1999a,b).
There are only a few published records of living shallow water benthic foraminifera along the
Skagerrak–Kattegat coast. Studies along the Norwegian Skagerrak coast (all Oslo Fjord) include Christiansen (1958), Risdal (1964), Alve and Nagy (1986,
1990), Alve (1990, 1995), studies on the Swedish
coast are Olsson et al. (1973), Nyholm and Olsson
(1973), Olsson (1976), Nyholm et al. (1977), Cato
et al. (1980). Except for the investigation of Hansen
(1965) in Øresund, there are no published records of
living shallow water assemblages along the Danish
coast of the Kattegat. All these studies deal primarily
with deeper water assemblages and generally there
are only brief comments on those living in shallow waters. There are no previous records of marsh
foraminiferal faunas.
Low salinity waters originating in the Baltic Sea
flow through the Danish Straits towards the northern
Kattegat and the North Sea (Fig. 1). Consequently, the
Kattegat and the Skagerrak waters form a transitional
zone between the Baltic Sea and the North Sea, with
surface salinities increasing from about 8–10‰ in the
southwestern Baltic Sea, to about 30‰ in the northern
Kattegat, due to entrainment from the higher salinity
bottom water (The Belt Project, 1981). The surface
water along the Swedish west coast has a salinity of
<30‰ and is dominated by Baltic Sea water (Lindahl
and Hernroth, 1983). The Jutland coastal water, with
salinities between 30 and 33‰, enters the Skagerrak–
Kattegat during periods of relatively strong winds
from the southwest (Pedersen et al., 1988) and can
be traced into the southwestern Kattegat (e.g., Aarhus
Bay, Lund-Hansen and Skyum, 1992; Lund-Hansen
et al., 1996). As the Baltic Water flows northwards,
it mixes with east-flowing Jutland coastal water at
the transition between the Kattegat and the Skagerrak
and turns westwards in the outer Oslo Fjord area as
the Norwegian Coastal Current. The degree of mixing between different shallow water masses increases
along the current direction and there is a generally increasing salinity gradient from the Kattegat and along
the Skagerrak coasts of Sweden and Norway (Anon.,
1997).
The area is microtidal with an astronomical tidal
range of 20–30 cm along the Skagerrak and about
40 cm along the Kattegat coast. This implies that
the general tidal effects cause virtually no mixing
of water masses and no transport of sediment (or
foraminiferal tests). However, meteorological effects
(surge, caused by wind and barometric pressure, 1
millibar D a change in level of 9.75 mm) generally
have a stronger impact on the prevailing sea level
in this area than the astronomical tide and ranges of
about 1 m occur several times a year along the Norwegian Skagerrak coast (Rune Braaten, pers. commun.,
1997). The general range of tides is less than 0.5 m
with the marshes being developed at the upper limit.
“The term (coastal) salt marsh is generally restricted to intertidal areas where the bottom (consisting of loose sedimentary material) is more or
less closely covered by a low, treeless vegetation of salt-tolerant or salt-requiring, predominantly
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
173
Fig. 1. Map of the Skagerrak–Kattegat with investigated areas shown by numbers from 1 to 27: 1 D Isefjærfjord; 2 D Kvastadkilen; 3 D
Dype Holla (Lyngør); 4 D Tøkersfjord (Lyngør); 5 D Hasdalen; 6 D Kilsfjord; 7 D Tjøme; 8 D Borre; 9 D Horten; 10 D Sandebukta;
11 D Bunnefjord; 12 D Hunnebotn; 13 D Hålkedalskilen; 14 D Tjärnö; 15 D Finnsbobukten; 16 D Gullmarsvik; 17 D Hafstensfjord; 18
D Kungsbackafjord; 19 D Jonstorp; 20 D Kildehuse; 21 D Kalundborg; 22 D Vejle Fjord; 23 D Havhuse (Kalø Vig); 24 D Vosnæs Pynt
(Kalø Vig); 25 D Løgstør; 26 D Valsted; 27 D Frederikshavn. Long arrows show general surface water circulation in the Skagerrak (after
Svansson, 1975). JCW D Jutland Coastal Water; BW D Baltic Water; NCC D Norwegian Coastal Current.
phanerogamic plants” (Fairbridge and Bourgeois,
1978). “Coastal marshes may be defined as areas,
vegetated by herbs, grasses or low shrubs, bordering
saline water bodies. Although such areas are exposed
to the air for the majority of the time, they are subjected to periodic flooding as a result of fluctuations
(tidal or non-tidal) in the level of the adjacent water
body” (Adam, 1990).
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E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
Table 1
Details of sample localities (salinity and temperature data from the time of collection unless otherwise indicated)
Area
Sample numbers
Salinity
(‰)
Temperature
(ºC)
Plants
Sampled
0 m: 4.7–20.8 a
5 m: 7.0–19.6 a
0 m: 0.2–21.0 b
5 m: 2.6–198 b
14.2–18.0
14.0–16.5
Zostera
14 July ’94
Zostera
4 July ’94
84–89
50–53
167
42–49
164–166
34–41
152–155
171
159–161
168–170
0 m: 0.7–30.0 a
5 m: 27.7–30.5 a
0 m: 0.1–28.6 b
5 m: 25.5–32.1 b
26–29
27–30
23
24–28
0 m: 3.9–31.2 a
5 m: 24.0–32.8 a
25.5 c
19–22
28
16–20
17
16
28
22–29 d 24
20
19
156–158
54–61
74–79
66–73
62–65
141–148
136–140
27
18–19
23–25
21–23
21–25
18–20
15
9.1
21.0–27.4
22.2–30.0
21.0–22.4
19.4–21.4
22.6–23.6
18.9
129–135
119–128
113–118
107–112
17–18
10–16
19–24
16–18
24–29 e
19–20
24–29 e
25
16
31
24.5
18.5–21.5
17.2–18.7
24.5–26.6
Potamogeton, Zostera
Salicornia
Zostera
Zostera
23 July ’94
23 July ’94
24 July ’94
21 July ’94
23.6–23.9
Zostera
21 July ’94
22.4
21.3
15.7
Cladophora
20 July ’94
20 July ’94
25 July ’94
Norway
1. Isafjærfjord
27–32
2. Kvastadkilen
14–19
3. Dype Holla, Lyngør
4. Tøkersfjord, Lyngør
5. Halsdalen
6. Kilsfjord
9–13
1–8
80–83
20–26
7. Tjøme
8. Borre
9. Horten
10. Sandebukta
11. Bunnefjord
12. Hunnebotn
Sweden
13. Hålkedalskilen
14. Tjärnö
15. Finnsbobukten
16. Gullmarsvik
17. Hafstensfjord
18. Kungsbackafjord
19. Jonstorp
Denmark
20. Kildehuse
21. Kalundborg
22. Vejle Fjord
23. Havhuse
Kalø Vig
24. Vosnaes Pynt
Kalø Vig
25. Løgstør
26. Valsted
27. Frederikshavn
a Bøhle
98–106
94–97
90–93
149–151
16.1–20.5
0 m: 2.8–16.8 a
5 m: 4.3–16.0 a
1–18 c
22
20.5–24.5
22.5
21.0–24.4
11.0
4–20 d 25
10.1
21.4
Zostera
Zostera
Zostera
Carex
Zostera
Phragmites, Carex
Zostera
Salicornia
Salicornia
Salicornia
Zostera
Potamogeton
Zostera
Potamogeton, Salicornia
3 July ’94
3 July ’94
25 July ’94
6 July ’94
8 Aug. ’94
19 July ’94
21 July ’96
18 July ’94
21 July ’96
18 July ’94
24 Oct. ’95
22 July ’96
25 Oct. ’95
22 July ’96
25 Oct. ’95
19 July ’94
20 July ’94
20 July ’94
20 July ’94
24 July ’94
24 July ’94
et al., 1989, 1990; b Bøhle, 1987; Bøhle et al., 1990; c Anon., 1997; d Magnusson et al., 1996; e Lund-Hansen et al., 1996.
In terms of classification based on vegetation, the
investigated marshes fall in the Scandinavian subgroup of the North European Group of Chapman
(1977) and in the boreal type of Adam (1990). In the
latter case, the examples from southern Norway and
Sweden are close to the boreal=temperate boundary
which lies in southern Sweden (Adam, 1990). Although marshes are known for plant zonation, there
is no regular pattern found in marshes in general.
The plants show spatial variation on three different
scales: micro-, meso- and macroscale. In the present
context, only the micro- and mesoscales are relevant.
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
On the microscale, patchiness of individuals of a
given plant species are on a scale of several centimetres. On a mesoscale, there may be zonation on a
scale of tens to hundreds of metres.
Apart from variables such as climate, elevation,
tidal range, wave and current energy, nutrients, etc.,
salinity is regarded as an important control. Elsewhere, the salinity of the sediment pore water does
not show any clear relationship with elevation and it
varies in a complex fashion both spatially and temporally (Adam, 1990). Apart from the input of marine
or brackish water from the sea, there is freshwater
input from the land. The presence of Phragmites
swamps has been attributed to pore water dilution by
freshwater (Adam, 1990).
Because of the very small tidal range, the vegetational zonation along the Skagerrak–Kattegat coast
is compressed and marshes are developed in small
patches (generally up to a few tens of m2 ) in sheltered bays and fjords. They have three dominant
phanerogamic plants: Phragmites australis, Carex
acuta, and Salicornia. In summer, the Phragmites
australis are commonly 2 m tall, the Carex acuta
about 50 cm and the Salicornia often not more than
10–15 cm. Each plant type may form a marsh by itself but sometimes the Carex acuta and Salicornia are
mixed. At the sediment surface, between the plants
shoots, there may be mats of filamentous green algae or cyanobacteria. At the landward margin there
is commonly several cm of leaf litter overlying the
moist sediment between the living marsh plants.
In this study, 27 geographical areas along the
Skagerrak–Kattegat coast were generally sampled
from the upper tidal limit to a water depth of 6
m (Fig. 1, Table 1). Areas 1–12 are located in
Norway, 13–19 in Sweden, and 20–27 in Denmark.
During the winter, for several months most of the
Skagerrak coast of Norway and the Kattegat coast of
Sweden is ice-covered to a depth of about 10–30 cm.
Consequently, the annual temperature range in the
nearshore, shallow water is from around 0 to >20ºC,
much more extreme than in the adjacent North Sea.
The Kattegat coast of Denmark is less subject to
freezing during the winter. Published hydrographic
information on shallow water areas is scarce and
that cited here generally represent data collected
<1 to several km away from the actual sampling
profiles. Because these shallow waters generally lack
175
tidal currents, they develop stratification during the
spring (when the snow melts) and during the warmer
summer months. The annual temperature range is
from at least 0 to 20ºC and salinities ranged from 10
(Kalundborg) to 30‰ (Frederikshavn) at the time of
sampling. The inshore waters are commonly clear,
enabling meadows of seagrass to develop, and this is
quite different from the open North Sea coasts.
3. Material and methods
3.1. Samples
A total of 171 sediment samples (each of around
100 cm3 ) were collected (see the Online Background Dataset 1 ), but two (162, 163) were not used.
The majority were collected in the month of July; the
exceptions were 84–89 (Aug.) and 152–159 (Oct.).
Collection was either with a small grab or, at exposed sites, simply by scraping off the oxygenated
surface 1–2 cm sediment. The grab was carefully operated so that negligible loss of surface sediment took
place. On removal from the water, the grab was gently
opened in a bowl and the surface sediment layer was
removed. All foraminiferal samples were preserved
in 70% ethanol. At most stations additional samples
were taken for grain size and total organic carbon
(TOC) analysis. Water temperature and salinity were
measured using a Salinity Temperature Bridge, type
M.C.5, or a thermometer and Atago salinity refractometer.
The water depths recorded are those at the time
of sampling. Because of the irregular tidal pattern, it
was difficult to know the exact position of average
sea level in the investigated areas. However, it was
clear that the water level was never very high or
very low because it was always within the range
of intertidal algal cover on rocks or jetties, i.e. the
maximum deviation from mean sea level was in the
range of about 20–30 cm at most stations. 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.
1
URL: http:==www.elsevier.nl=locate=palaeo; mirror site:
http:==www.elsevier.com=locate=palaeo.
176
PLATE I
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
Since the primary aim of this study has been to
investigate the environmental and biogeographic distribution of the various species, no effort was made
to collect quantitative samples representing an exact
sediment surface area nor to study vertical habitat partitioning between species. Fresh algal sheets
were collected at Hunnebotn (next to sample 169)
and Bunnefjord (next to sample 171) in order to
look at in-situ living foraminifera. These were studied directly under the microscope without further
preservation=processing.
3.2. Methods
In the laboratory, the foraminiferal samples were
washed on a 63-µm sieve (large fragments of organic
material were removed on a 1-mm sieve), stained for
1 h in rose Bengal (1 g=l), and washed again to
remove the surplus stain. Those samples not rich
in organic matter were dried at 40ºC, and then the
foraminifera were concentrated using carbon tetrachloride or tetrachloroethylene. Organic-rich samples were spread out over a large area (Lehmann and
Röttger, 1997) and allowed to dry without forming a
hard block. The dry material was then gently brushed
off into a container. It was not necessary to float such
samples in a heavy liquid.
For each sample, separate assemblages of living
and dead individuals were picked (ideally, at least
250 individuals in each, although some were too
small to yield this number). Of the 169 samples
studied, 47 yielded fewer than 50 live individuals
and are marked with a star in the Online Background Dataset 2 . Species diversity was calculated
for samples with >100 individuals using the Fisher
alpha index (Fisher et al., 1943) and the information
PLATE I
SEM photographs of calcareous taxa. The scale bar is 100 µm
for all except fig. 5 where it is 10 µm. # refers to sample number
(area number; see text Fig. 1).
1, 2.
Haynesina germanica (Ehrenberg), #94 (25).
3, 4.
Cibicides lobatulus (Walker and Jacob), #12 (3).
5–7. Rosalina anomala Terquem, #9 (3).
8–10. Elphidium excavatum (Terquem), #13 (3), live.
11.
Elphidium margaritaceum Cushman, #12 (3).
12, 13. Elphidium albiumbilicatum (Weiss), #94 (25).
14, 15. Elphidium oceanensis (d’Orbigny), #122 (21).
177
function H(S) (see Murray, 1991, for formula and
application).
Since many Reophax moniliformis tests were fragmented, only stained fragments with three or more
chambers were counted. Individuals of Goesella
waddensis lacking their initial chambers are impossible to distinguish from R. moniliformis so they
were counted together. However, where G. waddensis could be confidently identified, its presence was
noted in order to establish its distribution.
Sediment grain size analysis has been carried out
using standard sieving techniques, sieved on 63, 125,
250, 500, and 1000 µm meshes. TOC analyses were
made using the Leco combustion method.
SEM pictures of the most common calcareous
species are shown in Plates I and II, whereas those
of the agglutinated species are presented in Murray
and Alve (1999a,b).
4. Results
The species have been categorised according to
their environmental distribution based on presence or
absence of marsh plants, whether they are intertidal
or subtidal, and salinity (Table 2). Categories 1–5
comprise euryhaline and category 6 comprises essentially stenohaline species. We use the terms intertidal
and subtidal to differentiate between non-vegetated
areas which are frequently exposed (intertidal), even
though the exposure is not solely due to tidal cycles,
and areas which are not exposed (subtidal).
The category 1 species, Tiphotrocha comprimata and Balticammina pseudomacrescens, are
recorded only associated with marsh plants. Category 2, Jadammina macrescens, Trochammina inflata, and Haplophragmoides wilberti, comprises
species which are basically associated with marsh
plants but also occur outside the marshes. The category 3 species are characteristic of non-marsh areas,
but sometimes are present at the seaward edge of
marshes as noted below. The seven species listed
under category 4 are solely recorded in non-marsh
intertidal to subtidal environments, whereas category 5 comprises the euryhaline, subtidal species
2
URL: http:==www.elsevier.nl=locate=palaeo; mirror site:
http:==www.elsevier.com=locate=palaeo.
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E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
Table 2
Environmental and water depth distribution of principal living species (salinity measured at time of collection)
Species
Category
Depth range
(m)
Makes up
½5% of ass.
Makes up
½10% of ass.
As dominant
species
Salinity range
(‰)
B. pseudomacrescens
T. comprimata
1
IT*
IT*
IT*
IT*
IT*
IT*
IT*
IT*
20
20–28
T. inflata
J. macrescens
H. wilberti
2
IT–0.5
IT–1
IT–4
IT*
IT–0.3
IT–2
IT*
IT
IT*
ND
IT*
IT*
10–28
10–28
10–27
3
IT–1
IT–4
IT–6
IT–5
IT–6
IT–6
IT–1
IT–2
IT–5
IT–5
IT–6
IT–6
IT
IT–0.5
IT–5
IT–5
IT–4.5
IT–6
ND
IT
IT–3.5
IT–5
1–4.5
IT–6
16–26
15–27
10–29
10–31
15–31
15–31
E. pulchella
R. moniliformis
A. balkwilli
E. oceanensis
E. excavatum
P. (L.) haynesi
A. runiana
4
IT–4
IT–6
IT–5.5
IT–2
IT–6
IT–5
IT–6
IT–4
IT–5
IT–5
1–2
0.2–6
2
0.2–6
IT–4
IT–3
IT–4
1–2
0.4–6
2
2–5
IT–4
0.2–3
0.5
1–2
0.4–4.5
ND
2–2.5
15–27
15–28
15–29
15–24
15–31
18–28
16–28
O. kilianensis
E. scaber
A. cassis
5
1–5
2–4.5
3–6
3.5
–
3–6
–
–
3–6
ND
ND
ND
16–27
19–28
19–22
6
IT–4
IT–5
IT–5
1–5
IT–2
IT
2
4–5
2
–
–
4–5
2
ND
ND
5
24–28
25–28
26–28
24–28
E. albiumbilicatum
A. salsum
M. fusca
E. williamsoni
H. germanica
A. beccarii
C. lobatulus
E. margaritaceum
R. anomala
N. depressulus
Ophthalmina kilianensis, Eggerelloides scaber, and
Ammotium cassis, solely restricted to water depths
below 1, 2, and 3 m, respectively. Category 6, Cibicides lobatulus, Elphidium margaritaceum, Rosalina
anomala, and Nonion depressulus, are relatively
stenohaline and are found living only at stations
with >24‰ salinity at the time of sampling.
4.1. Marsh environments
Six marsh areas were sampled: Borre (area
8), Horten (9), Bunnefjord (11), Hunnebotn (12),
Hålkedalskilen (13), and Kalundborg (21) (Fig. 1,
Online Background Dataset 3 ). With the exception
of Borre and Hålkedalskilen, samples were collected
from dense Phragmites australis=Carex acuta marsh
3
URL: http:==www.elsevier.nl=locate=palaeo; mirror site:
http:==www.elsevier.com=locate=palaeo.
with some Salicornia on the seaward side. In most
instances, Phragmites australis was most abundant
landward of the Carex acuta marsh. The sediment
between the plants was usually covered with green
filamentous algae which could be removed in sheets.
The ‘marsh’ at Borre was only 1–2 m wide and there
was some green algal cover between the Carex acuta
but no Salicornia. At Hålkedalskilen, there were
only scattered Salicornia plants with no Phragmites
australis or Carex acuta. We include these localities under this heading even though they are not
well-developed marshes.
None of the typical marsh species of categories 1
and 2 was present at Borre. Elsewhere, J. macrescens
is invariably present but the associated species differ from one area to another. Balticammina pseudomacrescens was recorded only at Hunnebotn, T. comprimata at Horten, Bunnefjord and Hunnebotn, T. inflata at all except Borre, and H. wilberti at Horten,
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
179
Table 3
Geographical distribution of living species arranged in rank order
Area
Norway
E. williamsoni
M. fusca
A. beccarii
H. germanica
R. moniliformis
E. excavatum
A. runiana
A. balkwilli
A. salsum
J. macrescens
O. kilianensis
P. (L.) haynesi
E. oceanensis
H. wilberti
E. scaber
T. inflata
E. albiumbilicatum
E. pulchella
A. cassis
T. comprimata
C. lobatulus
E. margaritaceum
N. depressulus
B. pseudomacr.
R. anomala
a Number
Sweden
1 2
3
4 5
ð
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6 7
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Denmark
8 9
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
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No. of
areas a
ð ð ð 27
ð
25
ð ð ð 24
ð ð ð 24
19
ð ð ð 19
18
16
ð
14
ð ð
11
ð
9
ð
7
ð
7
7
6
5
ð
4
4
3
3
2
ð
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2
1
1
of areas in which the species occur.
Hålkedalskilen, and Kalundborg (Table 3). The category 3 species, Miliammina fusca and Elphidium
williamsoni, are also present (and may even dominate) at the seaward edge of any marsh, Ammonia beccarii and Haynesina germanica occur in the ‘marsh’
at Borre, Ammotium salsum and Elphidium albiumbilicatum are present at the seaward edges of the marsh
at Horten, and the former also at Hunnebotn.
The number of species ranges between 3 and 8
but is generally 4–6. The Fisher alpha and H(S)
species diversity values are 0.7–1.7 and 0.5–1.3,
respectively.
Examination of the fresh algal sheet from Hunnebotn (next to sta. 169) revealed the presence of
living individuals of J. macrescens and B. pseudomacrescens clinging in random orientation to the
filaments, while T. comprimata was more firmly attached by its umbilical side. Jadammina macrescens
has its maximum abundances on the degraded sur-
faces of decaying Carex acuta leaf debris. The same
was true for the algal sheet from Bunnefjord (next
to sta. 171) except that B. pseudomacrescens was not
observed and the density of individuals (although not
quantified) was several times higher.
The sediment median diameter shows a wide
range (from phi 0.2 to C4.2; very coarse sand to
coarse silt; see the Online Background Dataset 4 ).
The TOC values analysed in six of the fourteen
marsh samples, vary from 0.4 to 14.8%.
4.2. Non-marsh intertidal and subtidal environments
Samples were collected from non-marsh environments in all 27 areas and the live occurrence of
the principal species are summarised in Table 2,
4
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http:==www.elsevier.com=locate=palaeo.
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E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
PLATE II
PLATE II
SEM photographs of calcareous taxa. The scale bar is 100 µm for all except for figs. 3, 5 and 11 where it is 10 µm. # refers to sample
number (area number; see text Fig. 1).
1.
Ammonia beccarii (Linné), #80 (5).
2, 3.
Nonion depressulus (Walker and Jacob), #94 and 12 (25 and 3, respectively).
4.
Ophthalmina kilianensis Rhumbler, #11 (3), live.
5.
Eoeponidella pulchella (Parker), #9 (3).
6.
Elphidium williamsoni Haynes, #77 (15), live, typical variety.
7.
Elphidium williamsoni, #55 (14), typical variety.
8–11. Elphidium williamsoni, #23 (6), variety with fewer and shorter retral processes, all live showing etched surfaces and enlarged
pores.
11.
Detail of fig. 10.
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
categories 4–6. In Table 2 a distinction is made between presence and dominance. Although category
3 and 4 species are present in both intertidal and
subtidal environments, assemblages dominated by
certain taxa are more restricted. Some category 3
assemblages dominated by Miliammina fusca, Ammonia beccarii, Elphidium williamsoni or category
4 Eoeponidella pulchella, occur both in the intertidal and subtidal areas. Other assemblages are confined to subtidal areas where the dominant species
are either category 3 (Haynesina germanica), category 4 (Ammobaculites balkwilli, Ammoscalaria runiana, Reophax moniliformis, Elphidium excavatum,
E. oceanensis), or category 6 (Cibicides lobatulus,
Nonion depressulus).
Most live individuals of E. williamsoni in Kilsfjorden (area 6) show clear signs of chemical etching
(Plate II). Severe etching of living individuals is
also common in area 2. This is only reported as an
observation as it is discussed in Murray and Alve
(1999b).
The number of species ranges between 1 and 16
but is generally 4–9. The greater numbers, 13–16,
were recorded at Lyngør and Tjøme. The Fisher
alpha and H(S) species diversity values are 0.3–
181
4.0 (>2.7 only at Lyngør and Tjøme) and 0.1–1.9,
respectively.
The sediment median diameter is in the range
phi 0.4 to C4.5 (very coarse sand–coarse silt).
The TOC values vary from 0.1 to 39.1%, but the
maximum values (>9%) are only found in areas 1
and 2. The relative abundance of the most commonly
occurring species plotted against water depth, per
cent mud (clay and silt <63 µm) in the sediments,
and TOC are shown in Figs. 2–4, respectively.
4.3. Colour of the cytoplasm
After staining with rose Bengal, apart from the
red-stained final chamber, all E. albiumbilicatum, N.
depressulus, E. margaritaceum, and the vast majority
of E. williamsoni had a pale cream colour while only
scattered specimens of the latter were pale green in
the earlier chambers. Most A. beccarii and H. germanica were also pale cream but some individuals
of the former graded into brown whereas some of
the latter were light green. No particular colour was
seen in E. oceanensis but E. excavatum was usually
distinctly brown.
Fig. 2. Depth distribution of mean relative abundances of living species. The data are averaged over 1-m water depth intervals. Marshes
are treated as a separate category. Only those species with a mean value of ½20% within one or more depth intervals are included.
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E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
Fig. 3. Relative abundance of common species versus % mud (<63 µm) in the sediment.
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
Fig. 4. Common species versus TOC %. Note that the single sample with 40% TOC is omitted from the diagrams.
183
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E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
4.4. Rarer species
Living species recorded only as scattered individuals are Nodulina dentaliniformis, Psammosphaera bowmanni, Textularia truncata, Buliminella
elegantissima, Cornuspira involvens, Elphidium macellum, Fissurina marginata, Gavelinopsis praegeri,
Guttulina lactea, Pateoris hauerinoides, and some
unidentified textulariids, trochamminids, elphidiids,
Brizalina sp, Ammonia sp., and Quinqueloculina sp.
They are listed as ‘other agglutinated’ and ‘other
calcareous’ in the Online Background Dataset 5 .
Of these, T. truncata, C. involvens, E. macellum, G.
praegeri, and G. lactea are recorded only at Lyngør
(areas 3, 4).
5. Discussion
Marginal marine environments are very variable
in terms of physico-chemical conditions because
they respond to diurnal, seasonal and even longerterm changes. In a sense, the only constant attribute
of marginal marine environments is their variability.
As noted in Section 3.1, the majority of samples
were collected in July of successive years. Consequently, there should not be any major differences
caused by seasonality.
5.1. Marsh assemblages
In recent years much attention has been given
to using marsh foraminiferal assemblages as indices of former sea levels (e.g., Scott and Medioli,
1986; Williams, 1994; Guilbault et al., 1996). Most
marshes are found in tidal regions but even in the
essentially non-tidal Skagerrak–Kattegat there are
small patches of Phragmites australis, Carex acuta,
and Salicornia marsh developed in sheltered areas.
Unlike tidal marshes, these have no regular tidal
movement of water but they experience short-term
fluctuations related to winds and seiches. With the
exception of the small marsh at Borre and the most
seaward sample at Horten, the marsh assemblages
are strongly dominated by organo- and ferro-aggluti5
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http:==www.elsevier.com=locate=palaeo.
nated taxa (i.e., none with a calcareous cement). The
main species (J. macrescens, T. inflata, M. fusca) are
cosmopolitan (summary in Murray, 1991), whereas
T. comprimata and B. pseudomacrescens are reported
from this area for the first time.
Marsh environments have commonly been divided into high and low marshes associated with
characteristic dominant species and even more refined faunal subdivisions have been used in many
areas (e.g., Scott and Medioli, 1980; Williams, 1994;
Ozarko et al., 1997). Due to the small tidal range
and compressed nature of the vegetational zonation
in the Skagerrak–Kattegat area, refined subdivisions
are meaningless here. Despite this compressed zonation, the commonly recorded faunal distributional
trend is clearly present with J. macrescens and M.
fusca dominating the landward and seaward side of
the marshes, respectively. In contrast to most larger
marshes, which often show an irregular topography due to drainage creek systems, the surface of
these small marshes shows a generally decreasing
elevation from about 10–40 cm above mean sea
level down to the essentially non-vegetated, intertidal, more or less muddy sands.
The relative importance of salinity versus elevation above mean sea level for the distribution of
marsh species is a matter of current debate (e.g.,
Scott and Medioli, 1980; Scott et al., 1990; De Rijk,
1995; De Rijk and Troelstra, 1997). In the Skagerrak–
Kattegat area, the tides are too small to allow investigations about the importance of elevation. However, it is reasonable to assume that the sediment
pore water is relatively more influenced by infiltration of rain water than sea water at the most elevated sites compared to the lower sites, leaving the
former less saline. Balticammina pseudomacrescens
has been recorded in only one area (12) and shows
an increasing relative abundance towards the most
elevated parts of the marsh. Additionally, B. pseudomacrescens has recently been found living associated with damp, rotting leaf litter close to the upper
tidal limit in the River Cur, near Southampton, U.K
(Murray and Alve, unpublished data). In the work
of Scott et al. (1990) their Trochammina macrescens
f. macrescens is equivalent to B. pseudomacrescens
and their Trochammina macrescens f. polystoma is
equivalent to J. macrescens sensu Brönnimann and
Whittaker (1984) (D.B. Scott, pers. commun., 1997).
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
Also, De Rijk (1995) previously reported B. pseudomacrescens as Trochammina macrescens type A
(De Rijk and Troelstra, 1997). Balticammina pseudomacrescens should be a more low salinity species than
J. macrescens (De Rijk, 1995; De Rijk and Troelstra,
1997). This fits remarkably well with the distribution
of the two species in the present study area.
Investigation of the fresh algal sheets and the
degraded surfaces of decaying Carex acuta leaf debris revealed abundant, free living J. macrescens
with maximum abundances (although not quantified) on the latter. This reflects a typical epiphytal
mode of life and its high abundance on the decaying phanerogam leaf debris indicates that it probably
feeds on bacteria and decay products from rotting
plants, in addition to grazing on fresh algae. The
majority of marsh-dwelling macro-invertebrates are
either detritivores or they consume the microflora
(algae, cyanobacteria, bacteria) (Adam, 1990) and
the same is probably true of the foraminifera. An
epiphytal life style for J. macrescens was also observed by Matera and Lee (1972) but whereas we
also recorded abundant living individuals within the
sediments (between the plants), they did not record
it in the psammolittoral communities living in the
sediments below the plants. Recently, it has been
recorded living in sediments down to 30 cm below
the surface (Ozarko et al., 1997). In summary, these
findings imply that J. macrescens is a generalist,
with both an epiphytal as well as an infaunal mode
of life. However, regardless of whether it lives within
or above the sediment it seems to be able to flourish only in vegetated environments. The latter also
seems to hold for the other species of categories 1
and 2 (Table 2).
Trochammina inflata was abundant in only two areas (11 and 21), in sediments associated with marsh
plants but it was not recorded on the examined plant
fragments. Morphologically, T. inflata strongly resembles Ammonia beccarii in that it has the aperture
on the umbilical side, it is inflated to an extent that
it can be regarded as biconvex, and it has a large
interlocular space (deep umbilical cavity) which, according to Langer et al. (1989), is beneficial for locomotion. These features are in contrast to the attached,
immobile, trochospiral epifaunal species (e.g., Cibicides lobatulus, Rosalina spp; see Kitazato, 1988)
which are typically planoconvex. Although A. bec-
185
carii and T. inflata have been recorded in small
numbers in epiphytic communities (Matera and Lee,
1972), they are most commonly found living within
the sediments (e.g., Steineck and Bergstein, 1979;
Langer et al., 1989; Goldstein and Harben, 1993;
Ozarko et al., 1997) and none of them seems to live
in symbiotic association with algae or algal chloroplasts (Knight and Mantoura, 1985). Both in culture
and field studies, Langer et al. (1989) observed A.
beccarii to dig itself into the sediment surface with
a cork-screw movement, and the direction of the
rotation corresponds to the growth direction. It is
possible that this morphological similarity between
the agglutinated marsh species and the calcareous,
essentially intertidal species reflects adaption to the
same mode of life, just in different environments.
The wide range in sediment grain size and sediment TOC does not seem to have any obvious impact
on the assemblage compositions in the limited number of marsh samples where these were recorded.
5.2. Non-marsh intertidal and subtidal assemblages
Except for the most extremely marsh-bound
species of category 1 (Table 2), there are no species
solely restricted to the non-marsh intertidal environment. On the other hand, the three species of category 5 are completely restricted to and characteristic
of subtidal areas.
In the shallow waters of the Skagerrak–Kattegat
area, there are the typical shallow water species E.
williamsoni, M. fusca, and H. germanica, together
with species that are known to be widely distributed
onto continental shelves elsewhere, A. beccarii and
E. excavatum. These are the most abundant and
widely distributed species both environmentally and
geographically in the study area. Elphidium excavatum is common (>10% of living assemblage) only
in subtidal areas, whereas the others are common
from intertidal to 5–6 m water depth. However, assemblages dominated by E. williamsoni and M. fusca
are basically restricted to water depths of less than
2 m, those dominated by E. excavatum to 2–4 m,
whereas A. beccarii can dominate throughout the investigated depth interval. Although present in most
areas, H. germanica is dominant only in five samples, and generally more common along the Swedish
and Danish coasts than along the Norwegian coast.
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All five are well known euryhaline species (e.g.,
Murray, 1973, 1983), M. fusca being extremely so,
as it tolerates salinities of <1–35‰, but although
it is most common in hyposaline environments, no
correlation has been found between its frequency
distribution and salinity (De Rijk, 1995). The M.
fusca biofacies (unstained samples) in the Lower
Low Marsh Zone in the Fraser River Delta, British
Columbia, was generally found only in areas on tidal
flats where Zostera marina (eel grass) was present
to stabilize the substrate (Patterson, 1990). In the
Skagerrak–Kattegat it seems to live in all the environmental types investigated, including the Zostera
belt, but we did not record it on the algae or Carex
leaves. Ozarko et al. (1997) characterize M. fusca as
being primarily epifaunal to very slightly infaunal,
but it is not clear whether they actually observed it
attached or clinging to objects elevated above the
sediment surface or if they used the definition by
Corliss (1991) of the term epifaunal (i.e., the upper
0–1 cm of the sediment). On the other hand, Wefer
(1976) recorded it attached to algae where it could
make up to 50% of the epiphytic assemblage. It
has also been observed clinging to filamentous algae
from an intertidal marsh near Southampton, England
(Murray, unpublished observation).
No species showed any clear linear correlation
with sediment grain size distribution or TOC. However, some patterns are worthy of mention. Ammoscalaria runiana was common only in sediments
with less than 20% mud, a median diameter in the
medium to fine sand fraction, and low TOC values (0.2–0.7%), whereas abundant populations of E.
excavatum, A. beccarii, and H. germanica lived in
sediments with strongly variable TOC values and a
mud content ranging from <5 to >80% (Figs. 3
and 4). Elphidium williamsoni was never common in
sediments with >60% mud.
Elphidium albiumbilicatum, a typical brackish
water species (Lutze, 1965), occurs around the Skagerrak and in the Limfjord, Denmark, but basically
only down to 1 m water depth. On the other hand,
Lutze (1965) recorded it (as Cribrononion asklundi)
living down to >30 m water depth in the brackish
Baltic Sea, and in Drammensfjord, Norway, it lives
most commonly at 7–13 m in the lower part of the
brackish surface layer (Alve, 1995). This distributional pattern is a good example illustrating how hy-
drographical parameters, here probably mainly salinity rather than water depth, control the distribution.
This species has not been reported from the North
Sea. Therefore, its southern limit of distribution is in
the Kattegat–Baltic Sea.
Reophax moniliformis and Elphidium oceanensis
probably feed on plant debris (higher plants and
green algae) and the pigments in R. moniliformis,
E. williamsoni and H. germanica indicate that they
live in symbiotic association with diatoms or their
chloroplasts (Knight and Mantoura, 1985). On the
other hand, E. oceanensis does not seem to have this
symbiotic relation but they may concentrate algal
carotenoids with which they frequently cover themselves and living specimens appear bright red when
isolated from their environment (Knight and Mantoura, 1985). The colour of living E. williamsoni
(from southern England) studied by Knight and
Mantoura (1985) varied considerably, most being
dark or light green, but pale cream and orange
or red were also recorded and they showed that
these variations correspond to real differences in pigment content. Their H. germanica varied from dark
green to pale cream. In contrast, the vast majority of E. williamsoni and most H. germanica in the
Skagerrak–Kattegat area are pale cream coloured.
These differences possibly reflect seasonal differences in floral composition as indicated in a recently
completed seasonal sampling programme of H. germanica near Southampton, southern England (Murray and Alve, unpublished data). It might be argued
that the rose Bengal stain and subsequent drying of
the specimens can weaken the colour appearance of
the pigments but this does not seem to be the case,
as bright green individuals were commonly found in
our seasonal study. In an experimental study, Murray
(1963) determined that the colour of the cytoplasm in
Elphidium crispum was directly related to the colour
of the algal food.
Ammotium cassis characterizes vertically moving
discontinuity layers (e.g., Lutze, 1965) which may
cause organic material to be thrown into suspension
and thereby cause additional food supply (Olsson,
1976). In the Baltic, Lutze (1965) found A. cassis
at a depth of 17–23 m in the transition between the
surface water mass (14–19‰ salinity) and the deeper
water (20–24‰ salinity), while in Drammensfjord,
Norway, living individuals were present only at the
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
upper and lower border of the transitional water mass
with maximum frequency at the upper border (12.8–
25.9‰ salinity, Alve, 1995). Olsson (1976) recorded
peak abundances in connection with discontinuity
layers at 4–5 m, 12–20 m, and 25–30 m water depth
in Swedish estuaries. In the present study area, living
A. cassis was recorded only subtidal (½3 m water
depth) with a salinity range of 19–22‰ in the lower
part of or just below the Zostera belt, and it was
common in sediments with a wide range in mud and
TOC content (1–83% and 0.5–4.3%, respectively).
Only scattered living individuals of Eggerelloides
scaber were recorded at some of the Norwegian,
subtidal stations but higher abundances were found
in the corresponding dead assemblages (Murray and
Alve, 1999b). The apparently limited distribution is
due to the fact that we collected only from shallow
water (down to 6 m). Eggerelloides scaber requires
at least 24‰ salinity for a longer part of the year
(Lutze et al., 1983). It is otherwise a common species
in these deeper subtidal areas (e.g., Conradsen, 1993;
Alve, 1995).
Eoeponidella pulchella seems to be an euryhaline,
attached species characteristic of high-energy, intertidal to subtidal environments, as living specimens
were only found in sandy sediments (<6% mud) associated with wave-exposed beaches. These findings
agree with the single record by Lutze (1965) of this
species (as Asterellina) in the Baltic Sea.
Ophthalmina kilianensis is an epifaunal species
which settles solely on algae (Wefer, 1976). In the
western Baltic Sea it was typically associated with
red algae where Lutze (1965, p. 94) recorded abundances of about 100 individuals per 10 cm2 plant
surface. Wefer recorded only a few living individuals
in the sediments and thought these to have fallen off
the algae. This is probably also the case with the
few individuals (max. 6% of live assemblage in Kalø
Vig area) we recorded in the Skagerrak–Kattegat
sediments.
The nearly stenohaline species of category 6 basically live in the most open marine areas and show
restricted distributions. They are: Cibicides lobatulus, Rosalina anomala, Nonion depressulus, and Elphidium margaritaceum. The former two are known
to live epifaunally on substrates such as shells, hydroids, algae, bryozoans and other structures raised
above the sea floor. Because of their elevated, at-
187
tached mode of life, the presence of living individuals in the sediment is relatively uncommon. Elphidium margaritaceum was recorded only in the higher
salinity areas at Lyngør and Løgstør (areas 3, 25).
Nonion depressulus is much less tolerant to reduced salinities than truly estuarine species (Murray,
1983) which is probably the reason why we recorded
it only in the two most offshore areas on the Norwegian Skagerrak coast (areas 3, 4). Here, it was rare in
the landward area (4) but actually dominated one of
the samples on the island side (3) which also showed
one of the highest salinity values (28‰) recorded
throughout the study. The more marine character of
the Lyngør areas is also reflected by the fact that the
species diversities reach maximum values here and
by the presence of typically marine species (see Section 4) which are not recorded in the other areas. This
is also the only area with frequent Paratrochammina
(Lepidoparatrochammina) haynesi which has been
recorded living attached to larger objects (>1 cm)
such as stones and bivalve shells on the sea floor
exposed to strong tidal currents and in salinities of
32–33‰ (Murray and Alve, 1993; Alve and Murray,
1994).
The species diversity of the living assemblages is
low and this is consistent with other marginal marine
environments (summarised in Murray, 1991).
5.3. Biogeography
Living species not previously reported in the literature from the Skagerrak or Kattegat include Ammotium salsum, Balticammina pseudomacrescens,
Ophthalmina kilianensis, Elphidium oceanensis,
Tiphotrocha comprimata, Trochammina inflata, and
Eoeponidella pulchella. However, the latter three
species were recorded living by Christensen in Kalø
Vig (areas 23, 24), Denmark in 1971 (K.L. Knudsen,
pers. commun., 1996). All these species have now
been found living as far afield as the northern part of
the Skagerrak. None of the recorded living species
seem to reach their overall northern or southern distribution limit within the investigated area. They are
either distributed throughout the area or their distribution is limited due to their restriction to particular
local environmental characteristics.
Because marshes are restricted in their development, the occurrence of species which are entirely
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E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
associated with marsh plants, T. comprimata and B.
pseudomacrescens (category 1), is also patchy (Table 2). The marsh species of category 2, T. inflata,
H. wilberti, J. macrescens, are more widespread,
probably because they are not just associated with
the marsh vegetation proper. The most widespread
species belong to categories 3 and 4, whereas those
of category 5 have a more limited distribution because our records only represent the shallowest limit
of their distribution.
The majority of the common euryhaline species
are cosmopolitan and occur widely both in the
adjacent North Sea and in the Baltic (summarized in Murray, 1991). However, there is clearly
a faunal boundary between the Kattegat and the
Baltic Sea. The following species which occur
in the Kattegat have not been reported from the
Baltic: Ammobaculites balkwilli, Goesella waddensis, Haplophragmoides wilberti, Paratrochammina
(Lepidoparatrochammina) haynesi, Reophax moniliformis, Elphidium oceanensis, and Haynesina germanica. In view of the intensive study of the
Kiel Bay area in particular (Rhumbler, 1935, 1936;
Rottgardt, 1952; Lutze, 1965, 1968, 1974; Grabert,
1971; Wefer, 1976) it seems likely that these absences are real. Some can be explained by the low
salinity (P. (L.) haynesi, E. oceanensis, and possibly
A. balkwilli). For others, like H. germanica, which is
strongly euryhaline, the explanation is less obvious
but perhaps it needs higher salinities at least during
parts of the year for successful colonisation. Extreme
temperature conditions may also be a limiting factor or perhaps these species have not yet succeeded
in migrating into the area. Living H. germanica often has a characteristic vivid greenish colour and
lives in symbiotic association with diatoms or their
chloroplasts (Knight and Mantoura, 1985). In general, Baltic species live at greater water depths than
is normally recorded in other areas because of the
wider depth range of the brackish surface water.
Consequently, the absence of H. germanica in the
Baltic might be due to adverse light conditions below the permanently low salinity (<10–15‰) upper
part of the surface water.
Except for C. lobatulus (which also lives in
subarctic=arctic areas, Hansen and Knudsen, 1995)
the essentially stenohaline group are southern
species reaching their northeastern limits of distri-
bution although the cause of this is due largely to the
brackish salinities of the area. Stenohaline species
showing restricted distributions include: C. lobatulus, Nonion depressulus, R. anomala, and E. margaritaceum. They are abundantly present in Dype
Holla (area 3), and they typically occur in areas with
direct contact to the open Skagerrak (i.e., they are
not found in the inner parts of fjords). At Dype
Holla, they are also associated with normal marine
species (see Section 4) which are not found living in
any of the other areas. Here the sediments are shell
sands and the assemblages resemble those of similar sediments in the English Channel (Sturrock and
Murray, 1981, groups 1 and 3). Cibicides lobatulus
was not recorded by us in the Kattegat. However,
some living occurrences have previously been reported from the southern Kattegat (Conradsen, 1993)
and rare individuals were attached to red algae at 20
m water depth in the western Baltic (Lutze, 1965).
This implies that even though its main distribution is
in normal marine areas, minor populations may enter estuarine environments where there are suitable
weed substrates. Nonion depressulus, R. anomala,
and E. margaritaceum have not been reported from
the Baltic.
Cushman (1920) and Phleger and Walton (1950)
considered Ammotium cassis to be an arctic species.
According to Wefer (1976) it reproduces when the
temperature is <3ºC. Because Rhumbler (1935,
1936) did not record it in the Baltic Sea and
Rottgardt (1952) found only a few individuals, Lutze
(1965) considered it to be a recent introduction into
the Baltic. Olsson (1976) noted that this species
was not recorded either by Christiansen (1958) in
the Oslo Fjord or by Höglund (1947) in the Gullmar
Fjord. However, only a few years after Christiansen’s
investigation, Risdal (1964) recorded it as a dominant species at two shallow stations (3 and 4 m water
depth) in the Oslo Fjord. Thus, the idea of it being a
new introduction might apply for the Baltic, whereas
no conclusion can be drawn concerning the fjordic
areas of the Skagerrak and Kattegat.
Not surprisingly, there seems to be a delay between naming a species and determining its distribution. This applies both to A. balkwilli and to B.
pseudomacrescens. Ellison and Murray (1987) noted
that at that time A. balkwilli was known only from
the type locality (Dovey estuary, Wales, 1973) and
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
the Santoña estuary (N. Spain). However, living individuals had also been recorded at various localities
in the Oslo Fjord (as A. agglutinans by Risdal, 1964;
Alve and Nagy, 1986). Now it is also known from
southern England (Alve and Murray, 1994) and living specimens have been recorded in 16 of the 27
areas studied in the present investigation. Balticammina pseudomacrescens was described from, and
thought to be endemic to, the Baltic Sea (Brönnimann et al., 1989) but has recently been recorded
in high marshes in the Americas (see Section 5.1)
and it is abundant at one of our localities (area 12)
in the Oslo Fjord. Additionally, we have now found
it associated with rotting leaves in the highest part
of a Phragmites marsh in the River Cur, southern
England (Murray and Alve, unpublished data). It is
very significant that it is found in the most landward=
terrestrial parts of the intertidal zone and thus defines
the uppermost limit of the influence of marine water.
In summary, the Skagerrak–Kattegat coastline
does not show any marked biogeographic boundaries. The fact that some species are found in more
areas than others, is due to local environmental differences, e.g. presence=absence of a marsh, varying degrees of wave action, etc. This implies that
the most widely distributed species are those best
adapted to the most commonly occurring environment, which along the temperate Skagerrak–Kattegat
shore line is sand with a strongly variable mud content, usually partly covered by seagrasses and=or
Fucus attached to cobbles and large shells. There
is, however, a faunal boundary between the Kattegat
and the brackish Baltic Sea as 10 of the 25 species
which are commonly living in the former area have
not been reported from the Baltic.
6. Summary and conclusions
Analyses of living (stained) benthic foraminifera
collected from a range of different shallow water
environments (marsh to 6 m water depth) in 27 areas along the Skagerrak–Kattegat coast have given
a detailed distribution pattern of the 25 most common species (13 agglutinated; 12 calcareous). The
foraminiferal assemblage data are based on 169 sediment surface samples and some additional fresh
samples of marsh and subtidal plant material which
189
were also analysed. The salinity and temperature
were in the ranges 10–31‰ and 9–30ºC at the time
of sampling (July, August, October) but the temperature gets down to freezing during the winter. The
sediment grain size and total organic carbon (TOC)
content are extremely variable.
The species have been divided into six environmental categories based on presence or absence of
marsh plants, whether they are intertidal or subtidal,
and salinity. Categories 1–5 comprise euryhaline and
category 6 comprises essentially stenohaline species.
The marsh vegetational zonation is compressed,
due to the small tidal range (<40 cm), but the
species still show a general distributional trend with
Jadammina macrescens and Miliammina fusca dominating the landward and seaward sides, respectively.
Balticammina pseudomacrescens is reported both
from this area and from England for the first time. It
tolerates even lower salinities than J. macrescens
sensu Brönnimann and Whittaker (1984) and is
found in the most landward=terrestrial parts of the
intertidal zone.
Living M. fusca and Elphidium williamsoni were
recorded in all the investigated environments from
marsh, through non-vegetated intertidal, to subtidal areas in both sheltered, inner fjordic, and open
coastal settings, reflecting their wide environmental tolerance. The same distributional pattern was
recorded for Ammonia beccarii and Haynesina germanica, except that they were not found in the
typical marsh environments. Elphidium excavatum is
common only in subtidal areas although it is present
in low abundance in the intertidal zone.
The environmental data for the marsh samples
show very variable values and they are too few to
draw any firm conclusions about their significance
for the species distribution. None of the intertidal–
subtidal species showed any clear linear correlation
with sediment grain size or TOC. However, Ammoscalaria runiana was common only in medium to
fine sand sediments of low TOC. On the other hand,
E. excavatum, A. beccarii, and H. germanica lived
abundantly in a wide range of sediment types with
median values from coarse silt to very coarse sand,
and a variable TOC content, and showed the greatest
tolerance to substrate variability.
The number of living species at each station was
generally less than 9 but higher values of 13–16 were
190
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
recorded in the most open marine environments,
particularly at Lyngør, where normal marine species,
not present in the other areas, were found. The
species diversity values are low (Fisher alpha 0.3–
2.7, except for values of 2.7–4.0 at Lyngør, and H(S)
0.1–1.9).
The recorded distribution of shallow water species
along the Skagerrak–Kattegat coast has been extensively widened and the results show that there are
no marked biogeographic boundaries within the area.
However, several species, particularly those of categories 1, 5, and 6, show restricted distributions due to
local environmental conditions or simply to limited
sampling in particular environments. The following
species which live in the Skagerrak–Kattegat area
have not been reported from the brackish Baltic Sea:
Ammobaculites balkwilli, Goesella waddensis, Haplophragmoides wilberti, Reophax moniliformis, Haynesina germanica, Paratrochammina (Lepidoparatrochammina) haynesi, Elphidium oceanensis, Nonion depressulus, Rosalina anomala, and Elphidium
margaritaceum. The latter six are probably restricted
by the low salinity waters. It is very significant that
B. pseudomacrescens is found in the most landward=
terrestrial parts of the intertidal zone and thus defines
the uppermost limit of the influence of marine water. Ammotium cassis, Ophthalmina kilianensis, and
Elphidium albiumbilicatum have not been recorded
from the North Sea proper implying that the Kattegat
might be their southern limit.
Acknowledgements
We are pleased to acknowledge the Natural Environment Research Council for grant GR9=1591’A’
and the Industrial Liaison fund (Oslo University) for
financial support. We are grateful to Pål Glad for
help with sampling in Norway, to Eva Larsen, Siggurd Gausdal, Arne M. Svendsen, Bent Åsnes, Trond
Smith, John Ingebrigtsen, and Tjärnö Marinbiologiska Laboratorium for kindly letting us use their
boats, to Inger-Ann Hansen for performing the TOC
analyses, and to Daphne Woods for doing the grain
size analyses. Karen Luise Knudsen, Marit-Solveig
Seidenkrantz, Sacha DeRijk, and John E. Whittaker are kindly thanked for taxonomic help, and
Leif Ryvarden for advice on the marsh plants.The
manuscript was reviewed by Karen Luise Knudsen
and Sacha De Rijk and by referees Bill Berggren,
Jean-Pierre Debenay and Ron Martin; we kindly
thank them all for their comments.
Appendix A. Comments on taxonomy
Ammonia beccarii is a group represented by several variants
(e.g., Schnitker, 1974; Chang and Kaesler, 1974) and in the study
area, tepida is the principal one. Typical individuals of batavus
are present only in the dead assemblages of area 3. On the other
hand, there are numerous individuals throughout the area which
show some of the characteristics of both. Consequently, we have
not distinguished between them.
The Elphidium excavatum group comprises several forms
(see for example Feyling-Hanssen, 1972; Poag et al., 1980;
Miller et al., 1982), but in the present study the individuals seem
transitional so we have not used form names.
Most specimens of Elphidium williamsoni have unusually
short retral processes (Plate II, figs. 8–1) compared with what is
normally found in this species (Plate II, figs. 5 and 6).
Appendix B. Faunal reference list
Generic names are in accordance with Loeblich and Tappan
(1987) with the exception of Pateoris which is retained because
of common usage.
Ammobaculites balkwilli Haynes, 1973
Ammoscalaria runiana (Heron-Allen and Earland) D Haplophragmium runianum Heron-Allen and Earland, 1916
Ammotium cassis (Parker) D Lituola cassis Parker, 1870
Ammotium salsum (Cushman and Brönnimann) D Ammobaculites salsus Cushman and Brönnimann, 1948
Balticammina pseudomacrescens Brönnimann, Lutze and Whittaker, 1989
Eggerelloides scaber (Williamson) D Bulimina scabra Williamson, 1858
Goesella waddensis van Voorthuysen, 1960
Haplophragmoides wilberti Anderson, 1953
Jadammina macrescens (Brady) D Trochammina inflata (Montagu) var. macrescens Brady, 1870
Miliammina fusca (Brady) D Quinqueloculina fusca Brady, 1870
Nodulina dentaliniformis (Brady) D Reophax dentaliniformis
Brady, 1884
Paratrochammina (Lepidoparatrochammina) haynesi (Atkinson)
D Trochammina haynesi Atkinson, 1969
Psammosphaera bowmani Heron-Allen and Earland, 1912
Reophax moniliformis Siddall D Reophax moniliforme Siddall,
1886
Textularia truncata Höglund, 1947
Tiphotrocha comprimata (Cushman and Brönnimann) D Trochammina comprimata Cushman and Brönnimann, 1948
E. Alve, J.W. Murray / Palaeogeography, Palaeoclimatology, Palaeoecology 146 (1999) 171–193
Trochammina inflata (Montagu) D Nautilus inflatus Montagu,
1808
Ammonia beccarii (Linné) D Nautilus beccarii Linné, 1758
Buliminella elegantissima (d’Orbigny) D Bulimina elegantissima
d’Orbigny, 1839
Cibicides lobatulus (Walker and Jacob) D Nautilus lobatulus
Walker and Jacob, 1798
Cornuspira involvens (Reuss) D Operculina involvens Reuss,
1850
Elphidium albiumbilicatum (Weiss) D Nonion pauciloculum
Cushman, subsp. albiumbilicatum Weiss, 1954
Elphidium excavatum (Terquem) D Polystomella excavata Terquem, 1875
Elphidium macellum (Fichtel and Moll) D Nautilus macellus
Fichtel and Moll, 1798
Elphidium margaritaceum Cushman, 1930
Elphidium oceanensis (d’Orbigny) D Polystomella oceanensis
d’Orbigny, 1826
Elphidium williamsoni Haynes, 1973
Eoeponidella pulchella (Parker) D Pninaella? pulchella Parker,
1952
Fissurina marginata (Montagu) D Vermiculum marginatum
Montagu, 1803
Gavelinopsis praegeri (Heron-Allen and Earland) D Discorbina
praegeri Heron-Allen and Earland, 1913
Guttulina lactea (Walker and Jacob) D Serpula lactea Walker
and Jacob, 1798
Haynesina germanica (Ehrenberg) D Nonionina germanica
Ehrenberg, 1840
Nonion depressulus (Walker and Jacob) D Nautilus depressulus
Walker and Jacob, 1798
Ophthalmina kilianensis Rhumbler, 1936
Pateoris hauerinoides (Rhumbler) D Quinqueloculina subrotunda (Montagu) forma hauerinoides Rhumbler, 1936
Planorbulina mediterranensis d’Orbigny, 1826
Rosalina anomala Terquem, 1875
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