J. Nannoplankton Res. 29 (1), 2007, pp.5-18 © 2007 International Nannoplankton Association
ISSN 1210-8049 Printed by Cambridge University Press, UK

5Late Miocene dinoflagellate cysts and calcareous
nannoplankton from the eastern North Sea Basin margin:biostratigraphy and palaeoenvironmental interpretation(Breklum research borehole, Germany)
Angelika Köthe*, Harald Andruleit
Bundesanstalt für Geowissenschaften und Rohstoffe (BGR), Stilleweg 2, 30655 Hannover, Germany; *angelika.koethe@bgr.de
Manuscript received 13th February, 2006; revised manuscript accepted 8th November, 2006

Abstract Dinoflagellate cysts and calcareous nannoplankton were studied from sediments of the Upper Mica
Clay Member of the Breklum research well in northern Germany. The investigated section is of Late Miocene age(dinoflagellate cyst biozones DN8 and ?DN9 of Köthe, 2003). This age assignment is supported by calcareous
nannoplankton biostratigraphy (zones NN6–NN11 of Martini, 1971; Middle to Late Miocene) in the middle part ofthe section. The calcareous nannoplankton biozone assignment is based on the presence of Helicosphaera stalis,which was used as an additional marker.
As in other southern North Sea Basin countries, the diversity, preservation and number of dinoflagellate cysts are
good. Neogene records of calcareous nannoplankton in marginal North Sea Basin areas are known to be poor, but at
Breklum an almost continuous section was found, with a higher diversity than expected. The calcareous nannoplankton possibly benefited from a slightly stronger marine influence at this site, compared to currently known onshore
localities. Dinoflagellate cysts generally indicate a neritic environment, at least occasionally swept by oceanic
waters.

Keywords Late Miocene, biostratigraphy, North Sea Basin, dinoflagellate cysts, calcareous nannoplankton
1. Introduction
Studies on Late Miocene dinoflagellate cysts, and on
calcareous nannoplankton in particular, of the marginal
North Sea Basin are sparse. The scarcity of calcareous
nannoplankton in often non-calcareous sediments
increases from the Early to the Late Miocene and,
therefore, discourages investigation of this group.
However, calcareous nannoplankton are one of the mostimportant microfossil groups for worldwide correlation
and, therefore, any dating they provide for Neogene
sediments from the marginal North Sea is valuable. A
combined study of both groups enables results andinterpretations to be validated. The aim of this study wasto improve the understanding of Late Miocene
biostratigraphy by correlating dinoflagellate cysts andcalcareous nannoplankton, and evaluating theirpalaeoenvironmental potential.
2. Material and methods
The Breklum borehole is located in northern Germany on
the west coast of Schleswig-Holstein (TK25 1319
Bredstedt, Re: 3499968, H: 6053953; Figure 1). The
section comprises two boreholes drilled at the same
location by the Landesamt für Natur und Umwelt(LANU) of Schleswig-Holstein. The Breklum 2/96
borehole was drilled dry with an auger bit in 1996 to a
total depth of 40m. In 2001, the Breklum 2001 borehole
was flushed down to 40m and cored from 40–112m. As
the boreholes lie in the vicinity of the terminal moraine of
the Saalian Glacial Stage, Pleistocene faulting anddisturbance of the sediments cannot be excluded. The
boreholes (see Figure 2) were lithologically described andsampled by K. Gürs (pers. comm., 2004). The sediments
consist of micaceous clays and silts of the ‘Oberer
Glimmerton’ (Upper Mica Clay Member). Fourty-two
samples (13 ditch-cuttings and 29 core samples) were
investigated. In general, the samples were taken every
three metres. More closely spaced samples were taken insome cases when the lithology changed. The results from
Breklum were compared with previously investigatedsections (Figure 3).

Figure 1: Location of the Breklum borehole

2.1 Dinoflagellate cysts
All samples were processed using standard palynological
techniques (HCl, HF, KOH, ultrasonic sieving using a
10µm nylon screen) and mounted in glycerine jelly. All
slides were examined using a Leitz (Aristoplan) lightmicroscope with interference contrast at 250x and 400x
magnification. Two hundred dinoflagellate cyst

6

Köthe, Andruleit

can be found.
2.2 Calcareous
nannoplankton
Sample preparation followed
standard settling techniques(Hay, 1977; Bown & Young,1998). To avoid dissolution
during preparation, use wasmade of demineralised water
saturated with calcium
carbonate and buffered withammonia (Andruleit et al.,2000). All slides were
examined with a Zeiss
Axioplan 2 light-microscopeat 1000x magnification. All
slides are stored in the BGR
calcareous nannoplankton
collection. Total abundancesof the calcareous nannoplankton were defined as
specimens per field of view
(Table 2). Relative abundances were grouped intoclasses (Table 2). The
standard Neogene calcareous
nannoplankton biozonation,
as described by Martini(1971: NN zones), was used
as a basis for this study(Figure 4). Additionally, thebiostratigraphy and taxonomy presented by Young(1998) was used. Fullcitations of the identified
species can be found in
Bown (1998).
3. Palaeogeographyof the study area
Figure 2: Gamma-ray log, lithology and biostratigraphy of the Breklum borehole
After extensive transgression
specimens per sample were counted. The total in the Middle Miocene, the Late Miocene indicates aabundances of dinoflagellate cysts were defined as period of regression (Gramann & Kockel, 1988, p.439;
specimens per traverse (0.228cm2) of the slide, and Balson, 1999; Huuse, 2002). The sediments reflect
relative abundances were grouped into four classes (Table deposition during long-term climatic cooling and eustatic1).
sea-level fall (Huuse, 2002). The extent of the southern
All slides are stored in the dinoflagellate cyst North Sea was similar to that of today, but it is unlikely
collection of the Federal Institute for Geosciences and that there was a connection to the Bay of Biscay via the
NaturalResources,
Hannover(BGR).
The English Channel (Figure 3; Balson, 1999). Therefore, thebiostratigraphic zonation (DN zones) follows Köthe southern North Sea was probably a semi-enclosed(2003; see Figure 4). The biozonation proposed by embayment. The main depocentre lay along the axis of
Munsterman & Brinkhuis (2004) could not be applied, the subsiding Rhine Graben (Balson, 1999). Progressive
because the chronological order of their index-markers filling of the eastern and central North Sea Basin by large
contradicts observations from Germany (Köthe, 2000; deltas continued. During the Late Miocene, rivers were
Strauss et al., 2001). The taxonomy follows Williams et fed from the Baltic region. The Baltic River Systemal. (1998), where full citations of the identified species formed a westward prograding delta, which deposited

Late Miocene dinoflagellates and nannofossils...Germany

77), the interval from 111.45m to
22.50m is of early to middle Late
Miocene age (zone DN8). Because
specimens of Palaeocystodinium
are very rare, identification of
DN9 (middle to late Late
Miocene), from 19.50m to
10.50m, remains questionable.
The assemblages are dominated by Achomosphaera andSpiniferites. Almost continuously
present
andcommonare
Lingulodinium machaerophorum,
Nemato-sphaeropsis?
crassimuratus? and Operculodinium israelianum (Plate 2).
Spiniferites pseudofurcatus, Hystrichokolpoma salacia and Homotryblium tenuispinosum (Plate 3)
are not continuously present but
are very abundant in somesamples. The genera Impagidinium, Selenopemphix and
Sumatradinium are only sporadically present and occur in low
Figure 3: Palaeogeography of the marginal North Sea Basin in the late Middle Miocene (after Popovnumbers (Plate 4).
et al., 2004). Known locations analysed for dinoflagellate cysts and calcareous nannoplankton,
Dybkjær (2004) notes that, incovering the same time-interval as the Breklum research well, are depicted. The palaeogeographic
map and the investigated sections differ slightly in age
north-western Europe, the last
occurrence of H. tenuispinosum ishuge amounts of sediment in northern Germany (Huuse,distinctly younger than the 23.67Ma (around the2002). In the Lower Rhine Embayment, isochronous
Oligocene/Miocene boundary) suggested by Hardenbol etcoastal-deltaic and continental-fluvial beds compriseal. (1998, chart 3). The presence of H. tenuispinosum insands, clays and several thick lignite horizons
the Late Miocene in Breklum supports Dybkjær (2004).
(Herngreen, 1987) of economic importance. Apart from
Reworked specimens of Paleogene and Lower
seismic surveys, most publications covering the Late
Miocene strata are regularly distributed in the sequence,
Miocene in the southern North Sea Basin concern onshore
but they do not exceed 2% of the total dinoflagellate cyst
locations. The sediments consist ofnear-shore,
shallowmarinesediments, swept only occasionallyby oceanic waters.
4. Results
Taxa used for biozonation are shown
on Figure 2.
4.1 Dinoflagellate cysts
All samples contain common toabundant dinoflagellate cyst assemblages, except for the uppermost fivesamples that yield fewer specimens.
The two uppermost samples arealmost barren. Preservation is good.
The identified dinoflagellate
cysts are shown in Table 1. Based on
the co-occurrence of Achomosphaera
andalousiensis (Pl.1, figs 1–3) and
Palaeocystodinium spp. (Pl.1, figs 6,

Figure 4: Correlation between dinoflagellate cysts and calcareous nannoplankton zones (modified
after Köthe, 2005) and their markers in the Late Miocene. The time-scale follows Gradstein et al.
(2004)

8

Köthe, Andruleit

4.50*
8.50*
10.50*
13.50*
16.50*
19.50*
22.50*
25.50*
28.50*
31.50*
34.50*
37.50*
39.50*
41.28
42.50
45.50
48.20
48.50
51.50
54.50
60.50
63.50
66.40
66.50
70.00
72.50
75.50
78.50
82.35
83.35
86.50
89.50
93.50
95.10
95.40
97.75
100.65102.20105.55108.25109.55111.45

Achilleodinium biformoides
Achomosphaera alcicornu
Achomosphaera andalousiensis
Achomosphaera ramulifera
Achomosphaera spp.
Achomosphaera-Spiniferites 'complex'
Barssidinium pliocaenicum
Barssidinium-Sumatradinium 'complex'
Dapsilidinium pseudocolligerum
Filisphaera filifera
Habibacysta tectata
Homotryblium caliculum
Homotryblium plectilum
Homotryblium tenuispinosum
Homotryblium spp.
Hystrichokolpoma rigaudiae
Hystrichokolpoma salacia
Hystrichosphaeropsis obscura
Hystrichostrogylon membraniphorum
Impagidinium patulum
Impagidinium patulum 'complex'
Impagidinium spp.
Impletosphaeridium 'complex'
Invertocysta lacrymosa
Invertocysta tabulata
Labyrinthodinium truncatum modicum
Labyrinthodinium truncatum truncatum
Lejeunecysta sp.
Lingulodinium machaerophorum
Melitasphaeridium asterium
Melitasphaeridium choanophorum
Melitasphaeridium pseudorecurvatum
Nematosphaeropsis crassimuratus
Nematosphaeropsis lemniscata
Nematosphaeropsis ? crassimuratus ?
Operculodinium centrocarpum
Operculodinium crassum
Operculodinium israelianum
Operculodinium janduchenei
Operculodinium longispinigerum
Operculodinium piaseckii

Abundance†

Biozone*ditch-cuttings

Depth (m)autochthonous dinoflagellate cysts

--

DN9?

?

?
?

DN8

?
?
?

?

?
?

†specimens per traverse

?

>100

>40<100>15<40<15

Table 1: Range-chart of dinoflagellate cysts, Breklum boreholecounts (Pl.5, figs 7, 9). The presence of the fresh-wateralga Pediastrum kawraiskyi, at 4.50m and 8.50m, might
indicate a Pleistocene age, as this alga is only known fromthis period in Germany. The presence of P. kawraiskyi inthe core samples at 41.28m, 48.20m and, questionably, at
60.50m might indicate caving or Pleistocene faulting anddisturbance of the sediments. No other observations
support this interpretation.
4.2 Calcareous nannoplankton
The majority of samples yield at least sparse nannofloras,
but 15 were barren, and none of the samples contained
very abundant calcareous nannoplankton. The
autochthonous nannoplankton (Plate 6) generally show
species richness of a maximum of 12 species (Sample
66.50m), but usually no more than five species are found.
The most ubiquitous species is Coccolithus pelagicus,
followed by species of the genus Reticulofenestra. Other

species are generally rare and occur only sporadically.
Most
samples
containreworkedcalcareous
nannoplankton, mainly of Late Cretaceous age (Plate 7).
Preservation is generally moderate to poor, especiallywhen abundances are low.
The investigated interval can be subdivided using
nannoplankton (Table 2). The lowermost section, from111.45m to 86.50m, is no older than Early Miocene (zoneNN4), due to the presence of Reticulofenestra
pseudoumbilicus. The occurrence of Helicosphaera stalis
indicates a Middle to Late Miocene age (NN6 to NN11)
for the section from 83.35m to 66.50m. From core-depths
66.40m to 39.50m, the section is no younger than
Pliocene (NN15), due to the presence of R.
pseudoumbilicus. Further age determination using
nannoplankton cannot be given for the uppermost part ofthe section.

Late Miocene dinoflagellates and nannofossils...Germany

9

Operculodinium tegillatum ?
Operculodinium spp.
Palaeocystodinium golzowense
Palaeocystodinium miocaenicum ?
Palaeocystodinium powellense
Pentadinium laticinctum
Pentadinium taeniagerum
Reticulatosphaera actinocoronata
Selenopemphix brevispinosa
Spiniferella cornuta
Spiniferites gracilimembranaceus
Spiniferites nodosus
Spiniferites pseudofurcatus
Spiniferites pseudofurcatus (granulate)
Spiniferites spp.
Sumatradinium druggii ?
Sumatradinium pliocenicum
Surculosphaeridium alagoense
Tectatodinium pellitum
Tectatodinium psilatum
Tuberculodinium vancampoae

Achomosphaera crassipellis
Areoligera senonensis
Areosphaeridium diktyoplokus
Charlesdowniea clathrata
Charlesdowniea variabilisCharlesdowniea sp.
Chiropteridium lobospinosum
Cordosphaeridium exilimurum
Cordosphaeridium funiculatum
Cordosphaeridium gracile
Cordosphaeridium inodes
Distatodinium craterum
Enneadocysta pectiniformis
Exochosphaeridium insigne
Glaphyrocysta ordinata
Homotryblium sp. aff. H. abbreviatum
Homotryblium oceanicum
Polysphaeridium zoharyi
Spiniferites andalousiensis
Systematophora placacantha
Thalassiphora pelagica
Tityrosphaeridium cantharellum
Wetzeliella articulata
Wetzeliella symmetrica
Wetzeliella spp.
Pediastrum kawraiskyi
fresh-water algae
Pediastrum spp.

reworked dinoflagellate cysts

autochthonous dinoflagellate cysts

?

?

?

?
≥ 50%
≥ 10% < 50%≥ 2% < 10%> 0% < 2%?

determination questionable

?

?
?

?
?

?

?

Table 1 cont’d: Range-chart of dinoflagellate cysts, Breklum borehole

5. Discussion
5.1 Comparison with previous
investigations
Few published data from the southern North Sea Basinconcern Late Miocene dinoflagellate cysts, and evenfewer deal with calcareous nannoplankton (Figure 3). In
Denmark, dinoflagellate cysts of the isochronous Gram
Formation were studied by Piasecki (1980). According to
the first occurrence of Achomosphaera andalousiensis,
the base of the Gram Formation can be dated as DN8 andthe upper part of the Gram Formation as DN9, based on
the last occurrence of Palaeocystodinium golzowense(sensu Köthe, 2003). The Gram Formation yields richdinoflagellate cyst assemblages, dominated by
Homotryblium cf. H. plectilum and Gramocysta verricula.
Neither species are encountered at Breklum. Although
Breklum lies close to the Danish border (Figure 3), thedinoflagellate cyst assemblages differ considerably.

In Germany, the boreholes at Wursterheide (Cepek,
1989; Lund & Heilmann-Clausen, 1999), Nieder
Ochtenhausen (Martini, 2001; Strauss et al., 2001) and
Groß Pampau (Lund & Lund-Christensen in Daniels etal., 1990; Martini, 2001) were investigated, looking at
both fossil groups. Their positions are shown on Figure 3.
Zones DN8 and DN9 are present in the Nieder
Ochtenhausen borehole and DN9 in the Wursterheide
borehole (Köthe, 2005). In Nieder Ochtenhausen, thedinoflagellate cysts Lingulodinium machaerophorum andthe genus Spiniferites are abundant, although few
specimens of Gramocysta verricula were identified. Zone
DN8 (sensu Köthe, 2003) is also present in the Groß
Pampau clay pit, dominated by G. verricula (lower
sample only) and Spiniferites (Lund & Lund-Christensen
in Daniels et al., 1990). Dating by calcareous
nannoplankton was not possible in these three localities
due to the scarcity of this group. The Upper Pritzier Beds

Köthe, Andruleit

10

NN11
NN6

4.50*
8.50*
10.50*
13.50*
16.50*
19.50*?
22.50*
25.50*
28.50*
31.50*
34.50*
37.50*
39.50*
41.28
42.50
45.50
not
48.20
younger
than
NN15

48.50
51.50
54.50
60.50

63.50
66.40
66.50

70.00
72.50

75.50
78.50

82.35
83.35

86.50
89.50
93.50

95.10

95.40
97.75 not older
100.65 than NN4102.20105.55108.25109.55111.45*ditch-cuttings

BiozoneAbundance†

barrenbarrenbarrenbarren≥ 50%

barrenbarren≥ 10% < 50%≥ 2% < 10%> 0% < 2%<1 specimen per 20 fov

Ahmuellerella octoradiata
Arkhangelskiella cymbiformis
Biscutum constans
Biscutum cf. B. constans
Biscutum sp.
Broinsonia parca parca
Calculites obscurus
Chiastozygus trabalis
Cribrosphaerella ehrenbergii
Cyclagelosphaera margerelii
Eiffellithus eximius
Eiffellithus gorkae
Eiffellithus turriseiffelii
Eiffellithus sp.
Eprolithus floralis
Eprolithus sp.
Gartnerago segmentatum
Helicolithus anceps
Kamptnerius magnificus
Lithastrinus grillii
Lucianorhabdus cayeuxii
Manivitella pemmatoidea
Marthasterites furcatus
Micula staurophora
Micula spp.
Prediscosphaera cretacea
Prediscosphaera spinosa
Prediscosphaera sp.
Quadrum gartneri
Quadrum sp.
Reinhardtites anthophorus
Reinhardtites levis
Reinhardtites sp.
Retecapsa surirella
Retecapsa sp.
Staurolithites sp.
Tranolithus orionatus
Tranolithus sp.
Watznaueria barnesiae
Watznaueria fossacincta
Zeugrhabdotus sp.

reworked calcareous nannoplankton= 1-19 specimens per 20 fovbarren= 1-10 specimens

Table 2: Range-chart of calcareous nannoplankton, Breklum borehole

Braarudosphaera bigelowii
Calcidiscus macintyrei
Coccolithus pelagicus
Discoaster sp. cf. D. challengeri
Helicosphaera carteri
Helicosphaera kamptneri
Helicosphaera stalis
Helicosphaera sp.
Pontosphaera discopora
Pontosphaera multipora
Pontosphaera sp.
Reticulofenestra <3µm
Reticulofenestra <7µm
Reticulofenestra pseudoumbilicus
Sphenolithus abies
Sphenolithus moriformis
Syracosphaera sp.

autochthonous calc. nannoplankton†specimens per field of view (fov)Depth (m)Late Miocene dinoflagellates and nannofossils...Germany

11

Plate 1
Figs 1–8: dinoflagellate cyst index and secondary markers
Figs 9–15: dinoflagellate species of the genus Operculodinium

Achomosphaera andalousiensis
66.50m, DN8

Achomosphaera andalousiensis
34.50m, DN9?

Achomosphaera andalousiensis
89.50m, DN8

Habibacysta tectata
13.50m, DN9?

Habibacysta tectata
42.50m, DN9?

Palaeocystodinium
powellense
70.00m, DN8

Operculo. longispinigerum
102.20m, DN8Palaeocystodinium
golzowense
95.40m, DN8
Operculodinium
longispinigerum 89.50m, DN8

Habibacysta tectata
37.50m, DN9?

Operculodinium tegillatum?
60.50m, DN8

50µm

Operculodinium piaseckii 48.20m, DN9?

Operculodinium crassum 39.50m, DN9?

Operculodinium piaseckii 66.40m, DN8

in the Lübtheen 27/82 borehole (Figure 3) yield richdinoflagellate cyst assemblages of zones DN8 and DN9,
which are dominated by species of the AchomosphaeraSpiniferites complex, Habibacysta tectata, Hystrichokolpoma rigaudiae and L. machaerophorum (Köthe,2000).

Operculodinium centrocarpum
108.25m, DN8

In The Netherlands, two boreholes close to theGerman border yield Late Miocene dinoflagellate cysts(Munsterman & Brinkhuis, 2004), but a range-chart is notprovided. The determination of a Late Miocene age in the
Cuyk and Broeksittard boreholes (Herngreen, 1987) is notproven, according to the zonation of Köthe (2003).

Köthe, Andruleit

12

Plate 2
Dinoflagellate cysts which are almost continuously present throughout the borehole

Nematosphaeropsis? crassimuratus?
10.50m, DN9?
Nematosphaeropsis? crassimuratus?
34.50m, DN9?

Hystrichokolpoma rigaudiae
28.50m, DN9?

Nematosphaeropsis? crassimuratus?
22.50m, DN9?

50µm

Tuberculodinium vancampoae
102.20m, DN8

Lingulodinium machaerophorum
111.45m, DN8

In Belgium, the Bolderberg Formation is dated as
DN8 (Louwye et al., 1999) and the Diest Formation as top
DN9–DN10 (Louwye & Laga, 1998), that is, together
comprising DN8, DN9 and DN10 (Louwye et al., 1999).
The Deurne Sands of the Diest Formation (DN8; Louwye,2002) contain a diverse and well-preserved dinoflagellate
cyst assemblage. Selenopemphix brevispinosa and S.
armageddonensis are limited to the Diest Formation. A.
andalousiensis and Amiculosphaera umbracula arepresent, whereas P. golzowense and G. verricula are
missing. At Breklum, S. brevispinosa is restricted to thelower part of the studied section (DN8, 83.35m–95.10m).
G. verricula, the index marker of the Middle to Late
Miocene G. verricula Zone (Piasecki, 1980; Powell,1992) was not recorded in Breklum. Both resultscorrespond to the findings from the Diest Formation in
Belgium.
Miocene sediments are not present onshore GreatBritain, except for the Lenham Beds, which may be of
Late Miocene age (Balson, 1999, p.240).
The dinoflagellate cyst assemblages at Breklum show
affinities to those from other German and Belgian
localities. All these are dominated by Spiniferites andwere deposited in a shallow marine environment, whereas
the Danish localities indicate a shallower, restrictedmarine environment, due to the dominance of H. cf. H.

Spiniferites
gracilimembranaceus
72.50m, DN8

Reticulatosphaera
actinocoronata
34.50m, DN9?

plectilum.
Based on dinoflagellate cysts, the studied section at
Breklum is of Late Miocene age (DN8, ?DN9). This is inagreement with the biostratigraphy based on calcareous
nannoplankton (not older than NN4, NN6–NN11, notyounger than NN15), as DN8 is correlated internationallywith NN9 and NN10, and DN9 with the lower part of
NN11 (De Verteuil & Norris, 1996). The presence ofHelicosphaera stalis in particular, which was notdescribed until 1984, can be used as an additional marker.
As in other southern North Sea countries, the number ofdinoflagellate cysts in Breklum is high and thepreservation good, whereas the calcareous nannoplankton
were obviously affected by dissolution. Calcareous
nannoplankton are generally very rare in Neogene
sediments of North Germany and the North Sea Basin
margin, and subdivision by means of calcareous
nannoplankton was not thought to be possible (Köthe,1988). Considering the entire North Sea Basin, theNeogene assemblages appear to be poor, basin-wide, andage determinations are generally vague (Gallagher, 1990).
Sections younger than Middle Miocene are often barrenof autochthonous calcareous nannoplankton or
characterised by rare occurrences of long-ranging, andtherefore stratigraphically insignificant, taxa (Cepek,1989). Some Early and Middle Miocene north-westernLate Miocene dinoflagellates and nannofossils...Germany

13

Plate 3
Dinoflagellate cysts that are abundant in some samples

Homotryblium tenuispinosum 25.50m, DN9?

Hystrichokolpoma salacia 28.50m, DN9?
Spiniferites pseudofurcatus 75.50m, DN8European local stages were dated by nannoplankton
biostratigraphy (Martini & Müller, 1973; Müller et al.,1979), but nannoplankton biozonations are not available
for the local Late Miocene stages.
5.2 Environmental conditions during
deposition
The results herein do not contradict the general
impression of an unfavourable environment forcalcareous nannoplankton in the North Sea Basin during
the Late Miocene, but show that an almost continuous
section, with a higher diversity than expected, can befound (Müller et al., 1979). The calcareous
nannoplankton may possibly have benefited from
temporarily more offshore conditions at this site. Martini(2001) showed that abundances of autochthonous
calcareous nannoplankton assemblages from the southwestern North Sea area might be positively correlated to
Neogene sea-level high-stands. However, the distorting
influence of carbonate dissolution has to be consideredwhen interpreting the autochthonous calcareous
nannoplankton, especially since a number of samples are
barren of autochthonous, as well as reworked, calcareous
nannoplankton.
The dominance of the dinoflagellate genera
Operculodinium, Homotryblium and Spiniferites indicate
neritic conditions (Pross et al., 2004). The genera
Impagidinium and Nematosphaeropsis, which indicate,even in very small numbers, the influence of oceanic
waters (Dale, 1996), occur only sporadically at Breklum.
Impagidinium is notably present between 72.50m and111.45m, and Nematosphaeropsis (apart from
Nematosphaeropsis? crassimuratus?) between 83.35m
and 42.50m. There is no correlation between the presenceof calcareous nannoplankton and the presence of oceanic
dinoflagellate cysts.
The interpretation of the depositional environment of

50µm

the dinoflagellate genus Homotryblium has recently been
demonstrated by Dybkjær (2004). According to her
investigations, Homotryblium tenuispinosum is dominantand continuously present in the distal parts of thehighstand systems tract (HST) of a brackish sequence. H.
tenuispinosum is not continuously present in Breklum, but
most abundant in Samples 45.50m (54%) and 25.50m(90%). The presence of autochthonous calcareous
nannoplankton in Sample 45.50m also undermines theinterpretation of a brackish environment, whereas in
Sample 25.50m, calcareous nannoplankton are notpresent. Therefore, the interpretation of a brackish
environment during the deposition of these two samples
remains doubtful. The genus Homotryblium is very scarcein Wursterheide, Lübtheen and in Groß Pampau. In
Nieder Ochtenhausen, it is common to abundant in thelower part of DN8.
In conclusion, the calcareous nannoplankton possibly
benefited from a slightly stronger marine influence at thissite, compared to currently known onshore localities.
Dinoflagellate cysts generally indicate a neritic
environment, at least occasionally swept by oceanic
waters.
Acknowledgements
We are grateful to Dr. Karl Gürs (LANU) who kindly provided
the samples and the log data. The technical support by Angelika
Malik, Barbara Piesker and Sabine Stäger (BGR) is thankfully
acknowledged. Drs Liam Gallagher, Keith Gueinn (Network
Stratigraphic Consulting Limited, UK), and Emma Sheldon andStefan Piasecki (Geological Survey of Denmark and Greenland,
Denmark), improved the manuscript by constructive reviews.

References

Andruleit, H., Köthe, A., Stäger, S. & Bruns, A. 2000. Some
comments concerning sample processing and dissolution.
Journal of Nannoplankton Research, 22(3): 201.
Balson, P. 1999. The Neogene of eastern England. In: B. Daley

Köthe, Andruleit

14

Plate 4
Dinoflagellate cysts that are rarely present

Nematosphaeropsis
crassimuratus
42.50m, DN9?

Pentadinium taeniagerum 108.25m, DN8

Spiniferella cornuta 34.50m, DN9?

Labyrinthodinium truncatum
modicum 111.45m, DN8

Spiniferites nodosus 28.50m, DN9?

Imp. patulum ‘complex’
100.65m, DN8

50µm

Nematosphaeropsis crassimuratus
42.50m, DN9?
Impagidinium patulum
72.50m, DN8

Sumatradinium pliocenicum 108.25m, DN8Invertocysta lacrymosa 39.50m, DN9?

& P. Balson (Eds). British Tertiary Stratigraphy. GeologicalConservation Review Series, 15: 235–241.
Bown, P.R. (Ed.) 1998. Calcareous Nannofossil
Biostratigraphy. British Micropalaeontological Society
Series. Chapman & Hall/Kluwer Academic Press, London:
315pp.
Bown, P.R. & Young, J. 1998. Techniques. In: P.R. Bown (Ed.).
Calcareous
Nannofossil
Biostratigraphy.
BritishMicropalaeontological Society Series. Chapman &Hall/Kluwer Academic Press, London: 16–28.
Cepek, P. 1989. Das kalzitische Nannoplankton des Tertiär undder Kreide aus der Forschungsbohrung Wursterheide (NWDeutschland). Geologisches Jahrbuch, A111: 403–429.
Dale, B. 1996. Dinoflagellate cyst ecology: modeling andgeological applications. In: J. Jansonius & D.C. McGregor(Eds). Palynology: principles and applications. American

Sel. brevispinosa
95.10m, DN8

Selenopemphix brevispinosa
86.50m, DN8

Nematosphaeropsis lemniscata 48.20m, DN9?

Association of Stratigraphic Palynologists Foundation, 3:
1249–1275.
von Daniels, C.H., Lund, J., Lund-Christensen, J.J. &Uffenorde, H. 1990. The Langenfelde (Miocene) of GroßPampau, Schleswig-Holstein. Foraminifer, dinocyst andostracod stratigraphy and paleoecology (preliminaryaccount). Veröffentlichungen Übersee-Museum Bremen,
A10: 11–38.
De Verteuil, L. & Norris, G. 1996. Dinoflagellate cyst zonation
and allostratigraphy of the Chesapeake Group. In: L. De
Verteuil & G. Norris (Eds). Miocene dinoflagellate
stratigraphy and systematics of Maryland and Virginia.
Micropaleontology, 42, suppl., part 1: 1–82.
Dybkjær, K. 2004. Morphological and abundance variations in
Homotryblium-cyst assemblages related to depositional
environments; uppermost Oligocene–Lower Miocene,Late Miocene dinoflagellates and nannofossils...Germany

15

Plate 5
Figs 1-6, 8, 10, 11: autochthonous dinoflagellate cysts
Figs 7, 9: reworked Paleogene dinoflagellate cysts

Hystrichosphaeropsis obscura
66.50m, DN8Hystrichosphaeropsis obscura 86.50m, DN8
Spiniferites pseudofurcatus ‘granulate’ 16.50m, DN9?

50µm

Melitasphaeridium
choanophorum 60.50m, DN8Melitasphaeridium
choanophorum 25.50m, DN9?
Spiniferites pseudofurcatus ‘granulate’ 19.50m, DN9?

Labyrinthodinium
truncatum truncatum
42.50m, DN9?

Achomosphaera crassipellis 109.55m, DN8

Homotryblium sp. aff. H.
abbreviatum 25.50m, DN9?

Labyrinthodinium truncatum Filisphaera filifera 34.50m, DN9?
truncatum 93.50m, DN8

Köthe, Andruleit

16

Plate 6
Autochthonous calcareous nannoplankton
BF = bright-field, PC = phase-contrast, XPL = cross-polarised light

Helicosphaera carteri 45.50m (XPL/BF)

Braarudosphaera bigelowii
66.40m (XPL/BF)

Coccolithus pelagicus 34.50m (XPL/BF)

Reticulofenestra <3µm86.50m (XPL/BF)

Helicosphaera stalis
66.50m (XPL/BF)

Calcidiscus macintyrei 66.50m (XPL/BF)

10µm

Sphenolithus abies 66.50m (XPL/XPL/BF)

Sphenolithus moriformis
66.50m (XPL/XPL)

Jylland, Denmark. Palaeogeography, Palaeoclimatology,
Palaeocology, 206: 41–58.
Gallagher, L.T. 1990. Calcareous nannofossil Biozonation of the
Tertiary of the North Sea Basin. Newsletters on
Stratigraphy, 22(1): 21–41.
Gradstein, F., Ogg, J. & Smith, A. (Eds) 2004. A Geologic Time
Scale 2004. Cambridge University Press, Cambridge:
589pp.
Gramann, F. & Kockel, F. 1988. Palaeogeographical,
lithological, palaeoecological and palaeoclimaticdevelopment of the Northwest European Tertiary Basin. In:
R. Vinken (Ed.). The Northwest European Tertiary Basin.
Results of the International Geological Correlation
Programme Project No. 124. Geologisches Jahrbuch, A100:
428–441.
Hardenbol, J., Thierry, J., Farley, M.B., De Graciansky, P.-C. &Vail, P.R. 1998. Mesozoic and Cenozoic sequencechronostratigraphic framework of European basins. In: P.C.
DeGraciansky, J. Hardenbol, T. Jacquin & P.R. Vail (Eds).
Mesozoic and Cenozoic sequence stratigraphy of European
basins. Society of Economic Paleontologists andMineralogists (Society for Sedimentary Geology), Special
Publication, 60: 3–13.
Hay, W.W. 1977. Calcareous nannofossils. In: A.T.S. Ramsay(Ed.). Oceanic Micropaleontology. Academic Press,

Reticulofenestra <7µm34.50m (XPL/PC)

Reticulofenestra pseudoumbilicus 66.50m (XPL/BF)

London: 1055–1200.
Herngreen, G.F.W. 1987. Correlation between Miocene beds of
the SE Netherlands and Italy based on dinoflagellatebiozonation. Mededelingen van de Werkgroep voor Tertiaire
en Kwartaire Geologie, 24(1-2): 31–40.
Huuse, M. 2002. Late Cenozoic palaeogeography of the eastern
North Sea Basin: climatic vs. tectonic forcing of basin
margin uplift and deltaic progradation. Bulletin of theGeological Society of Denmark, 49: 145–170.
Köthe, A. 1988. Nannoplankton. In: R. Vinken (Ed.). The
Northwest European Tertiary Basin. Results of theInternational Geological Correlation Programme ProjectNo. 124. Geologisches Jahrbuch, A100: 253–288.
Köthe, A. 2000. Dinozysten der Pritzierer Schichten in derBohrung Lübtheen 27/82. In: W. von Bülow (Ed.).
Geologische Entwicklung Südwest Mecklenburgs seit dem
Ober-Oligozän. Schriftenreihe für Geowissenschaften, 11:
157–179.
Köthe, A. 2003. Dinozysten-Zonierung im Tertiär
Norddeutschlands. Revue de Paléobiologie, 22(2): 895–923.
Köthe, A. 2005. Korrelation der Dinozysten-Zonen mit anderen
biostratigraphisch wichtigen Zonierungen im Tertiär
Norddeutschlands. Revue de Paléobiologie, 24(2): 697–718.
Louwye, S. 2002. Dinoflagellate cyst biostratigraphy of theUpper Miocene Deurne Sands (Diest Formation) of northernLate Miocene dinoflagellates and nannofossils...Germany

17

Plate 7
Reworked calcareous nannoplankton
BF = bright-field, PC = phase-contrast, XPL = cross-polarised light

Tranolithus orionatus 86.50m (XPL/BF)

Watznaueria barnesiae 86.50m (XPL/BF)

Helicolithus anceps 19.50m (XPL/PC).

Micula staurophora 19.50m (XPL/PC)

Cyclagelosphaera margerelii
60.50m (XPL/BF)

Calculites obscurus 63.50m (XPL/BF)

Eiffellithus gorkae
86.50m (XPL/BF)

Lucianorhabdus cayeuxii 34.50m (XPL/BF)

Ahmuellerella octoradiata 45.50m (XPL/PC)

10µm

Arkhangelskiella cymbiformis 100.65m (XPL/BF)

Kamptnerius magnificus 66.50m (XPL/BF)

Belgium, southern North Sea Basin. Geological Journal, 37:
55–67.
Louwye, S., De Coninck, J. & Verniers, J. 1999. Dinoflagellate
cyst stratigraphy and depositional history of Miocene andLower Pliocene formations in northern Belgium (southern
North Sea Basin). Geologie en Mijnbouw, 78: 31–46.
Louwye, S. & Laga, P. 1998. Dinoflagellate cysts of the shallow
marine Neogene succession in the Kalmthout well, northern
Belgium. Bulletin of the Geological Society of Denmark, 45:
73–86.
Lund, J.J. & Heilmann-Clausen, C. 1999. Dinoflagellate
correlation between the research borehole Nieder
Ochtenhausen and Wursterheide, Miocene, NW Germany.
Aardkundige Mededelingen, 11: 51–60.
Martini, E. 1971. Standard Tertiary and Quaternary Calcareous
Nannoplankton Zonation. Proceedings of the 2nd
Planktonic Conference, Roma, 1970(2): 739–785.

Martini, E. 2001. Kalkiges Nannoplankton aus dem Neogen derForschungsbohrung Nieder Ochtenhausen sowie Groß
Pampau und Hohen Woos (Norddeutschland). Geologisches
Jahrbuch, A152: 357–377.
Martini, E. & Müller, C. 1973. Nannoplankton-Gemeinschaften
im Miozän und Pliozän des Nordseebeckens. Neues
Jahrbuch für Geologie und Paläontologie Mitteilungen, 9:555–564.
Müller, C., Kuster, H., Spiegler, D., von Daniels, C.H. 1979.
Nachweis von Nannoplankton-Zonen und ihre Korrelation
mit lokalen Stufen im Neogen NW-Deutschlands.
Newsletters on Stratigraphy, 8(2): 153–158.
Munsterman, D.K. & Brinkhuis, H. 2004. A southern North Sea
Miocene dinoflagellate cyst zonation. Geologie en
Mijnbouw, 83(4): 267–285.
Piasecki, S. 1980. Dinoflagellate cyst stratigraphy of the
Miocene Hodde and Gram Formations, Denmark. Bulletin

18

Köthe, Andruleitof the Geological Society of Denmark, 29: 53–76.
Popov, S.V., Rögl, F., Rozanov, A.Y., Steininger, F.F., Shcherba,
I.G. & Kovac, M. (Eds) 2004. LithologicalPalaeogeographic maps of Paratethys – 10 maps Late
Eocene to Pliocene. Courier Forschungsinstitut
Senckenberg, 250: 1–46.
Powell, A.J. (Ed.) 1992. A Stratigraphic Index of Dinoflagellate
Cysts. British Micropalaeontological Society Series,
Chapman & Hall, London: 281pp.
Pross, J., Kotthoff, U. & Zonneveld, K. 2004. Die Anwendungorganischwandiger
Dinoflagellatenzysten
zur
Rekonstruktion von Paläoumwelt, Paläoklima und
Paläozeanographie:
Möglichkeiten
undGrenzen.
Paläontologische Zeitschrift, 78(1): 5-39.
Strauss, C., Lund, J.J. & Lund-Christensen, J. 2001. Miocene
Dinoflagellate Cyst Biostratigraphy of the Nieder
Ochtenhausen Research Borehole (NW Germany).
Geologisches Jahrbuch, A152: 395–447.
Young, J.R. 1998. Neogene. In: Bown, P.R. (Ed.). Calcareous
Nannofossil Biostratigraphy. British MicropalaeontologicalSociety Series. Chapman & Hall/Kluwer Academic Press,
London: 225–265.
Williams, G.L., Lentin, J.K. & Fensome, R.A. 1998. The Lentin
and Williams Index of Fossil Dinoflagellates (1998 Edition).
American Association of Stratigraphic Palynologists
Contributions Series, 34: 1–817.

Figured dinocyst taxa
Achomosphaera andalousiensis Jan du Chene, 1977
Achomosphaera crassipellis (Deflandre & Cookson, 1955)
Stover & Evitt, 1978
Filisphaera filifera Bujak, 1984
Habibacysta tectata Head et al., 1989
Homotryblium sp. aff. H. abbreviatum Eaton, 1976
Homotryblium tenuispinosum Davey & Williams, 1966
Hystrichokolpoma rigaudiae Deflandre & Cookson, 1955
Hystrichokolpoma salacia Eaton, 1976
Hystrichosphaeropsis obscura Habib, 1972
Impagidinium patulum (Wall, 1967) Stover & Evitt, 1978
Invertocysta lacrymosa Edwards, 1984
Labyrinthodinium truncatum modicum DeVerteuil & Norris,
1996
Labyrinthodinium truncatum truncatum Piasecki, 1980
Lingulodinium machaerophorum (Deflandre & Cookson, 1955)
Wall, 1967
Melitasphaeridium choanophorum (Deflandre & Cookson,1955) Harland & Hill, 1979
Nematosphaeropsis crassimuratus Strauss et al., 2001
Nematosphaeropsis? crassimuratus? Strauss et al., 2001
Nematosphaeropsis lemniscata Bujak, 1984
Operculodinium centrocarpum (Deflandre & Cookson, 1955)
Wall, 1967
Operculodinium crassum Harland, 1979
Operculodinium longispinigerum Matsuoka, 1983
Operculodinium piaseckii Strauss & Lund, 1992
Operculodinium tegillatum? Head, 1997
Palaeocystodinium golzowense Alberti, 1961
Palaeocystodinium powellense Strauss et al., 2001
Pentadinium taeniagerum Gerlach, 1961
Reticulatosphaera actinocoronata (Benedek, 1972) Bujak &Matsuoka, 1986
Selenopemphix brevispinosa Head et al., 1989
Spiniferella cornuta (Gerlach, 1961) Stover & Hardenbol,

1994
Spiniferites nodosus (Wall, 1967) Sarjeant, 1970
Spiniferites gracilimembranaceus Strauss & Lund, 1992
Spiniferites pseudofurcatus (Klumpp, 1953) Sarjeant, 1970
Spiniferites pseudofurcatus ‘granulate’ (Klumpp, 1953)
Sarjeant, 1970
Sumatradinium pliocenicum Head, 1993
Tuberculodinium vancampoae (Rossignol, 1962) Wall, 1967