Early Miocene pollen and spores from Central Jylland, Denmark – environmental and climatic implications

Full text

Early Miocene pollen and spores from western Jylland, Denmark –
environmental and climatic implications
LINDA M. LARSSON1, VIVI VAJDA1 and ERIK S. RASMUSSEN2
Larsson, L.M., Vajda, V. & Rasmussen, E.S., 2006: Early Miocene pollen and spores from western Jylland, Denmark – en-
vironmental and climatic implications. GFF, Vol. 128 (Pt. 3, September), pp. 261–272. Stockholm. ISSN 1103-5897.
Abstract: A palynological analysis of a Lower Miocene cored section from Sønder Vium in western
Jylland, Denmark, provides new data regarding the vegetation and climate during the earliest Neogene.
Most samples yielded well-preserved palynomorphs. Terrestrial pollen and spores dominate, with lesser
proportions of dinoflagellates. A fluvial input into the marine setting is corroborated by the presence of
freshwater algae, indicating an inner-neritic setting. A level containing comparatively abundant dino-
flagellate cysts probably represents a transgressional event. The late Aquitanian age of the sequence as
suggested by previous studies is supported by the composition of the palynoflora, e.g., by the presence
of Ephedripites, Platycarya, and the relatively frequent occurrence of Engelhardtia. The pollen record is
dominated by Taxodiaceae-Cupressaceae suggesting that swamp forests dominated the onshore region,
which is consistent with previous results from central and northern Europe. Besides Taxodium, the swamp
forest also contained angiosperm taxa such as Myricaceae, Nyssa, Betula, and Alnus. Elevated or better
drained hinterland areas hosted a diverse mesophytic forest, with a ground cover of reeds, sedges and
pteridophytes. Abundant pollen taxa derived from mesophytic forests indicates the presence of evergreen
conifers, such as Pinus, Sequoia and Sciadopitys, and deciduous angiosperms, including Fagus and
Quercus. A decrease in relative abundances of thermophilous elements such as Arecaceae (palms), Ilex,
Mastixiaceae and Engelhardtia, in the middle part of the studied succession indicates a possible cor-
relation to the late Aquitanian climatic deterioration. The composition of the palynological assemblages
including widely distributed Taxodium swamps, suggests a warm, frost-free temperate climate during the
Aquitanian in Denmark.
Keywords: Sønder Vium, Miocene, Denmark, palynology, paleoclimate, Taxodiaceae, Cupressaceae,
swamp forest.
1 GeoBiosphere Science Centre, Department of Geology, Lund University, Sölvegatan 12, SE-223 62 Lund,
Sweden; linda.larsson@geol.lu.se
2 Geological Survey of Denmark and Greenland (GEUS), Øster Voldgade 10, DK-1350 Copenhagen,
Denmark
Manuscript received 27 January 2005. Revised manuscript accepted 24 August 2006.
Introduction
The Neogene has a complex history of global climate evolu-
tion and one of the most remarkable episodes was the Miocene
Climatic Optimum, which peaked 17–15 million years ago. In
Danish sediments this is reflected in the mid-Miocene Falsterholt
flora indicating warm temperate to subtropical climatic condi-
tions (Friis 1975). However, this interval was preceded by an
earliest Miocene cooling event, and global climate oscillations
are characteristic for the whole epoch (Kashiwagi & Shikazono
2003). As Antarctica became isolated, the circulation patterns
in the oceans changed to “modern” configurations and the
south circum-polar ocean current became established. This sig-
nificantly inhibited the mixing of cold polar water with warmer
tropical water, which led to the buildup of the Antarctic polar cap
(e.g., Lear et al. 2003; Roberts et al. 2003). In central Europe,
the climate changed from subtropical to cool temperate during
the transition from Late Oligocene to Miocene. During the Late
Aquitanian and Early Burdigalian a severe temperature decrease
is evident from central European sediments, which is correlated
to a glacial maximum (Miller et al. 1991; Rasmussen 2004b).
Palynology is an excellent tool for interpreting paleoclimate
variations as the worldʼs climatic zones are closely reflected
by the distribution of vegetation types. Despite comprehensive
Neogene vegetation and climate records from west-central Eu-
rope (e.g., Thomson & Pflug 1953; Mai 1965, 1995; von der
Brelie 1967; Krutzsch 1971; Ashraf & Mosbrugger 1995, 1996;
Sadowska 1997; Zetter 1998; Kolcon & Sachsenhofer 1999;
Figueiral et al. 1999; Kovar-Eder et al. 2001; Ivanov et al. 2002),
few data have been forthcoming from Scandinavia. So far, paleo-
botanical investigations of the Danish Miocene have been based
mainly upon plant macrofossils collected from lignite-bearing
GFF
volume 128 (2006), pp. 261–272. Article

Middle Miocene deposits in central Denmark (e.g., Ingwersen
1954; Koch et al. 1973; Christensen 1974, 1976; Friis 1975,
1977a, 1977b, 1978).
The extensive palynological material investigated in this study
is derived from a cored section in the Sønder Vium borehole (Fig.
1) penetrating a pro-deltaic unit. Rasmussen (2003) interpreted
the predominantly silty sediments to have been deposited during
a relatively short interval in the early part of the Miocene. This
study aims to reconstruct the vegetation, paleoenvironment and
climate for the earliest part of the Neogene.
Geological setting
Tectonic evolution
During the Cenozoic the North Sea area was developed as an
epicontinental basin with periodic connections to the Tethys Sea
towards the south and east and the North Atlantic through a nar-
row strait between present-day Norway and Scotland (Ziegler
1990). The basin was bounded towards the north and north-east
by the Fennoscandian Shield. The southern limit was located
within the area of northern Poland, Germany, and northern France
and the western border was probably located in the British Isles.
The shape of the basin was influenced by several tectonic events
associated with the Alpine Orogeny (Vejbæk & Andersen 1987;
Ziegler 1990; Rasmussen 2004a). Most distinctive was the Early
Paleocene inversion tectonism, which reshaped the basin along
older fault zones such as the Sorgenfrei-Tornquist Zone and Cof-
fee Soil Fault. Uplift of the Fennoscandian Shield took place in
the Late Eocene (Michelsen & Nielsen 1993). Mid-Oligocene
rejuvenation of older faults resulted in the formation of a topo-
graphic relief especially on the Ringkøbing-Fyn High and within
the Norwegian-Danish Basin where older salt structures were
reactivated. The relief formed prior to the Miocene strongly con-
trolled the deposition of Miocene sediments (Rasmussen 2004a).
Tectonic movements and initial subsidence of the central North
Sea area occurred during the late Early–Middle Miocene (Koch
1989; Rasmussen 2004a). This phase resulted in a minor shift
in sediment transport direction towards a more easterly source
area. In addition, thicker coal-rich deposits were concentrated
along reactivated older fault trends. The overall subsidence of
the North Sea area resulted in a major flooding of the Danish and
southern Scandinavian area in the Late Miocene. Strong tilting
of the eastern North Sea area occurred around the Pliocene to
Pleistocene transition (Japsen 1993; Rasmussen 2005).
Sedimentation
Cold-water carbonates dominated sedimentation in the basin
during the Danian (Surlyk 1997) and the area adjacent to the
Fennoscandian Shield was dominated by accumulation of bryo-
zoan banks and coral reefs. This pattern changed dramatically
in the Selandian to a predominance of fine-grained siliciclastic
sedimentation (Heilmann-Clausen et al. 1985). Paleocene and
Eocene clay-rich deposits where laid down in deep marine en-
vironments that probably covered major parts of the Fennoscan-
dian Shield as evidenced by relictual outliers in southern Finland
(Fenner 1988). The uplift of the Fennoscandian Shield during the
Late Eocene (Michelsen & Nielsen 1993; Michelsen et al. 1998)
resulted in progradation of major deltas in the Oligocene. In the
Early Miocene, three phases of delta progradation occurred,
reaching the area of modern Denmark. The progradation of
deltas occurred from the north and north-east towards the south
and south-west and consequently a northwest–southeast trend on
the shoreline. The large fetch across the North Sea resulted in
wave-dominated deltas with barrier/lagoonal complexes located
on the eastern side on the main delta lobes. Development of this
Lower Miocene succession was controlled by a combination of
sea-level changes and tectonism (Rasmussen 2004a). A major
transgression in the Middle Miocene resulted in deposition of
the clayey deposits in the North Sea area and persistence of high-
stand deposits for the remainder of the epoch. At the termination
of the Miocene a distinct progradation of deltas from the Fen-
noscandian Shield and Central Europe reached the central part
of the North Sea area. Infilling of the central part of the North
Sea area continued from the Pliocene to the Holocene and was
punctuated only by minor transgressions.
Stratigraphy of the Danish Miocene succession
The Miocene succession, the focus of this study, is characterised
by several transgressive-regressive cycles that generated five
distinct depositional sequences named B to F in ascending order
by Rasmussen (2004b) and Rasmussen & Dybkjaer (2005). An
approximate correlation of these sequences to the lithostratig-
raphy of Denmark (Rasmussen 1961; Rasmussen 2004b) is
presented here (Fig. 2).
262 Larsson et al.: Early Miocene pollen and spores from western Jylland, Denmark GFF 128 (2006)
Fig. 1. Location of the Sønder Vium borehole in Denmark, with the
structural elements mentioned in the text (mainly after Dybkjaer
2004).

The earliest Miocene transgression resulted in the deposi-
tion of marine clayey prodelta sediments of the Vejle Fjord and
Klintinghoved formations (Sorgenfrei 1958; Larsen & Dinesen
1959; Rasmussen 1961; Rasmussen 2004b) and sand-rich fluvio-
deltaic deposits of the Ribe Formation (sequence B; Sorgenfrei
1958; Rasmussen 2004b). Subsequently, a minor transgression
resulted in the accumulation of the marine Arnum Formation.
This was followed by the deposition of a wedge of sediment,
informally named the Bastrup sand (Rasmussen 2003), which
represents a second progradation of a delta complex in the Early
Miocene (sequence C). This unit was overlain by the upper part
of the marine, clay-rich Arnum Formation. The third and final
deltaic progradation occurred at the Early to Middle Miocene
transition and is represented by the coal-bearing Odderup For-
mation (sequence D; Rasmussen 1961; Koch 1989). A warming
phase in the early part of the Middle Miocene corresponded to
a major transgression and deposition of the marine Hodde and
Gram formations (sequences E and F; Rasmussen 1961; Piasecki
1980).
Material and methods
This study is based on palynomorphs extracted from the suc-
cession referred to as sequence B in the Sønder Vium bore hole
(DGU 102.948; Piasecki et al. 2004). The lowermost interval
(288–193 m), is characterised by dark, silty clays in some cases
interbedded with sandy layers. The succeeding interval (193–183
m) is a medium to coarse-grained sand (Fig. 3). All samples were
processed at the Geological Survey of Denmark and Greenland
in Copenhagen using standard palynological preparation tech-
niques including treatment with HCl, HF, HNO3 and sieving on
11 µm nylon filters (Poulsen et al. 1990). Forty-one samples col-
lected from the 288 to 183 metre interval were examined. Only
two samples were examined from the arenitic 192 to 183 metre
interval of the studied succession (Fig. 3) as these were the only
ones recovered from the drilling.
The taxa used herein represent a combination of natural taxa
and parataxa, and the taxonomy is primarily based on the works
of Thomson and Pflug (1953), Krutzsch (1971), Thiele-Pfeiffer
1980), Nagy (1985), Traverse (1988), Koch (1989), and Kolcon
and Sachsenhofer (1999). Between 340 and 600 terrestrial paly-
nomorphs and additional marine palynomorphs were counted in
each sample to interpret the depositional environment.
Because the assemblages are strongly dominated by the coni-
fers Taxodiaceae-Cupressaceae they mask the perturbation in
the pollen spectra of other groups (Fig. 4A, B). Therefore, we
have excluded Taxodiaceae-Cupressaceae in one of the data sets
(Fig. 4B) in order to better illustrate the quantitative variations
in the other taxa. Paleoenvironmental and vegetation analyses
involved grouping taxa into distinctive plant associations, which
are modified from Kohlman-Adamska (1993) and Kolcon &
Sachsenhofer (1999). In addition, the quantitative relationship
between marine and terrestrial palynomorphs is calculated for
paleoenvironmental purposes (Fig. 3).
The botanical affinities of pollen and spores from recent plant
taxa can be used to aid interpretation of the structure of paleo-
vegetation and ecological tolerances of individual ancient plant
taxa. This can further be used to interpret the ancient climatic
conditions. All taxa were categorized according to botanical af-
finities, in most cases to family or generic level (Table 1), by
using palynological works by, e.g., Thomson & Pflug (1953),
Thiele-Pfeiffer (1980), and Kolcon & Sachsenhofer (1999).
Palynology
All samples contain well-preserved marine and terrestrial
palynomorphs. A total of 95 terrestrial palynomorph taxa were
identified and assigned to 54 botanical groups (Table 1, Fig. 4A,
GFF 128 (2006) Larsson et al.: Early Miocene pollen and spores from western Jylland, Denmark 263
Fig. 2. Upper Oligocene
and Miocene lithostratigra-
phy of Denmark (modified
from Dybkjaer 2004; Ras-
mussen & Dybkjaer 2004).

B). The marine palynomorphs are mainly dinoflagellate cysts,
but algae, such as Tasmanites, and acritarchs also occur. Marine
palynomorphs comprise 20–39% of the palynomorphs in the
three lowermost assemblages, (286–284 m) and 22% at the 260
m level, but the relative abundances of marine palynomorphs
decrease markedly upsection. The present study focuses on the
terrestrial palynomorphs. The dinoflagellate cyst assemblages in
the studied section were studied by Piasecki et al. (2004).
Although vegetation types certainly interact with each other,
five distinct plant associations are recognized here based on
their growth habits and ecology (Fig. 4). Naturally, there will be
overlap in ecological requirements for several taxa and in order
to interpret climate we have been obliged to segregate taxa that
have traditionally been assigned to “arctotertiary” and “paleo-
tropic” associations (Kolcon & Sachsenhofer 1999).
Plants have been grouped into following associations; (1)
Swamp forest association, (2) Cool temperate” mixed meso-
phytic” forest canopy association, (3) Warm temperate plant
association, (4) Mesophytic understorey plant association, and
(5) Aquatic plant and algal association (modified from Kolcon
& Sachsenhofer 1999).
264 Larsson et al.: Early Miocene pollen and spores from western Jylland, Denmark GFF 128 (2006)
Fig. 3. Simplified sedimentary log of the studied suc-
cession (Sequence B) in the Sønder Vium borehole,
showing the relationships between marine and terres-
trial palynomorphs, and between the different plant
associations mentioned in the text.

GFF 128 (2006) Larsson et al.: Early Miocene pollen and spores from western Jylland, Denmark 265
Fig. 4. A. Relative abundance diagram of pollen and spores recorded in the studied succession. B. Relative abundance diagram of pollen and spores
recorded in the lowermost part of the Vium drill core with the Taxodiaceae-Cupressaceae group exluded.

Swamp forest association (Figs. 3, 4A,B)
Pollen from plants that predominantly grow in mire habitats are
by far the dominant palynomorphs in the studied succession
(Figs. 3, 4A). Eight pollen taxa are assigned to this association,
which mainly includes pollen derived from Taxodiaceae-Cu-
pressaceae and Nyssa. Although Taxodiaceae is considered to be
nested within Cupressaceae according to some molecular and
morphological studies (Gadek et al. 2000) it is not always ap-
propriate to group these traditionally separate families, because
their representatives may have different environmental toler-
ances. However, they are grouped here because the preservation
of these palynomorphs is poor in some cases and they are not
always distinguishable at a generic level (Zetter 1998). Note
that Sequoia, which is well preserved, is segregated from this
grouping. Taxodiaceae-Cupressaceae are the most abundant taxa
in this association comprising 42–76% of all identified pollen
grains. The highest relative abundances of Taxodiaceae-Cupres-
266 Larsson et al.: Early Miocene pollen and spores from western Jylland, Denmark GFF 128 (2006)
Table 1. Recorded palynomorph taxa in the lowermost interval of the Vium
drillcore, placed within their known plant affinity (mainly after Thomson &
Pflug 1953; Tschudy & Scott 1969; Krutzsch 1971; Figueiral et al. 1999).
Gymnosperms
Abies
Abiespollenites sp
Cedrus
Cedripites sp.
Larix
Laricipollenites gerceensis Nagy, 1985
Picea
Piceaepollis sp. (Fig. 7P)
Piceaepollis planoides Krutzsch, 1971
Pityosporites alatus (Potonié, 1931) Thomson & Pflug, 1953
Pinus
Pinuspollenites sp.
Pinuspollenites labdacus Potonié, 1958 (Fig. 6O)
Pityosporites microalatus (Potonié, 1931) Thomson & Pflug, 1953, (Fig. 6
P)
Podocarpus
Podocarpidites sp. (Fig. 7O)
Sciadopitys
Sciadopityspollenites serratus (Potonié & Venitz, 1934) Raatz, 1937 (Fig. 6J)
Sciadopityspollenites sp.
Sequoia
Sequoiapollenites sp.
Sequoiapollenites polyformosus Thiergart, 1937 (Fig. 6L)
Taxodiaceae-Cupressaceae
Inaperturopollenites sp. (Fig. 6K)
Inaperturopollenites hiatus Thomson & Pflug, 1953 (Fig. 6N)
Inaperturopollenites dubius Potonié & Venitz, 1934
Inaperturopollenites concepidites Wodehouse, 1933
Inaperturopollenites verrupapillatus Trevisan, 1967
Inaperturopollenites incertus Thomson & Pflug, 1953
Cupressacites sp.
Cupressacites bockwitsensis Krutzsch, 1971 (Fig. 6M)
Tsuga
Tsugaepollenites sp.
Tsugaepollenites robustus Krutzsch, 1971
Zonalapollis sp. (Fig 7Q)
Angiosperms
Acer
Aceripollenites sp.
Alnus
Alnipollenites sp. (Fig. 6F)
Alnipollenites verus Potonié, 1934
Polyvestibulopollenites sp.
Anacardiaceae
Rhoipites pseudocingulum Potonié, 1960 (Fig. 7G)
Araliaceae
Araliaceoipollenites sp.
Arecaceae
“Arecaceaepollenites”
Arecipites sp.
Sabalpollenites papillosus Nagy, 1969
Monocolpopollenites sp.
Monocolpopollenites tranquillus Thomson & Pflug, 1953 (Fig. 7B)
Fig. 5. Stratigraphically important pollen taxa and zones in the upper-
most Oligocene and Miocene of central Europe (from von der Brelie
1966; von der Brelie et al. 1988). Line thickness indicates relative
abudance but this has not been defined quantitatively.

saceae were found in the basal interval between 286–284 m,
although three additional peaks occur at the levels of 260, 257
and 254 metres (Fig. 4A). The occurrence of Nyssa is gener-
ally low, (below 4.5%); however, it should be pointed out that
its representation in this association increases up-section (Fig.
4A). Also included in this association are small trees and shrubs,
such as Betula, Alnus, Myricaceae, Salix, Malvaceae (other than
Tilia), and Cyrilla, which grow in periodically flooded riparian
environments. These palynomorphs are present in low quantities
and show no significant variation in abundance throughout the
investigated succession (Fig. 4A).
When the Taxodiaceae-Cupressaceae group is excluded, the
other elements of the swamp forest association reach a relative
abundance of 6–28% (Fig. 4B). The palynological signals are
enhanced by this procedure and spikes of taxa that previously
where hidden by the overwhelming abundance of Taxodiaceae-
Cupressaceae are now amplified. Most significantly, important
peaks in the typical swamp genera Alnus, Salix, Malvaceae and
Cyrilla are revealed around the 260 m level.
Cool temperate” mixed mesophytic” forest canopy
association, (Figs. 3, 4A, B)
This association comprises 20 identified taxa and constitutes
between 9 and 34% of the total miospore assemblage. Pollen
in this association is derived from conifers and other evergreen
plants but also deciduous angiosperm trees and shrubs, often
referred to as “arctotertiary taxa” (Kolcon & Sachsenhofer
GFF 128 (2006) Larsson et al.: Early Miocene pollen and spores from western Jylland, Denmark 267
Monocolpopollenites areolatus Potonié, 1934
Betula
Trivestibulopollenites betuloides Pflug, 1953
Betulaepollenites betuloides Nagy, 1953 (Fig. 6E)
Carpinus
Carpinipites sp.
Carya
Caryapollenites sp. (Fig. 6C)
Corylus
Triporopollenites coryloides Pflug, 1953
Asteraceae
Artemiseaepollenites sp.
Cyrilla
Cyrillacaepollenites exactus Potonié, 1960
Engelhardtia
Engelhardtioidites sp. (Fig. 7J)
Ephedraceae
Ephedripites sp.
Ericaceae
Ericipites sp. (Fig. 7M)
Fagaceae?
Tricolporopollenites pusillus (Potonié, 1931) Thomson & Pflug, 1953 (Fig. 6G)
Tricolporopollenites fusus Potonié, 1934
Fagus
Faguspollenites sp. (Fig. 6I)
Poaceae
Graminidites sp. (Fig. 7L)
Triatriopollenites coryloides Pflug, 1953 (Fig. 7K)
Ilex
Ilexpollenites sp.
Tricolpopollenites iliacus (Thiergart, 1937) Potonié, 1960
Liquidambar
Liquidambarpollenites sp.
Magnoliaceae
Liriodendroipollis semiverrucatus Krutzsch, 1971
Magnolipollis neogenicus minor Krutzsch, 1971
Malvaceae
Malvacearumpollis sp.
Mastixiaceae?
Tricolporopollenites edmundii (Potonié, 1931) Thomson & Plug, 1953 (Fig.
7C)
Myricaceae
Triatriopollenites myricoides Kremp, 1949
Triatriopollenites bituitus (Potonié, 1931) Thomson & Pflug, 1953 (Fig. 6D)
Triatriopollenites rurensis Thomson & Pflug, 1953
Nymphaeaceae
Nupharipollenites kedvesi Nagy, 1969
Nymphaeaepollis minor Nagy, 1985
Nyssa
Nyssapollenites sp. 1960 (Fig. 6H)
Plantaginaceae
Plantaginacearumpollis miocaenicus Nagy, 1963
Platanaceae
Platanus sp.
Platycarya
Platycarypollenites sp. (Fig. 7I)
Pterocarya
Pterocaryapollenites sp.
Quercus
Quercopollenites sp. (Fig. 7D)
Quercus?
Tricolpopollenites henrici (Potonié, 1931) Thomson & Pflug, 1953 (Fig. 7H)
Tricolpopollenites microhenrici (Potonié, 1931) Thomson & Pflug, 1953 (Fig. 7F)
Rosaceae
Rosaceae-type
Salix
Salixipollenites sp.
Sapotaceae
Sapotaceoidaepollenites sp. (Fig. 7E)
Sparganiaceae
Sparganiaceaepollenites sp.
Symplocaceae
Porocolpopollenites sp.
Tilia
Tilia sp. (Fig. 7N)
Ulmus
Ulmipollenites sp.
Zelkova
Zelkovaepollenites potoniei Nagy, 1969
Pteridophytes
Lycopodiaceae
Polypodiaceae
Polypodiisporites sp.
Foveotriletes sp.
Baculatisporites sp.
Gleicheniidites senonicus Ross, 1949
Laevigatosporites sp. (Fig. 6B)
Laevigatosporites hardtii Thomson & Pflug, 1953
Leiotriletes sp.
Osmundaceae
Triplanosporites sp.
Bryophytes
Stereisporites sp. (Fig. 6A)
Algae
Sigmopollis sp.
Ovoidites sp.
Botryococcus braunii Kützing, 1849
Tasmanites sp.
Acritarchs
Dinoflagellates
Fungal spores

268 Larsson et al.: Early Miocene pollen and spores from western Jylland, Denmark GFF 128 (2006)
1999). The association is dominated by conifer pollen such as
Pinus (1–16%), Abies (0–4%), Picea (0–2%), Tsuga (0–4%), and
Sequoia (0.3–4%); (Fig. 4). Pollen from deciduous trees such
as Fagus (0–8%) and Quercus (up to 3%) are also relatively
frequent. Minor constituents include, among others, Carpinus,
Ulmus, Corylus, Magnoliaceae, and Tilia. Other significant taxa
include Araliaceae and Rosaceae. Where the Taxodiaceae-Cu-
pressaceae is omitted (Fig. 4B) a more pronounced but basically
identical palynological signal is recorded.
Fig. 6. Photomicrographs of selected palynomorph taxa from the studied succession, magnification ×700. A. Stereisporites sp. B. Laevigatosporites
sp. C. Caryapollenites sp. D. Triatriopollenites bituitus (Potonié 1931) Thomson & Plug, 1953. E. Betulaepollenites betuloides Nagy, 1969. F.
Alnipollenites sp. G. Tricolporopollenites pusillus (Potonié, 1931) Thomson & Pflug, 1953. H. Nyssapollenites sp. I. Faguspollenites sp. J. Scia-
dopityspollenites serratus (Potonié &Venitz, 1934) Raatz, 1937. K. Inaperturopollenites sp. L. Sequoiapollenites polyformosus Thiergart, 1937 M.
Cupressacites bockwitzensis Krutzsch, 1971. N. Inaperturopollenites hiatus Thomson & Pflug, 1953. O. Pinuspollenites labdacus Potonié, 1958.
P. Pityosporites microalatus (Potonié, 1931) Thomson & Pflug, 1953.

GFF 128 (2006) Larsson et al.: Early Miocene pollen and spores from western Jylland, Denmark 269
Fig. 7. Photomicrographs of selected palynomorph taxa from the studied succession, magnification ×700. A. Liriodendroipollis semiverricatus
Krutzsch, 1971. B. Monocolpopollenites tranquillus Thomson & Pflug, 1953. C. Tricolporopollenites edmundii (Potonié, 1931) Thomson &
Pflug, 1953. D. Quercopollenites sp. E. Sapotaceoidaepollenites sp. F. Tricolpopollenites microhenrici (Potonié, 1931) Thomson & Pflug, 1953.
G. Rhoipites pseudocingulum Potonié, 1960. H. Tricolpopollenites henrici (Potonié, 1931) Thomson & Pflug, 1953. I. Platycaryapollenites sp. J.
Engelhardtioidites sp. Potonié, 1960. K. Triatriopollenites coryloides Pflug, 1953. L. Graminidites sp. Cookson, 1947. M. Ericipites sp. Wode-
house, 1933. N. Tilia sp. O. Podocarpidites sp. P. Piceapollis sp. Q. Zonalapollis sp.

270 Larsson et al.: Early Miocene pollen and spores from western Jylland, Denmark GFF 128 (2006)
Warm temperate plant association (Figs. 3, 4A, B)
This association comprises 15 thermophilous taxa, representing
the “palaeotropic flora” (Kolcon & Sachsenhofer 1999), and
4–16% of the total miospore assemblage. The main constituents
are angiosperm taxa, among others, Arecaceae (1–8%), Mastix-
iaceae (0–3%), Engelhardtia (0–3%), Ilex (0–1%) and Platy-
carya (0–1%). Gymnosperms are represented by the presence of
conifer pollen such as Sciadopitys (0.2–4.5%), Cedrus (0–1.5)
and Podocarpus (0–1%). The warm temperate taxa included in
this group grew in different habitats. Thus, the quantitative vari-
ations mainly serve to interpret the paleoclimate. After exclud-
ing the Taxodiaceae-Cupressaceae, a significant decrease in the
relative abundance of the warm temperate plant association is
revealed above the 247 m level (Fig. 4B).
Mesophytic understorey plant association (Figs. 3, 4A, B)
This association consists of five taxa that constituted ground-
cover vegetation in the mesophytic forests. This group makes
the smallest contribution to the total palynoflora comprising
only 1–7% of the total miospore assemblage. Pteridophytes are
the dominant group in this association, comprising up to 6% of
total terrestrial palynomorphs (Fig. 4A). Other taxa contributing
to the association are bryophytes, Asteraceae, Ericaceae, and
Plantaginaceae. When excluding Taxodiaceae-Cupressaceae,
the same general trend in relative abundance is evident for this
association (Fig. 4B).
Aquatic plants and algal association (Figs. 3, 4A, B)
This association includes palynomorphs from plants that prefer
mainly moist environments (although Poaceae frequently occur
as undergrowth plants in other communities). In the studied suc-
cession this association varies between 1 and 6% (Figs. 3, 4A).
Taxa included are Poaceae, Sparganiaceae, and Nymphaeaceae.
Poaceae occur in almost every sample, comprising up to 4%
of the total miospore assemblage (Fig. 4A), although it is most
abundant in the lowermost samples. The aquatic taxon Nym-
phaeaceae is present in almost every sample although it peaks
at the 260 m level where it comprises 3% of the total miospore
assemblage (Fig. 4A).
Since this study deals primarily with paleovegetational recon-
struction, freshwater taxa, such as Sigmopollis, Ovoidites, and
Botryococcus are included in this association. Freshwater taxa
are present in almost every sample, except in those from the basal
part of the succession (Fig. 4A). After excluding Taxodiaceae-
Cupressaceae the previously mentioned spike of Nymphaeaceae
is enhanced and an additional spike of Sigmopollis occurs at the
260 m level (Fig. 4B).
Discussion
Age of the succession
Palynological subdivision of the northern European Miocene is
difficult for several reasons. Firstly, most species range through-
out the epoch (von der Brelie 1967). Secondly, major differences
in phytogeographic distributions and depositional environments
occurred in Europe during this time, hence there are difficulties
in correlating the pollen spectra at different locations (Traverse
1988; Sadowska 1995).
Based on quantitative differences in the pollen spectra of cen-
tral Europe, von der Brelie (1967) established five pollen zones
for the Miocene, which was enhanced by the more general study
of von der Brelie et al. (1988; Fig. 5). The palynological results
from the Sønder Vium succession have been compared with the
quantitative results of von der Brelie (1967) and von der Brelie
et al. (1988) in order to assess the applicability of the central
European zonation to Denmark (Fig. 5). Von der Brelie (1967)
indicated that the ranges and relative abundance of certain pol-
len taxa such as Tricolpopollenites henrici and T. microhenrici
were stratigraphically significant as they show a slight increase
in abundance at the Aquitanian–Burdigalian boundary. Results
from the studied succession indicate that these species represent
consistent but subsidiary elements throughout the sampled suc-
cession. In addition, Paleogene key pollen, such as Ephedripites,
disappear by the end of the Aquitanian whereas Engelhardtia
occurs throughout the Miocene, but is most common during the
Early Miocene (Fig. 5; von der Brelie 1967; von der Brelie et
al. 1988). These taxa were consistently recorded throughout the
sampled portion of the Sønder Vium core.
An Early Miocene age is indicated by the presence of Ephe-
dripites, Platycarya, and the consistent occurrence of Engelhar-
dtia. Based on the known range of Ephedripites (von der Brelie
1967), the studied section should be assigned to the Aquitanian.
In addition, the increasing level of Rhoipites pseudocingulum
(Anacardiaceae), accompanied by successively decreasing
amounts of palm pollen suggest that the studied succession is
possibly of late Aquitanian age.
According to Mai (1995), the climate in Europe was warm
temperate to humid subtropical, with slight climate deterioration
at the end of the Aquitanian. Climate change is reflected in the
studied succession by a general reduction in warm temperate taxa
starting at the level of 262 metres. However, a more pronounced
reduction in warm temperate taxa above 247 metres (sample
133) is revealed when excluding Taxodiaceae-Cupressaceae
from the pollen signal (Fig. 4B). We contend that this climatic
deterioration may indicate a late Aquitanian age for this part of
the succession.
The new spore-pollen data supports a late Aquitanian age for
the studied succession in the Sønder Vium bore core (sequence
B), as proposed by previous authors (Dybkjaer 2004; Piasecki et
al. 2004) based on dinoflagellate cyst assemblages.
Depositional environment, vegetation and climatic
conditions
The consistent occurrence of marine dinoflagellate cysts in all
the studied samples, combined with the strong dominance of
terrestrial palynomorphs implies that deposition occurred in an
inner-neritic setting. This interpretation agrees well with seismic
data from the studied area. Here southwards dipping clinoforms
indicate an Early Miocene delta progradation (Rasmussen
2004a). Samples showing increased relative abundances of dino-
flagellate cysts, e.g., the level of 286–284 m, probably reflect a
minor transgressional phase resulting in less terrestrial influence
in the studied area (Fig. 3).
The Sønder Vium terrestrial palynomorph assemblages indi-
cate no major change in diversity as most taxa persist through
the investigated succession. However, the relative abundance of
taxa varies. All five palynological associations show pronounced
fluctuations in relative abundance in the lower one-third of the
studied succession compared to the more stable composition in

GFF 128 (2006) Larsson et al.: Early Miocene pollen and spores from western Jylland, Denmark 271
the upper two-thirds (Fig. 4A, B). This signal is enhanced when
the Taxodiaceae-Cupressaceae are excluded. The transition from
variable to more uniform assemblages occurs at the 247 m level
and is coincident with the vegetational signal of climatic deterio-
ration. We contend that the relatively uniform palynofloras from
the upper part of the succession reflect a more stable climate
during the time of deposition, but their homogeneity could also
indicate a higher sedimentation rate (i.e., shorter period of depo-
sition).
A warm and humid climate during the Early Miocene favoured
the widespread development of herbaceous mires and peat-
forming Taxodium swamp forests in adjacent coastal lowlands
of central Europe and northwestern Germany (Ziegler 1990;
Figueiral et al. 1999; Kolcon & Sachsenhofer 1999; Bechtel
et al. 2002). These wet lowlands were surrounded by elevated
forests incorporating both palaeotropical and arctotertiary ele-
ments (e.g., Thiele-Pfeiffer 1980; Sadowska 1995; Zetter 1998;
Kovar-Eder et al. 2001; Bechtel et al. 2002). The Sønder Vium
assemblages extend the distribution of this paleovegetation type
and imply swamp forests in coastal Denmark in the earliest Mi-
ocene. Modern analogues for these are the river swamp forests
of the Mississippi delta, where the canopy is co-dominated by
Taxodium and Nyssa.
Mesophytic forests dominated the vegetation in moderately wet
areas further inland. The pollen spectra indicate that these were
rather diverse forests incorporating both warm and cool temperate
elements, although the latter dominate the assemblage. The most
abundant warm temperate taxa in the mesophytic assemblage,
besides Arecaceae, are Mastixiaceae, Ilex and Engelhardtia. Pi-
nus, which is the most abundant taxon in the mesophytic forest,
might also have been quite common in the better drained, sandy
parts of the swampy forest (Zetter 1998).
The ground vegetation of the mesophytic forests and open areas
of swamp forests was probably dominated by reed, sedges and
pteridophytes. These groups maintain stable relative abundances
and do not appear to have been influenced by transgressive/re-
gressive events. This category of plants undoubtedly includes
taxa derived from many habitats with varied characteristics.
In the lower part of the succession dinoflagellates are abundant,
whereas fresh and brackish water algae are absent. An increase
in fluvial input up-section is corroborated by higher percentages
of Botryococcus (Fig. 4A, B). Higher in the studied succession
freshwater palynomorphs such as Sigmopollis become more
abundant, together with pollen derived from aquatic plants, such
as Nymphaeaceae, indicating the occurrence of open freshwater
ponds in the Taxodium swamp forests (Zetter 1998; Masselter &
Hofmann 2005).
The individual elements of the family Poaceae are difficult
to identify as the pollen grains of different genera do not show
significant morphological differences. However, comparisons
with the floristic composition of extant riparian swamp forests
provide clues to the likely representatives of this family in the
Miocene flora. In the palynological assemblages from Sønder
Vium, it is most likely that the Poaceae would have been repre-
sented mostly by reeds and other hydrophilous graminoids rather
than xeric grasses. As the relatively high abundance of Poaceae
coincides with the strong representation of warm temperate taxa
and Nymphaeaceae, it is possible that bamboo was a constitu-
ent of this swamp forest community. Bamboo has previously
been recorded from several Neogene localities in Europe, both
as macro and microfossils and coexisting with typical swamp
forest taxa, such as Taxodiaceae-Cupresseace, Alnus and Salix
(Worobiec & Worobiec 2005). In addition, the entomophilous
pollen taxon Mastixiaceae, an autochthonous element, common
in warm and humid climates (Masselter & Hofmann 2005), is
also relatively abundant in the Sønder Vium assemblages. We
contend that the palynological signal reflects a transition to a
slightly cooler climate at the end of the Aquitanian.
The ratio between warm and cool-temperate taxa in the Lower
Miocene of central Europe is typically 1:1 (Kolcon & Sach-
senhofer 1999). This is not the case for the palynofloras in the
succession studied here, which show a slight dominance of cool
temperate elements. This is probably due to the higher paleolati-
tude of the studied area compared to previous studies carried out
on sediments from central Europe.
The increased abundances of swamp forest pollen (Taxo-
diaceae-Cupressaceae group) at the levels of 286–284 m, 260
m, 257 m, and 254 m correlates well with increased relative
abundances of dinoflagellates. Taxodiaceae-Cupressaceae pollen
probably derives from swamp forest communities that dominated
the lower delta plain. Interestingly, around the 260 m level the
palynofloras contain relatively high percentages of hydrophilous
taxa (Nymphaeaceae, Arecaceae, Alnus, Salix, Malvaceae and
Cyrilla) presumably derived from riparian and coastal settings.
This reinforces our interpretation that the lower portion of the
studied succession was deposited in an inner-neritic environment
during a transgressional event. In contrast, samples with abun-
dant pollen of mesophytic forest plants above the 247 m level
have low levels of dinoflagellates and swamp forest pollen, and
these intervals may correspond to delta progradation.
Conclusions
The miospore assemblages from the studied succession of
Sønder Vium are mainly well preserved and incorporate 95 ter-
restrial taxa. A near-shore pro-deltaic environment is suggested
by dinoflagellates in all samples. The lowermost samples in the
succession include abundant dinoflagellate cysts and probably
represent a transgressional event. A late Aquitanian age is indi-
cated by the presence of Ephedripites, Platycarya, and relatively
frequent occurrences of Engelhardtia, together with the increas-
ing levels of Rhoipites pseudocingulum (Anacardiaceae), and
coeval decreasing amounts of Arecaceae.
Five distinct terrestrial plant associations, can be recognised
(1) Swamp forest canopy association, (2) Cool temperate” mixed
mesophytic” forest canopy association, (3) Warm temperate
plant association, (4) Mesophytic understorey plant association,
and (5) Aquatic plants and algal association. Previous work in
Europe by Bechtel et al. (2002) and Kolcon & Sachsenhofer
(1999) have shown that Taxodium swamp forests dominated
central Europe. This study extends the northern limit of this
swamp forest vegetation, at least as far as present Denmark.
Better drained areas further inland hosted diverse mesophytic
forests. They included both warm and cool temperate taxa, with
a dominance of the latter. The presence of Arecaceae (palms)
and other warm temperate taxa such as Ilex and Engelhardtia,
Mastixiaceae together with the widely distributed Taxodium
swamps, suggest that a warm temperate climate prevailed with
slight climatic deterioration reflected in the upper part of the suc-
cession possibly corresponding to the climatic deterioration in
the late Aquitanian.
Acknowledgements. – Professors Else Marie-Friis is gratefully acknowledged for carefully
reviewing and improving this paper with her comments and suggestions. Karen Dybkjaer
and Stefan Piasecki [Geological Survey of Greenland and Denmark (GEUS)] are thanked
for providing the samples and for sharing their expertise on the Danish Miocene. Steve

McLoughlin is acknowledged for sharing his botanical knowledge and for improving an
earlier version of the manuscript together with Johan Lindgren and Anita Löfgren. This
project was partly supported by the Royal Physiographic Society and Lunds Geologiska
Fältklubb (L.L) and by the Swedish Research Council (V.V.).
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