Palaeogeography, Palaeoclimatology, Palaeoecology 365366 (2012) 8198

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Palaeogeography, Palaeoclimatology, Palaeoecology

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Paleoclimatology and Paleoecology across the Eocene/Oligocene boundary, Thrace

Basin, Northeast Aegean Sea, Greece
Angelos Maravelis , Avraam Zelilidis
Laboratory of Sedimentology, Department of Geology, University of Patras, 26500 Rion, Greece

a r t i c l e

i n f o

Article history:
Received 30 January 2012
Received in revised form 8 August 2012
Accepted 14 September 2012
Available online 23 September 2012
Keywords:
Biostratigraphy
Ichnology
Calcareous nannofossils
Eocene
Oligocene
Lemnos Island

a b s t r a c t
The detailed biostratigraphic and ichnological analysis of the Upper Eocene/Lower Oligocene forearc sedimentary sequence in the southern Greek part of the Thrace Basin on Lemnos Island reveals a relationship between eco-sedimentary factors and the base-level dynamics inuencing calcareous nannofossils and trace
fossil assemblages. The lower part of the ~ 350 m thick succession was deposited in a sand-rich submarine
fan system and upper part in a shelf environment. It contains nannofossil assemblages and distributions typical of the NP19NP21 biozones. The Eocene/Oligocene (E/O) boundary (top of NP20 biozone) is clearly distinguished within the upper parts of the turbidite system where a decrease in water temperature, salinity,
nutrient supply and oxygenation level is also revealed. Additionally, 19 ichnospecies, mostly graphoglyptids
typical of deep-sea fan deposits, have been distinguished. The trace fossil diversity and abundance decrease
upward reecting the impact of the changing depositional environments from basin oor fans, to slope systems and then shelfal environments. The regional basin-ll history, rather than global climatic changes, was
probably the fundamental controlling factor on the distribution of graphoglyptids and other trace fossils
across the E/O boundary.
2012 Elsevier B.V. All rights reserved.

1. Introduction
The Eocene/Oligocene (E/O) boundary is of broad interest because it
marks the transition between the Paleogene greenhouse and Neogene
icehouse climates. The Oligocene in particular, is recognized as a period
marked by large and abrupt paleogeographic and climatic changes including the opening of the Tasmanian Gateway and Drake Passage
and large uctuations in the Antarctic Ice Sheet volume after its initiation in the earliest Oligocene, with associated eustatic changes at orbital
frequencies (Wade and Plike, 2004). Although traditionally it has been
argued that glaciation in Antarctica started much earlier than in the
northern hemisphere (Zachos et al., 2001), more recent evidence suggests that at least some level of glaciation in the Northern Hemisphere
started at approximately the same time as in the southern hemisphere,
during the middle Eocene to early Oligocene (Moran et al., 2006; Eldrett
et al., 2007).
Studies of modern calcareous nannofossil assemblages indicate that
the species makeup and community structure of assemblages are highly
sensitive to climate related factors, such as surface water stratication,
nutricline depth and nutrient availability (Okada and Honjo, 1973).
The response of the calcareous phytoplankton to global change through
the E/O boundary is poorly understood compared to other critical intervals such as the Paleocene/Eocene boundary (e.g. Bralower, 2002; Gibbs
et al., 2006). One set of high-resolution records from the Southern
Corresponding author. Tel./fax: +30 2610996272.
E-mail address: mgalith@gmail.com (A. Maravelis).
0031-0182/$ see front matter 2012 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.palaeo.2012.09.015

Ocean has shown a signicant fall in nannofossil diversity and an increase in cold water taxa directly coincident with the onset of the Oligocene (Persico and Villa, 2004). Currently, there are no comparable data
sets from low latitudes, with previous quantitative records being either
low resolution (Wei and Wise, 1990; Wei et al., 1992) or severely
compromised by poor preservation (Monechi, 1986; Coccioni et al.,
1988). There was a deterioration of microbenthic faunal environmental
conditions (Jovane et al., 2009).
The ethological behavior of turbidite infauna is greatly inuenced by
water depth. As depth increases, bioturbation becomes more sensitive
to changes in sedimentation rates and substrate consistency as well as
other chemical and physical parameters such as temperature, salinity,
oxygenation levels and pressure conditions (Seilacher, 1967; Uchman,
2007). Ichnology is today recognized as an important tool for environmental analysis (Savrda, 1995) essentially because most trace fossils
are autochthonous indicators of ecologic conditions. Environmental
changes recorded by trace fossils can be revealed through analysis of
ichnofabrics (Bromley and Ekdale, 1986). Ichnofabrics result from bioturbation processes that vary in response to changes in sediment accumulation rates and other environmental factors (Taylor and Goldring,
1993). Ecologic conditions change with depth below the sediment
water interface. The sediment strength increases, organic matter becomes decomposed, porosity and permeability decrease while the
pore-water oxygenation level decreases (Bromley, 1996).
The Upper Eocene/Lower Oligocene deep-sea fan and shelf sequence
in the southern part of the Greek Thrace Basin on Lemnos Island provides a great opportunity to analyze the calcareous nannofossil and

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trace fossil distributions across the E/O boundary in a low-latitude

setting. The biostratigraphy of these deposits has previously been
discussed by Maravelis et al. (2007) but as of yet, systematic ichnology
has not been documented. This research, by integrating sedimentology,
ichnology, and biostratigraphy, attempts to study the calcareous
nannofossil assemblages and deep-sea trace fossil distribution in relation to factors caused by variations in depositional environments,
sea-level dynamics and climate changes.
2. Tectonostratigraphy
The study area lies in the southern Greek part of the Thrace Basin. The
plate conguration of the Aegean region consists of the Aegean Plate to
the south separated by a strikeslip boundary from the Eurasian Plate
to the north, which encompasses the north Aegean, Rhodope and adjacent area (McKenzie, 1970) (Fig. 1). Convergence between the Eurasian
and African plates has played a key role in controlling magmatism in the
Balkan Peninsula since the Late Cretaceous. During the Late Eocene/Early
Oligocene, because of the subduction of the African Plate beneath the
Eurasian Plate, magmatic activity occurred in the Macedonian
RhodopeNorth Aegean region (Harkovska et al., 1989; Marchev and
Shanov, 1991). The magmatic belt extends to the northwest into Skopje
and Serbia, crossing the Vardar Zone and continues towards the southeast, in the Thraciane Basin and Western Anatolia (Yilmaz and Polat,
1998; Aldanmaz et al., 2000).
The Thrace Basin has been considered as a fore arc basin of the
contracted type (Grr and Okay, 1996; Maravelis and Zelilidis,
2010a) and is the largest and thickest Tertiary sedimentary basin
of the eastern Balkan region (Turgut et al., 1991; Turgut and Eseller,
2000). It is exposed on Greek, Turkish and Bulgarian territory and comprises an important hydrocarbon province (e.g. Turgut and Eseller,
2000; Siyako and Huvaz, 2007; Maravelis and Zelilidis, 2010b;
Maravelis and Zelilidis, 2012). A large accretionary prism in the Central Aegean region bounds the basin on the seaward side, and the
landward ank was bound by an active volcanic arc, the Rhodope

Zone. Most of the basin strata are Lower Eocene to Upper Oligocene
and form thick sedimentary successions (up to 9000 m) made of
deep-sea fan deposits (Turgut et al., 1991; Turgut and Eseller,
2000). Sedimentation along the basin margin was dominated by deposition of carbonates during the Eocene and by deltaic bodies,
prograding towards the basin center, in the Oligocene (Smengen
and Terlemez, 1991; Turgut et al., 1991).
The Greek part of the Thrace Basin in Lemnos Island and Rhodope
was mainly inuenced by two major sediment inputs (Maravelis and
Zelilidis, 2010a, 2012). The southern part (Lemnos) was signicantly
affected by the accretionary prism and associated ophiolitic units
e.g. on Lesvos Island (Maravelis and Zelilidis, 2010a, 2012) while the
northern part (Greek Rhodope) reects a CircumRhodope Belt inuence (Maravelis and Zelilidis, 2010a; Caracciolo et al., 2011). On Lemnos,
the source area contributed signicant a volume of ultramac, gabbro,
basalt, chert, and possibly some volcaniclastic detritus of variable grain
size into the forearc basin and was probably located south-southwest
of Lemnos (Maravelis and Zelilidis, 2010a). The source area was probably rugged and rapidly eroded, causing the ophiolitic bedrock to be
deeply incised, enabling a signicant amount of coarse-grained material
from the uplifting source area to be transported to the Lemnos area
(Maravelis and Zelilidis, 2010a). This material was deposited in submarine fans in the Lemnos area, which, as a result of tectonic activity, are
overlain by shelf deposits (Maravelis et al., 2007) (Fig. 2). The submarine
fan is a sand-rich system, which comprises a basin-oor fan overlain by
a slope fan, suggesting submarine fan progradation. Both basin-oor
and slope fans are characterized by monotonous alternations of sandstone and mudstone beds (Maravelis et al., 2007).
During the Miocene, Lemnos was the site of volcanic activity resulting
in accumulation of magmatic rocks (Pe-Piper and Piper, 2001). Both plutonic and volcanic rocks accumulated, principally trachyandesites and
dacites, and cover a large part of the studied area. These rocks have
been interpreted as belonging to the high-K province along the
AegeanAnatolian boundary (Pe-Piper et al., 2009). In particular,
they belong to the northern, shoshonitic province that includes

Fig. 1. Simplied sketch map and plate tectonic conguration of the Eastern Mediterranean region and adjacent regions. Inset: Schematic diagram illustrating the depositional setting of
the Lemnos Island forearc region (Modied from Maravelis and Zelilidis, 2010a). SK Skopje, RZ Rhodope Zone, VR Vardar Zone, TB-WA Thrace Basin-Western Anatolia.

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83

Fig. 2. Generalized chart of the Late Eocene to Holocene stratigraphy of Lemnos.

the islands of Samothrace, Lemnos and Lesvos (Pe-Piper et al., 2009).

These high-K rocks, mostly of intermediate composition, indicate ensuing calc-alkaline orogenic volcanism, emitted from large volcanic centers. Upwelling of asthenospheric mantle has been invoked to account
for their genesis (Pe-Piper et al., 2009). The end of the Miocene is characterized by the deposition of conglomerates, marls and calcareous
sandstones. Local Pleistocene porous calcareous and locally oolitic limestones and Holocene alluvial, coastal deposits and dunes are sparse.
3. Methods
For nannofossil analysis, forty-seven (47) samples were collected
from Lemnos (Fig. 3). These samples were mudstones from the
sand-rich turbidite system and the overlying shelf deposits (Fig. 4).
Ichnofaunal research conducted on these deposits was based on extensive eldwork and observation of the trace fossil communities.
The samples were prepared from unprocessed sediment as smear slides
using standard techniques (Bown and Young, 1998). Smear slides were
then analyzed with an optical microscope, at 1250 magnication.
Biostratigraphic data were obtained using the taxonomic concepts of
Perch-Nielsen (1985), Young (1998), Aubry (1999) and Bown (2005).
Biostratigraphic events are dened as species rst appearance datum
(FAD) and last appearance datum (LAD). The schemes of Martini
(1971) were adopted for the biostratigraphic study. Moreover, the high
latitude zonal scheme of Wei and Wise (1990) proved to be suitable
for the detailed biozonal attribution of the studied nannofossil assemblages. In fact, both classical biozonal events of low latitude (Martini,
1971; Okada and Bukry, 1980) and high latitude (Wei and Wise, 1990)
schemes were recognized, dening all biozones from the Upper Eocene
to Lower Oligocene interval. The essential FAD and LAD events have
been distinguished, and a biostratigraphic zonation according to the classical denition of the standard zonation, can be established. Where index
species were not observed, secondary index and characteristic species
were used.
The trace fossils are classied in the morphological groups introduced by Ksikiewicz (1977) and later modied by Uchman (1995a).
The toponomic classication of Martinsson (1970) is used to describe
the position of the trace fossils in relation to the sandstone beds. This
scheme describes four trace fossil positions: endichnia (within the
bed), hypichnia (at the base of the bed), epichnia (on the top of the
bed), and exichnia (external to the bed).
4. Sedimentology
The sand-rich submarine fan system on Lemnos can be subdivided
into a basal basin-oor fan and an overlying slope fan. The slope fan is

Fig. 3. Chronostratigraphy, sedimentary environments and lithology of the Upper Eocene/

Lower Oligocene sedimentary succession, Lemnos Island, Northeast Aegean Sea, Greece.
Red circles mark the sample position (modied from Maravelis and Zelilidis, 2010a).

further subdivided into a channelll and levee system that evolves

upwards into a conglomeratic channelll system (Maravelis et al.,
2007; Maravelis and Zelilidis, 2011).

4.1. Basin-oor fan

4.1.1. Description
The basin-oor fan is up to ~150 m thick and is characterized by light
brown to light green, very ne-grained sandstones which are overlain by
ne to medium-grained sandstones, interbedded with hemipelagic
mudstones. The sandstones have at bases, are laterally continuous
and accumulated in sedimentary bodies. Some of these bodies exhibit
distinct coarsening and thickening upward trend (Fig. 5A, B) although
deposits with no similar trend are common (Fig. 5C). The sandstone
beds exhibit parallel cross- and/or climbing ripple-laminations (Bouma
Tb and Tc sandstones) (Fig. 5D, E) and the cross-laminations are often
overlain by convolute laminations (Fig. 5F). The mudstones are brown
in color and typically lack internal structures, although beds with silt
laminae occur. To provide insights into the paleoow direction,
paleocurrent data were collected from several outcrops. Approximately one hundred and fty ute and groove marks were measured
and plotted in rose diagrams that exhibited a NE/NNE trend (Maravelis
et al., 2007).

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Fig. 4. Geologic map of Lemnos Island displaying the distribution of the sedimentary environments. Red circles mark the sample position (modied from Maravelis and Zelilidis,
2010a, 2010b).

4.1.2. Interpretation
The sedimentary facies observed suggest deposition from both low
and high-density turbidity currents. The common occurrence of convolute laminations and/or climbing ripples indicates rapid deposition of
sediment from suspension. This is interpreted to be related to the dilution of ows which is associated with the transition from high gradient,
conned and channelized environments, to unconned, gently dipping depositional lobe areas. The occurrence of distinct, small-scale,
thickening-upward cycles within major thickening-upward cycles,
indicates a lobe sequence with both aggradation and progradation,
been responsible for lobe development (Mutti and Sonnino, 1981).
Sedimentary bodies with lack of well-dened trends in bed thickness
and grain-size, indicate formation by vertical aggradation, rather than
basinward progradation, typical of lobe-fringe and fan-fringe facies
(Hiscott, 1981; Maravelis et al., 2007).
4.2. Slope fan
4.2.1. Description
The slope fan is up to 100 m thick and is characterized by very thin
to very thick-bedded sandstones and conglomerates interbedded
with hemipelagic mudstones. The sandstone beds have sharp bases
and are commonly erosive (Fig. 6A), laterally discontinuous and accumulated in packages which exhibit distinct ning and thinning upward trend (Fig. 6B, C). Sandstones are light brown to light green in
color and display both complete and incomplete Bouma sequences
(Fig. 6D, E, F). Laterally, these deposits evolve into a slump unit of
interbedded, ne-grained sandstones and mudstones. Sandstones
have lenticular geometry exhibiting cross-lamination. Conglomerates
are disorganized or have rare inverse to normal grading, polymict,
and consist of gneissic schist, radiolarian chert, calcareous, arenaceous,
or quartzitic cobbles in arenaceous cement. Mudstones are brownish
and typically lack internal structure although beds with silt laminae
occur.

4.2.2. Interpretation
Both the thick-bedded, structureless sandstones and the normally graded sandstones with parallel laminations and/or ripple crosslaminations are interpreted as having been deposited by high density, waning turbidity currents. The conglomerates were deposited by
traction-carpets at the base of high concentration turbidity currents,
non-cohesive sandy debris ows or grain ows. The sharp and commonly erosionally based thick-bedded sandstones, which exhibit laterally discontinuous geometries, are interpreted to represent channelll
deposits. The thin-bedded sandstones with parallel laminations and/or
ripple cross-laminations are interpreted as being channel-overbank deposits. The slump unit is interpreted as having developed in response to
failure of aggraded overbank deposits (Maravelis et al., 2007).
4.3. Shelf
4.3.1. Description
The overlying shelf deposits are 70/100 m in total thickness and characterized by a general ning upward trend (Fig. 7A, B). The basal deposits
comprise sandstones interbedded with very thin-bedded mudstones.
Most of the sandstones are structureless, however, some display grooves
and tool marks. Internal structures are dominated by prominent parallel
lamination and the top of sandstone beds commonly exhibit a single set
of ripple cross-laminae. The mudstones commonly contain a high proportion of plant debris (Fig. 7C). The upper part of the shelf succession
is mud-dominant, consisting of massive, homogeneous green or greengray mudstones.
4.3.2. Interpretation
The sandstones of this unit differ signicantly from the turbidite
sandstones of the underlying units and are interpreted as storm-surge
sandstones deposited on outer shelf. The overall ning upward trend
observed in outer shelf deposits has been attributed to turbidity currents and to shelf storm currents (Maravelis et al., 2007).

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85

Fig. 5. Outcrop photographs of the basin-oor fan deposits. (A), (B) Lobe deposits with typical upward increase in sandstone/mudstone ratio. (C) Lobe fringe facies. Note the absence of a distinct thickening and coarsening upward trend. (D), (E) Typical thin-bedded, parallel and ripple-laminated sandstones in the basin-oor fan. (F) Medium-bedded, normal grading, sandstone that exhibits cross-laminations overlaying by convolute laminations.

5. Calcareous nannofossil biostratigraphy

The nannofossil assemblage is generally rich and well diversied, with
429 taxa in the entire dataset. The recovery varies from poor to very good
and most of the samples (41) contain nannofossil taxa. Six (6) samples
contain rare calcareous nannofossils with abundant siliciclastic material,
and/or reworked specimens, making the biostratigraphic identication
very difcult (barren or essentially barren). The average sample species
richness, excluding obviously reworked species, for the dataset is 10
to12, with a maximum observed value of 16 (samples H9 and D5) and
no slide had less than 3 species (E8), if reworked taxa were excluded.
The nannofossil assemblage is indicative of Late Eocene/Early Oligocene
(NP19/NP21 biozones) age. Reworked forms derived mainly from the
Cretaceous, the Paleocene and a few from the Early/Mid Eocene.
Cretaceous species are: Arkhangelskiella cymbiformis, Bomolithus
elegans, Cretarhabdus crenulatus, Cribrosphaera ehrenbergii,
Cyclagelosphaera margerelii, Eifellithus eximius, Eifellithus turriseifellii,
Fasciculithus tympaniformis, Lithraphidites acutus, Lucianorhabdus

cayeuxii, Microrhabdulus decoratus, Micula concave, Micula decussata,

Micula murus, Prediscosphaera cretacea, Quadrum tridum, Quadrum
gartneri, Reinhardites levis, Rhagodiscus angustus and Watznaueria
barnesae, whereas Biantholithus sparsus, B. elegans, Cruciplacolithus
tenuis, Prinsius spp., Heliolithus kleinpelli, Heliolithus cantabriae and
Prinsius dimorphosus are the dominant species of the reworked Paleocene nannooral assemblage. EarlyMid Eocene is mainly attributed
to species such as Tribrachiatus orthostylus (Early) and Coccolithus
staurion (Mid).
Calcareous nannofossil assemblages are dominated by the genera
Coccolithus, Discoaster, Ericsonia, Reticulofenestra and Sphenolithus
while Helicosphaera and Cyclicargolithus also occur. Calcareous
nannofossil assemblages mainly consist of Coccolithus eopelagicus,
Coccolithus pelagicus, Chiasmolithus titus, Cribrocentrum reticulatum,
Cyclicargolithus oridanus, Ericsonia formosa, Ericsonia subdisticha,
Helicosphaera compacta, Helicosphaera euphratis, Helicosphaera
intermedia, Pontosphaera spp., Reticulofenestra bisecta, Reticulofenestra
ovata, Reticulofenestra umbilica, Sphenolithus moriformis, Toweius

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Fig. 6. Outcrop photographs of the slope fan facies. (A) Thick-bedded, structureless sandstone. (B), (C) Channelll facies with conventional thinning and ning upward trend. Note
on (A) and (C), the sharp contact between the sandy ll and the underlying mudstone. (D), (E) Thin-bedded sandstones with parallel and/or cross lamination. Note on (E) the siltstone laminations within the mudstone facies. (F) Normally graded, amalgamated sandstones of the channelll.

gammation, Transversopontis pulcheroides and Zygrhablithus bijugatus.

Forty species, of diverse genera, were recognized (Fig. 8). Smear slides
were studied in light microscopy and microphotographs of signicant
calcareous nannofossil taxa are illustrated in Fig. 9.
The observed calcareous nannofossil assemblages are typical of the
NP19, NP20 and NP21 biozones. The boundary of the NP19 and NP20
biozones, marked by the FAD of Sphenolithus pseudoradians, is widely
considered unreliable being this species already observed in Middle
Eocene rocks (Aubry, 1992; Luciani et al., 2002), so we recognize a combined NP19/NP20 zone. The base of this biozone is marked by the FAD of
Isthmolithus recurvus (Perch-Nielsen, 1985). Supporting evidence also
comes from the occurrence of Reticulofenestra oamaruensis, which also
marks the onset of the NP19 biozone. This species is common through
Upper Eocene to Lower Oligocene and very well-documented geochronologically. Its FAD yields an age of 35.4 Ma while its LAD is above the
E/O boundary (Wei and Wise, 1992; Berggren et al., 1995). The E/O
boundary, in terms of calcareous nannofossil events, is drawn at the

top of NP20 zone and is dened by the LAD of the disk-shaped

discoasters represented by Discoaster saipanensis (Perch-Nielsen, 1985).
The base of the Early Oligocene NP22 zone is marked by the LAD of
Ericsonia formosa (Perch-Nielsen, 1985).
On Lemnos, the FAD of Reticulofenestra oamaruensis appears at the
base of the studied basin-oor fan (samples E5 and E7), indicating that
the accumulation of the submarine fan system was initiated ~35.4 Ma
ago. The LAD of Discoaster saipanensis occurs within the conglomeratic
slope fan (sample H9). These lines of evidence indicate that the
basin-oor fan, the slope fan system and the lower part of the conglomeratic fan formed during the NP19/NP20 zones (Late Eocene).
These sub-environments contain nannofossil assemblages that include
biostratigraphically signicant nannofossils (e.g. D. saipanensis, Discoaster
barbadiensis and Discoaster tanii). These taxa last occur (LO) within the
Early Oligocene, suggesting a Late Eocene age. Additional taxa common
in the NP19/NP20 biozones have been observed (e.g. Sphenolithus
predistentus, Cyclicargolithus oridanus and Zygrhablithus bijugatus).

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87

Fig. 7. Outcrop photographs of shelf facies. (A) Sandstone dominated sequence that progressively grades upwards into a mudstone dominated sequence (B) where organic-rich
beds (C) occur.

Furthermore, cosmopolitan nannofossils (e.g. C. oridanus, Sphenolithus

moriformis and Coccolithus pelagicus) are often present and reect a
common global feature for the Late Eocene nannofossils. Upslope, the
remainder deep-sea fan system and shelf environment contain the
taxa R. oamaruensis, Ericsonia formosa and Reticulofenestra umbilica. Upwards, these rocks exhibit a general decrease in the frequency of
Discoaster and Sphenolithus, followed by extinction of the disk-shaped
D. saipanensis. The only discoasters observed are Discoaster deandrei,
D. barbadiensis and D. tanii. Compared to the Upper Eocene deposits,
these sediments are characterized by lower diversity in calcareous
nannofossil assemblages (Fig. 8). The nannofossil assemblage indicates
that the upper part of the deep-sea fans and overlying shelfal deposits
were formed during the NP21 biozone (Early Oligocene), based on similar assemblages elsewhere (e.g. Marino and Flores, 2002; Watkins,
2007).
Summarizing, on Lemnos Island, during E/O boundary interval, a
sand-rich submarine fan system accumulated. During the Late Eocene
(NP19/NP20) the basin-oor system, the slope fan and the lower part
of the conglomeratic fan formed. Subsequently, the remainder of the
deep-sea fan system and upslope shelfal deposits accumulated during
the Early Oligocene (NP21) (Fig. 10).
6. Trace fossil assemblages
6.1. Systematic ichnology
The Upper Eocene/Lower Oligocene sedimentary succession on
Lemnos Island contains 19 ichnospecies, mostly graphoglyptids. These
graphoglyptids are interpreted as pre-turbidite trace fossils, formed during the deposition of background deposits between turbidity currents.
Graphoglyptid trace fossils have been documented in detail in other
studies (e.g. Uchman, 1998; Rodrguez-Tovar et al., 2011) and represent
three dimensional burrow systems preserved as bi-dimensional casts on
the soles of thin-bedded turbidite sandstones, not only in the distal basin
plain but also in other more proximal settings (Heard and Pickering,
2008). They are considered to be pre-turbidite trace fossils that develop
in the background mud as agrichnial burrow systems for microbial farming or trapping (Seilacher, 1977, 2007).
6.1.1. Simple and branched structures
Planolites beverleyensis (Billings, 1866) (Fig. 11A) occurs as
hypichnial, straight to slightly winding ridges preserved in sandstone

beds. The burrows are predominantly horizontal, long, cylindrical,

smooth walled, unlined, unbranched and oriented more or less parallel
to the bedding plane. They occur as a single isolated specimen. The
length of the burrows varies from 18 to 45 mm and the diameter from
2.5 to 5 mm. Planolites is generally regarded as a feeding structure of deposit feeders, mainly worms (Pemberton and Frey, 1982), or possibly larval insects in continental deposits (Buatois et al., 1998).
Ophiomorpha rudis (Ksikiewicz, 1977) (Fig. 11B) is a hypichnial
straight, branched trace fossil. It forms sand-lled cylinders, 520 mm
in diameter, smooth or covered locally with irregular granules or with
short oblique ridges, which are according to Rodrguez-Tovar et al.
(2010), casts of scratch marks. The granules are 2 mm long and
1.5 mm wide, and are arranged perpendicularly to the burrow axis.
Ophiomorpha rudis is a typical species of sandy deep-sea sediments
(Uchman, 2001).
Ophiomorpha isp. (Ksikiewicz, 1977) (Fig. 11C) is a hypichnial
horizontal, straight or slightly winding trace fossil, usually smooth
or covered with poorly visible knobs. Trace fossil diameter ranges
from 10 to 20 mm. The trace fossils are elliptical to rarely circular in
cross section and display rare Y-shaped branching. Ophiomorpha isp. is
typical post-depositional trace fossil of high-energy regimes (Uchman,
1999; Wetzel and Uchman, 2001).
Thalassinoides paradoxicus (Woodward, 1830) (Fig. 11D) is a
hypichnial trace fossil. Three dimensional structures forming horizontal networks, smooth-walled, irregularly branched. The branches are
Y- to T-shaped and occasionally enlarged at the bifurcations points.
Burrow diameter varies from 3 mm to 8 mm.
Thalassinoides is produced by scavenging and deposit-feeder crustaceans in marine environments of different water depths (Ekdale,
1992; Bromley, 1996). This trace fossil occurs in oxygenated, soft
but cohesive sediment (Wetzel, 2008).
Halopoa imbricata (Torell, 1870) (Fig. 11E) is a hypichnial, cylindrical, ~ 20 cm long, simple trace fossil, 8 to 9 mm wide. This trace
fossil is generally horizontal, covered with longitudinal irregular ridges
or wrinkles. Halopoa is formed by large, robust, and deeply burrowing
organisms which exploited organic rich layers within the sand beds.
They are typical of proximal axial sections, at least in part, because the
trace-making organisms were well-adapted to surviving burial by
newly deposited thick sand beds (Wetzel and Uchman, 2001).
Thalassinoides isp. (Ehrenberg, 1944). Hypichnial, slightly winding
trace fossil. Thalassinoides isp. is predominantly horizontal, more or less
regularly branched, essentially cylindrical burrow system consisting of

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Fig. 8. Detailed nannofossil assemblages of the selected samples.

a horizontal network connected to the surface by a more or less vertical

shaft. The branches are Y- to T-shaped and usually enlarged at the bifurcations points. A horizontal branching polygonal network is dominant,
with vertical shafts connected to the surface. The diameter of branches
varies from 2 to 5 mm (Fig. 13B).
6.1.2. Winding and meandering structures
Helminthorhaphe exuosa (Uchman, 1995a) (Fig. 12A, B) is a
hypichnial, string-shaped, tightly meandering trace fossil with relatively deep meanders. This trace fossil consists of a string that follows
narrow and high amplitude meanders, which lacks distinct bulges in

the curved portions. Seilacher (1977) suggested that H. exuosa is

generally characterized by widely spaced meanders. The studied material also contains specimens with relatively tightly spaced meanders. However, distance between meanders is a variable feature and
thus, it is not regarded as a diagnostic feature (Uchman, 1998). Strings
are 2 to 6 mm wide and meanders are smooth and maintain fairly uniform amplitude.
Helminthorhaphe japonica (Seilacher, 1977) (Fig. 12C) is a
hypichnial, non-branching graphoglyptid trace fossil showing only
one order of meanders. Tunnels are 2 to 3 mm wide and rather straight.
Meanders are smooth, locally pressed and maintain fairly uniform

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89

Fig. 9. Light microscope microphotographs of some selected calcareous nannofossils. 12. Zygrhablithus bijugatus, 3. Cyclicargolithus oridanus, 4. Reticulofenestra wadae,
5. Reticulofenestra oamaruensis, 6. Reticulofenestra umbilica, 7. Pemma papillata, 8. Ericsonia subdisticha, 9. Cribrocentrum reticulatum, 10. Pontosphaera multipora, 11. Reticulofenestra
bisecta, 12. Zygrhablithus aff. bijugatus ssp., 13. Reticulofenestra hillae, 14. Braarudosphaera bigelowii, 15. Coccolithus eopelagicus.

amplitude. The observed specimens show well preserved and regular

meanders but are partially destroyed. Smoothing is a typical feature in
studied specimens. Monaco (2008) proposed that smoothing resulted
from a uting process that caused partial destruction of the downcurrent side of tunnels and produced a broad, apron-shaped structure
or tail.
Desmograpton ichthyforme (Fuchs, 1895) (Fig. 12D) is preserved as
hypicnial double rows of string-like, U-shaped semi-meanders.
Desmograpton ichthyforme displays angular narrow semi-meanders
which appear as parallel ridges. On Lemnos, the studied specimen
appears to cross a minigroove induced by a pre-turbidite current.
Supporting evidence comes from the occurrence of undisturbed strings.
Desmograpton is interpreted as a three-dimensional graphoglyptic trace
fossil (Seilacher, 1977). It occurs in submarine fan deposits from the
Silurian to the Miocene (Uchman, 1995a).
Scolicia prisca (de Quatrefages, 1849) (Fig. 12 E) is preserved as
hypichnial, trilobate furrow with slightly concave, semicircular bottom and oblique slopes. This trace fossil is characterized by a densely
packed ne transverse ribs at the bottom and loose, asymmetrical and
thicker ribs on the slopes. Uchman (1998) proposed that the densely

packed ribs at the bottom are probably produced by locomotion organs of the producer while the asymmetric thicker ribs on slopes
are remnants of the edges of backll menisci.
Scolicia vertebralis (de Quatrefages, 1849) (Fig. 13A) is a
preservational variant of Scolicia prisca (Uchman, 1998). The distinguished furrow has one central narrow string or suture-structured
which passes into the furrow having indistinct trilobite structure.
Cosmorhaphe lobata (Seilacher, 1977) (Fig. 13B) is a hypichnial,
partly eroded, semicircular string, which is generally arranged in twoorder meanders. In general, C. lobata forms widely spaced, rst-order
meanders. These meanders contain turns of regular second-order undulations which are slightly higher than wide (Seilacher, 1977). The distinguished specimen exhibits a second-order meander, which is well
developed as strings of 2 to 3 mm in diameter and wavelength of
~7 mm. Compared to the wavelengths (~7 mm), the amplitudes of
the second-order meanders are distinctly larger (~9 mm).
Cosmorhaphe sinuosa (Seilacher, 1977) (Fig. 13C) is a hypichnial,
partly eroded, trace fossil that consists of a 7 to 10 mm diameter burrow.
This particular trace fossil form widely spaced, rst-order meanders,
which consist of second-order undulations of greater wavelength than

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Fig. 10. Calcareous nannofossil biostratigraphic zonation and sequence stratigraphic framework for the Upper Eocene/Lower Oligocene studied deposits.

amplitude. Seilacher (1977) suggested that C. sinuosa can be distinguished from related forms by the low amplitude of the secondary undulations and the relatively small burrow diameter. The form illustrated
here has low amplitude secondary undulations but a relatively large burrow diameter. However, size alone is not an entirely satisfactory
ichnospecic criterion.
6.1.3. Networks
Paleodictyon (Menghini, 1850) is a distinct trace fossil preserved
commonly as casts on the sole of ne- to medium-grained turbidites
(Peruzzi, 1881). It is a three-dimensional burrow system made of a
regular, repetitive net made up of horizontally distributed hexagonal
meshes and, if preserved, vertical outlets. This group, represented by
many ichnospecies, is very commonly recovered in the studied sequences as pre-depositional trace fossils at the soles of turbidites, mainly in deep-sea fan deposits. The ichnotaxonomy of Paleodictyon is based
on the maximum mesh size and the string diameter (Uchman, 1995a).
In the studied deposits several forms occur and Paleodictyon regulare,
Paleodictyon strozzi, Paleodictyon minimum, Paleodictyon hexagonum
and Protopaleodictyon have been distinguished.
Paleodictyon regulare (Ksikiewicz, 1977) (Fig. 14A) is hypichnial,
with a horizontal hexagonal net of semicircular ridges (strings). The
meshes are ~ 8 mm wide and the string ~ 2 mm in diameter.
Paleodictyon strozzi (Ksikiewicz, 1977) (Fig. 14B, C) differs in
that the meshes are 3 to 5.5 mm wide and the string is 0.3 to 0.5 mm
in diameter.

The studied specimen of Paleodictyon minimum (Sacco, 1888)

(Fig. 14C) is a very small Paleodictyon, with a mesh-size up to 1 to
2 mm and string diameter 0.2 to 0.4 mm.
Paleodictyon (Ramodictyon) hexagonum (Menghini, 1850) (Fig. 14D)
is a hypichnial, knob shaped trace fossil with vertical outlets. Bending
(Bnd) and smoothing (Smo) occurs at the shallow level where shafts
developed. This was interpreted by Monaco (2008) to be a result of
the effects of utes and suggests the existence of currents with multi directional currents. Bending and smoothing are often associated with the
same fossil specimen, but exhibit different orientations. This association
of bending and smoothing as products of unidirectional currents leads
to the exclusion of compaction as the primary factor.
Protopaleodictyon (Ksikiewicz, 1970) (Fig. 14E) is preserved as
hypichnial, wide rst-order meanders and more or less regular secondorder meanders with one or two appendages usually branching from
the apex of the second-order meanders.
6.1.4. Radial structures
Asterozoa (Bell, 2004) (Fig. 14F) occurs as hypichnial trace fossil
with attened exible body and long slender arms. The studied specimen typically has a central disk and arms, which are longer than the
diameter of the disk. Furthermore, this particular specimen suggests
some entrainment by a turbidity current, which has repositioned
the distal ends of three of its arms into the direction of current ow.
Specimens of Asterozoan are sparse because they suffer rapid destruction after death (e.g. Twitchett et al., 2005).

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6.2. Trace fossil distribution

The types of trace fossils identied, as well as trace fossil diversity
and abundance, signicantly changes throughout the studied sedimentary succession (Fig. 15). Some of the trace fossils such as Ophiomorpha
isp. and Scolicia prisca are continuously recorded and can be considered
as the permanent background components of the trace fossil assemblage. Thalassinoides isp., Ophiomorpha rudis and Scolicia vertebralis
also occur frequently and continuously throughout the deep-sea fan,
but are absent in the upslope shelf deposits. Thalassinoides paradoxicus
also occur throughout the submarine fan system, but less frequently
and less continuously. The basin-oor fan is characterized by an increased trace fossil diversity and abundance, showing the highest
values throughout the studied section. Together with the background
ichnotaxa, 19 ichnospecies have been identied (Fig. 15). In the slope
system and the conglomeratic slope fan, trace fossil diversity and abundance signicantly decrease. Apart from the background assemblage,

91

the ichnotaxa known already from basin-oor fan (T. paradoxicus,

Thalassinoides isp., O. rudis and S. vertebralis) occur. The trace fossils include Desmograpton, Helminthorhaphe, Cosmorhaphe and Paleodictyon
are not present. These ichnotaxa belong to pre depositional, predominantly winding and meandering forms. The upslope shelf deposits are
characterized by a decrease of turbidite content. The trace fossil abundance and diversity further decrease and only the background assemblage (Ophiomorpha isp. and S. prisca) occur.
7. Discussion
7.1. Calcareous nannofossil and trace fossil distributions in different
depositional environments
Turbidity currents imply reworking and transport of material from
the continental shelf and shallow upper slope towards the deeper
parts of the basin, where the allochthonous material is redeposited

Fig. 11. Simple and branched trace fossils developed in the Lemnos sand-rich deep-sea fan system. (A) Planolites beverleyensis. (B) Ophiomorpha rudis (C) Ophiomorpha isp.
(D) Thalassinoides paradoxicus. (E) Halopoa imbricata.

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Fig. 12. Winding and meandering trace fossils that were distinguished in the studied sand-rich submarine fan system. (A), (B) Helminthorhaphe exuosa. (C) Helminthorhaphe japonica.
(D) Desmograpton ichthyforme. (E) Scolicia strozzii.

together with the autochthonous sediment (Ortiz et al., 2011). Therefore,

in order to elucidate the environmental conditions that prevailed in the
deep sea during the deposition of the studied deep-sea fans and shelfal
deposits on Lemnos, we deal with the distinction between allochthonous
from autochthonous taxa. The occurrence of shallow water calcareous nannofossil taxa within the studied turbidite system, such as
Braarudosphaera bigelowii, Pontosphaera spp. and Thoracosphaera
spp., implies that turbidite accumulation inuenced the abundance
and distribution of the calcareous nannofossils. Such taxa are considered as reworked taxa that were transported down-slope by turbidity currents. Within the shelf sediments, shallow water taxa, such as
Micrantholithus spp. and B. bigelowii indicate a basinward shift of the
coastline and reect the inuence of the variations in depositional

environments in the distribution of the nannofossils. In particular,

Micrantholithus spp. is associated with shelf environments and is extremely rare in open ocean deposits (Bown, 2005).
The fundamental control on trace fossil assemblages across the
Eocene/Oligocene boundary in studies elsewhere is interpreted as
sedimentation rate and variations in depositional environments
(Uchman, 1995b; Tchoumatchenco and Uchman, 2001). Global environmental changes have been also been invoked to account for the distinct
drop in diversity and even more pronounced drop in the abundance of
graphoglyptids that occurred in the Oligocene (Uchman, 1999). On
Lemnos, the decrease in ichnofossil abundance and ichnodiversity suggests that conditions for macrobenthic fauna deteriorated since the Late
Eocene.

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During the Late Eocene, ~200 m of deep-water sediments accumulated in a basin-oor fan (~150 m) and overlying slope fan (~75 m). During
the Early Oligocene, the remainder of the slope turbidite system and upslope shelfal deposits ~125 m thick was deposited. In the slope fan,
higher sediment accumulation rates associated with proximal and axial
environments may have inuenced the nature of the trace fossil assemblages. The time between sand deposition may have been insufcient for
colonizers to migrate from areas unaffected by the newly deposited
sediment (Uchman, 1995b) resulting in the low diversity trace fossil
assemblage dominated by traces formed by organisms well adapted to
the depositional environment (e.g. Thalassinoides, Ophiomorpha and
Scolicia). In addition, the coarse-grained nature of the deposits in the
proximal and axial environments may have excluded burrowers adapted
to ne-grained sediments (Tchoumatchenco and Uchman, 2001) and
may have prevented the preservation of the activities of smaller metazoans. In contrast, basin-oor fans are associated with lower sedimentation rates relative to slope fans suggesting organisms had more time to
colonize and bioturbate the newly deposited sediments (Heard and
Pickering, 2008).
The distribution of average bioturbation intensity through different
sub-environments may be also related, at least in part, to bed thickness
and grain-size distribution (Heard and Pickering, 2008). The observations of bioturbation in this study are simply qualitative rather than
quantitative. The general reduction in bed thicknesses and grain-size
in the studied basin-oor fan compared to the slope fan (Maravelis
and Zelilidis, 2012), may have also promoted the higher bioturbation
in the basin-oor fan. Such information has been used in several studies
to make similar inferences and is interpreted as a result of enabling a
greater variety of organisms to survive burial, compared to the more
proximal slope fan environments (Wetzel, 1991; Heard and Pickering,
2008). The basin-oor fan was closer to areas unaffected by the depositing sediment gravity ows and the newly deposited sediments, due to

93

the reduced migration distances, may have been colonized by migrating

organisms more quickly. The reduced migration distances have been invoked to account for the high diversity and abundance of the trace fossils (Heard and Pickering, 2008).
The distribution of trace fossils in a submarine fan and related environments is also linked to environmental stability (Uchman, 1995a).
The ichnofauna community on basin-oor fan is mainly associated
with an abundance of pre-depositional ichnotaxa. The trace fossil assemblages are low-diversity and are characterized by high abundances of individual trace fossils. In contrast, on slope fan and shelfal environment,
the trace fossils are rare. Compared to the slope fans, the basin oor
fans have increased trace fossil abundance as a result of predicted supply
of organic matter to the deep-sea oor in a short span of time (Uchman,
2003). The fairly constant organic material supply on the studied basin
oor fan might have been an evolutionary factor promoting farming activity. Summarizing, on Lemnos, the decrease in ichnofossil abundance
and ichnodiversity suggests that conditions for macrobenthic fauna deteriorated since the Late Eocene. This observation is interpreted to be a result of regional changing in depositional environments across the E/O
boundary rather than the global climatic changes.

7.2. Trace fossil distribution in relation to base-level dynamics

Some of the mentioned paleoecological features, such as oxygenation level and organic matter are conditioned by the base-level dynamics (Rodrguez-Tovar et al., 2010). Thus, the vertical distribution
of trace fossils can be related to the sequence stratigraphic framework
proposed for the studied sedimentary sequence. According to this
framework, the studied sand-rich submarine fan system and shelfal
environment accumulated during the base-level fall and subsequent
base-level rise (Maravelis and Zelilidis, 2011).

Fig. 13. Winding and meandering trace fossils that were distinguished in the studied sand-rich submarine fan system. (A) Scolicia vertebralis. (B) Cosmorhaphe lobata (CL) and
Thalassinoides isp. (TI). (C) Cosmorhaphe sinuosa.

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Fig. 14. Network trace fossils. (A) Paleodictyon regulare. (B) Paleodictyon strozzi. (C) Paleodictyon minimum. (D) Paleodictyon (Ramodictyon) hexagonum. (E) Protopaleodictyon.
(F) Asterozoan fossil external mold, showing possible current entrainment probably within, or at the base of, a turbidity-current as displayed by the curved specimen arms
(white arrows).

The basin-oor fan corresponds to the development of the lower

middle part of the low-stand system tract (Maravelis and Zelilidis,
2011). The ichnofaunal research suggests moderately oligotrophic and
well oxygenated conditions. Explanation of this situation by base-level
dynamics is not obvious, and several scenarios have been invoked to account for the high trace fossil abundance and diversity. It has been proposed that less mud is resuspended during development of low-stand
systems tracts, and in consequence more oligotrophic conditions appear
(Rodrguez-Tovar et al., 2010). Trace makers may adapt to the lateral inux of organic matter and to increasing siliciclastic interturbiditic mud. It
is also possible that prolonging lithological diversication of sea oor due
to deposition of turbidites promoted new behavioral adaptations, and
consequently increased trace fossil diversity (Rodrguez-Tovar et al.,
2010).
The slope system and part of the conglomeratic slope fan correspond to the upper part of the low-stand system tract and consist
of turbidite-rich sediments deposited during the relative base-level
fall (Maravelis and Zelilidis, 2011) and contain less trace fossils.

The remainder of the conglomeratic slope fan and the shelf deposits
are characterized by a further decrease in trace fossil abundance and diversity. These depositional sub-environments are turbidite-poor and
correspond to the uppermost part of the low-stand system tract and
the rising sea-level during the subsequent transgressive high-stand systems tracts (Maravelis and Zelilidis, 2011). Progressive eutrophization of
the environment has been proposed by Rodrguez-Tovar et al. (2010) to
explain the decrease in ichnofossil abundance and ichnodiversity. On the
Upper Oligocene conglomeratic slope fan and upslope shelfal environment, such an explanation is possible and compatible with the
nutrient-enriched cool surface waters interpreted on the basis of
the absence of warm and well-oxygenated calcareous nannofossils.
7.3. Paleoecology
Coccolithophorids distribution is largely controlled by temperature,
water mass conditions and trophic resources (Aubry, 1992; Persico and
Villa, 2004). The distribution pattern of the calcareous nannofossil

A. Maravelis, A. Zelilidis / Palaeogeography, Palaeoclimatology, Palaeoecology 365366 (2012) 8198

assemblages recorded on Lemnos reects the environmental changes,

both climatic and oceanographic, that characterized the E/O boundary.
The Upper Eocene basin-oor fan and the lower part of the slope fan
are dominated by the Sphenolithus and Discoaster genera. These taxa
are mostly conned to warm and well-oxygenated surface waters
(Aubry, 1992). In particular, the species Discoaster is mostly ecologically
controlled, and indicative of oligotrophic conditions (Aubry, 1992). In
the Lower Oligocene conglomeratic slope fan and shelf deposits, the occurrence of these taxa decreases. This observation that is followed by the
extinction of most discoasterids (only Discoaster deandrei and Discoaster
tanii are present), which indicates progressive eutrophization and

95

decrease in water temperature and salinity (Rodrguez-Tovar et al.,

2010). The genus Sphenolithus is represented by the long ranging and
diagenetically-resistant species Sphenolithus moriformis (Sphenolithus
celsus and Sphenolithus radians are rare).
The climate changes across the E/O boundary are further
established by the occurrence of the taxa Braarudosphaera bigelowii
and Micrantholithus spp. in shelfal deposits. These species, which
are typical of low oxygen and salinity waters (Peleo-Alampay et al.,
1999), indicate inux of terrigenous material (vabenicka, 1999) and
eutrophication (Cunha and Shimabukuro, 1997). Even though the paleoecological preferences of B. bigelowii have been studied in detail,

Fig. 15. Synthetic log of the Lemnos section with the vertical distribution of trace fossils.

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A. Maravelis, A. Zelilidis / Palaeogeography, Palaeoclimatology, Palaeoecology 365366 (2012) 8198

not much is known about the paleoecology of the genus Micrantholithus

(Bartol et al., 2008). Its ecology is probably related to neritic factors such
as lowered salinity and eutrophic waters (Street and Bown, 2000). Even
though the occurrence of Micrantholithus indicates warm water, the
broad latitudinal range of the genus seems to reect its tolerance to relatively large amplitude of temperatures (Bartol et al., 2008).
During the Early Oligocene, paleoclimatic cooling lead to
paleoenvironmental and biotic changes in the open seas (Aubry,
1992; Zachos et al., 2001). Northern Hemisphere glaciation was initiated during the Middle Eocene to Early Oligocene, while cold polar
water entered several deep-sea basins (e.g. Carpathians and Alps),
and local anoxic conditions occurred, inuencing trace fossil assemblages (Leszczyski, 1997). Graphoglyptid trace making organisms, associated with the infaunal farming activity exploiting short seasonal
rains of phytoplankton, were successfully adapted to oligotrophic conditions (Uchman, 1999). In the described habitat, a progressive deterioration in macrobenthos suitability occurs. This is interpreted to be
fundamentally related to variations in depositional environments. Nevertheless, the changes of climatic conditions may have partly inuenced
the decrease of graphoglyptids and other trace fossils. In particular, the
E/O crisis inuenced the Lower Oligocene deep-sea fans and shelf deposits promoting the deterioration in macrobenthos suitability. The
ichnofossil distribution suggests a decrease in temperature, salinity, oxygen levels and nutrient supply.
During periods of high primary production, so much organic material is provided and thus, the oxygen ux into the sediment is lowered
(Wetzel, 2002). Increases in primary production are probably driven
by cooling and increase the surfaceocean divergence (Salamy and
Zachos, 1999). The associated upwelling of nutrient-rich deep waters
and/or enhanced uxes of weathering derived nutrients during rapid
sea level fall lead to the concentration of benthic food in shallow sediment surface layers (Salamy and Zachos, 1999). The lower nutriment
supply and oxygenation level could induce the decrease of deeper
trace makers, while those living in shallower substrates, close to the
sea oor, although with benthic food available, could not survive because of no sufcient oxygen supply.
8. Conclusions
The detailed biostratigraphic and ichnofaunal research on the
sand-rich submarine fan system and upslope shelfal deposits on the
southern part of the Greek Thrace Basin (Lemnos Island) indicates:
1) The age of the succession is Late Eocene/Early Oligocene (NP19NP21
biozones). The E/O boundary is drawn at the top of NP20 biozone and
is dened within the conglomeratic slope fan. Ages are established on
the youngest index species of calcareous nannofossils found in the
mudstone intervals.
2) The calcareous nannofossil assemblages indicate a gradual decrease
in temperature, salinity, oxygenation level and nutrient supply across
the E/O boundary. Compared to the Lower Oligocene, the Late Eocene
nannooras contain genera diagnostic of warm and well-oxygenated
surface waters.
3) A distinct drop in diversity and abundance of graphoglyptids and
other trace fossils towards the Lower Oligocene is noted. This observation reects the impact of the changing in the depositional
environments from basin-oor fan to slope turbidite system and
nally shelfal environment. Among the interpreted limiting factors affecting the macrobenthic habitat are the bed thickness,
grain-size and sedimentation rates. Compared to the basin-oor
fan, the slope system has thicker and coarser sandstone beds, limiting the abundance of trace fossils.
4) Additional factors that incidentally affected the macrobenthic habitat
are changes of trophic level, temperature and ecological disturbances,
which in turn are partly determined by the base-level dynamics. Increased diversity of trace fossils in the basin-oor fan is interpreted

to be a result of moderate oligotrophy and stabilization of ecological

conditions. Lowered oxygenation and associated decrease of temperature, typical during low sea-level, may limit the diversity of trace
fossils in slope turbidite system and upslope shelfal environment.
The biostratigraphic and ichnofaunal features presented in this research reveal the usefulness of the fully integrated sedimentological,
biostratigraphic and ichnological approach to interpret changes in
the atmosphereocean system during the past, including climatic
dynamics.
Acknowledgments
The work presented in this article presents the research by the authors as part of the Department of Geology, University of Patras, Greece
internal and externally funded studies. Funding was supported by EC
(European Commission) and GSRT (Greek Ministry of Development).
The authors thank journal Editor (F. Surlyk), F.J. Rodrguez-Tovar
(Univ. Granada) and the anonymous reviewer for their valuable suggestions and comments on the manuscript. The authors acknowledge
K. Stoykova (Univ. Soa) for the determination of calcareous
nannofossils. The authors gratefully acknowledge D.J.W. Piper (Univ.
Dalhousie) and Georgia Pe-Piper (Univ. Saint Mary) for their thorough
editing and suggested changes to English.
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